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. . the best single guidebook to American mushrooms."
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 ushrooms
Demystified
 S.s.F. Public Library
West Orange
840 West Orange Ave
Coin FrnnciSCO, CA 94Q8C

David Arora

Mushrooms
Demystified
A Comprehensive Guide
to the
Fleshy Fungi

SECOND EDITION

Ten Speed Press

Berkeley
 Copyright © 1979, 1986 by David Arora
All rights reserved. No part of this book may be reproduced in any form, or by any means elec¬
tronic or mechanical, including photocopying, recording or by any information storage and
retrieval system, without written permission from the Publisher.

1€>
TEN SPEED PRESS
Post Office Box 7123
Berkeley, California 94707
www.tenspeed.com

Distributed in Australia by Simon and Schuster Australia, in Canada by Ten Speed Press Canada, in
New Zealand by Southern Publishers Group, in South Africa by Real Books, and in the United
Kingdom and Europe by Airlift Book Company.

Cover design by Brenton Beck,

Fifth Street Design Associates.
Illustrations by Michael Cabaniss.
All photographs by David Arora
unless otherwise indicated.
Front cover photograph or Agaricus augustus by Bill Donaldson.

Grateful acknowledgement is made to Alan Snitow of the San Francisco Bay Guardian and to the
Peoples Press, in whose pages some of the introductory material originally appeared.

Library of Congress Catalog Number 79-85123

The author welcomes information and feedback from readers.

You can write to him care of the publisher.

Library of Congress Cataloging-in-Publication Data

Arora, David
Mushrooms demystified.

Bibliography.
Includes index.
l.Mushrooms-California-Identification.
2. Mushrooms-Pacific States-Identification.
I. Title
QK617 .A69 1986 589.2’223’09794 86-5917
ISBN-13: 978-0-89815-169-5
ISBN-10: 0-89815-169-4

11 12 13 14 15 - 10 09 08 07 06

Printed and bound in Hong Kong.

The information in this book is accurate to the best of the author’s knowledge.
However, neither the author nor the publisher accepts responsibility for mistakes in
identification or idiosyncratic reactions to mushrooms. It is certainly not necessary to
eat mushrooms in order to enjoy them. People who choose to eat mushrooms do so
at their own risk.
 I dedicate this book
with love
to my mother and father,
whose admonitions to me as a teen-ager
to stay away from mushrooms
inspired me to get closer
 PREFACE TO THE SECOND EDITION
THE first edition of Mushrooms Demystified was designed primarily for
California. The second edition is useful throughout the United States and
Canada. The distinction is not as great as it sounds because most of the mush¬
rooms found in California are widely distributed; the text has undergone con¬
siderable change nevertheless. Prominent and distinctive species are included
from all parts of the continent: New England, the South, Midwest, Southwest,
Rocky Mountains, Pacific Northwest, and of course, California. In addition,
the nomenclature has been updated, over 200 new color plates have been added,
most of the original black-and-white photographs have been replaced, and
certain groups such as the truffles are treated in greater detail. One thing that
has not changed is the tone of the tome—I have once again made a special effort
to keep the terminology simple and the language entertaining. The result is a
bigger book, and I think, a better one.
Of course, in trying to become more things to more people, one runs the risk
of becoming less to some. Despite a California slant (intended or not, every
field guide has built-in geographical bias because every author has more field
experience in some regions than others), the inclusion of species from across the
country may disappoint Californians who found the limited scope of the first
edition reassuring. The text and keys are longer because of the larger number of
species covered, but they are not any more difficult to use. Simple geography,
in fact, often facilitates recognition (e.g., Suillus lakei of western North America
and S. pictus, its eastern counterpart).
I have, however, maintained a certain provincial flavor by providing detailed
season and habitat information for the Central California Coast (hereafter
referred to as “our area”). Southern Californians, northern Californians, eastern
Californians, non-Californians, and worldly central Californians may find this
disconcerting, but there are good reasons for doing so. The terrain, climate, and
vegetation of North America are remarkably varied. Anyone who has used or
written books of national scope knows that in striving to cover such a diverse
area, one is often forced to present information in an extremely generalized,
vague way. Just how useful is it to say of a particular mushroom, “found in woods
across the country in spring, summer, fall, or winter,” when in any one area it is
likely to have a favorite host tree and limited fruiting period? Instead of pro¬
viding information on habitat and season that is marginally useful to all readers,
I have decided to supply information that is very useful to readers fortunate
enough to live in or visit “our area,” and marginally useful (i.e., as useful as that
in any other national field guide) to the rest. Besides, any mushroom hunter
worth her salt-and-butter quickly learns when and where mushrooms grow in
her area.
I have tried to make Mushrooms Demystified as accessible as it is compre¬
hensive. For the uninitiated, there are special chapters on mushroom termi¬
nology and classification, when and where to find mushrooms, how to collect
and identify mushrooms, how to use the keys (which are the backbone of the
book), mushroom cookery and mushroom toxins, and what the scientific (Latin)
names of mushrooms mean. The book is comprehensive enough to interest
intermediate and advanced mushroom hunters. However, beginners intimidated
by its length can selectively ignore the more technical discussions and concentrate
on learning the “Seventy Distinctive Mushrooms” listed on pp. 48-51.
In using this book it is important to realize that the identification of mush¬
rooms differs in several key respects from the identification of, say, birds or
wildflowers. For one thing, there is no book that “has them all.” Over 2000
species are described, illustrated, or mentioned in this book—more than in any
other North American field guide—but you will quickly discover that many of
the mushrooms you find are not included in this book and many of the included
mushrooms do not occur in your area (unless it is “our area”). This will be truer
of some regions (e.g., the Southeast) than others, and there is no way it can be
avoided—there are just too many mushrooms! As consolation I offer the words
of Gary Lincoff (author of another national field guide): “It is better for a book
to contain good descriptions of mushrooms that don’t occur in your area than
bad descriptions of ones that do!” As every author has a different set of ex¬
periences to share, I recommend that all mushroom hunters, even those in
California, supplement this book with others, particularly regional guides
(if they exist) because their limited scope enables them to present more specific
information.
It will be many years before we have a complete inventory of North American
mushrooms. More than anything this is a tribute to the elusive nature of mush¬
rooms. They are difficult to study because they are ephemeral and unpredictable,
and so much depends on being in the right place at the right time. It is also a
comment on the fact that few people take mushrooms seriously. As a result, the
documented distribution of the lesser-known mushrooms corresponds to the
undocumented distribution of the better-known “mushroomologists.” Further¬
more, many mushroom species still await classification. While these factors
make identification more difficult, they also add an element of suspense to the
hunt. One is continually finding species that are new to science, or new to North
America, or new to one’s area, or at least new to oneself. Mycology (the study of
fungi) is a field to which discerning amateurs can make significant contributions!
Another important point to realize is that many mushrooms cannot be
positively identified unless you have access to a microscope and technical
literature and know how to use them. Microscopic characteristics are not
stressed in this book. This means that many of the species mentioned briefly are
merely suggestions as to what an unidentified mushroom might be, and many of
the species that are fully described are actually “complexes”—groups of closely
related species whose exact identities are a matter for the specialist.
In other words, I’ve chosen to sacrifice a certain degree of exactitude by inter¬
preting many species broadly. For instance, the description of Agaricus silvicola
embraces a confusing group of white woodland mushrooms with a sweet odor
and yellow-staining skin. Whether or not the “true” A. silvicola occurs in North
America, I see no harm in applying the name to the “complzx”providing readers
are made aware of the situation (for instance, by appending the word “group” to
the name) and any attendant risks. Purists may object to this approach, but there
are many amateurs who would like to apply names to these “complexes” even if
they are unequipped to make the subtler distinctions between species or varieties
within a complex. Those who wish to verify identifications, look up species’
authorities, or otherwise pursue the matter further can do so by making use of the
literature listed in the bibliography. Students and professionals interested in
the more unusual species I have collected can write to me care of the publisher—
I will be happy to furnish the names of herbariums to which specimens have been
sent.
I would also like to say a few words about the title of this book, since there
have been varied reactions to it. One person has stated publicly that mushrooms
cannot be demystified because there is nothing mystifying or mysterious about
them; others have questioned the desirability of demystification and suggested
that I call the second edition “Mushrooms demystified.” I, for one, was first
attracted to mushrooms because I found them mysterious. I still find them
mysterious and I see no danger that this book or any other will deprive them of
their mystery. Mushrooms, like other forms of life, are miracles—miracles
which we can explain but not fully comprehend. Anyone who has studied
anything knows that answers beget questions as surely as questions lead to
answers, that the more one knows about a subject, the larger and more chal¬
lenging it becomes. In addition to being an informative home and field com¬
panion, I hope Mushrooms Demystified will be an inspiration—to look more
closely, ask more questions, seek more answers. Identification, after all, is not
an end in itself, but a means toward acquiring a deeper knowledge and keener
appreciation of our co-inhabitants on this planet.
The selection of color plates reflects my hope that this book will be more than
a tool for identification. Many plates were chosen for their usefulness, but
some were picked for their beauty, dramatic effect, or charm. (A book does little
good if it does not invite one to take it off the shelf!) Mushrooms Demystified is
also a vehicle for expressing my love for (and exasperation with) mushrooms
and for people who love mushrooms, my respect for life and for people who
respect life. Much of the pleasure in getting to know wild mushrooms is directly
attributable to the “wild” companionship of fellow fungophiles, and I have paid
tribute to these folks wherever possible.

Acknowledgments
Mushrooms Demystified is a compilation of information from many diverse
sources. I have acted as assembler and interpreter, supplementing the infor¬
mation with my own knowledge, experience, keys, comments, and photographs,
and binding it together in an accessible, useful, and cohesive (assuming the
pages don’t fall out) form. I get credit for writing the book; the many mycologists
whose monographs and articles form its factual foundation do not. Yet com¬
pared to the vast fund of knowledge painstakingly accumulated by these
mycologists, my own two cents’ worth (15 years, actually) is insignificant. I have
provided a bibliography of primary references, but in the tradition of other field
guides, I have not made extensive use of footnotes or included species’ author¬
ities (the names of those who described them). I would like to express my
appreciation, then, to all the authors listed in the bibliography plus any I have
inadvertently omitted. I want them to know that they have my congratulations
and respect, and I want my readers to know that this book would not have been
possible without their dedicated efforts. If at times I seem irreverent in my
discussions of mushrooms and mycologists, it is only because I am so by nature,
and a sense of humor helps me to cope with the more exasperating aspects of
mushroom scrutiny and book-writing.
 I want to extend special thanks to those mycologists who directly contributed
to this book by giving me their time and expertise, providing identifications of
“problem” mushrooms, and offering suggestions on the manuscript. These
mycologists are (in alphabetical order, with the areas in which they have helped):
Prof. Joseph Ammirati of the University of Washington, Cortinarius\ Chuck
Barrows of Santa Fe, New Mexico, mushrooms of the Southwest; Prof. Howard
Bigelow of the University of Massachusetts, Clitocybe and Marasmius; William
Burk of the University of North Carolina, stinkhorns; Dennis Desjardin of
San Francisco State University, Marasmius and Collybia; Prof. Robert Fogel
of the University of Michigan, Hymenogaster and Destuntzia; Prof. Robert
Gilbertson of the U niversity of Arizona, resupinate (crust) fungi and mushrooms
of southern Arizona; Bill Isaacs of Santa Fe, New Mexico, Agaricus and En-
doptychum\ Prof. David Jenkins of the University of Alabama-Birmingham,
Amanita; Rick Kerrigan of the University of California-Santa Barbara,
Agaricus; Prof. David Largent of Humboldt State University, Leptonia,
Nolanea, Entoloma, and Hygrophorus; Herb Saylor of San Francisco State
University, truffles, false truffles, and coral fungi; Prof. Robert Shaffer of the
University of Michigan, Russula,; Prof. Emeritus Alexander Smith of the
University of Michigan, Cortinarius and many other mushrooms; Prof. James
Trappe of the U.S.D.A. Forestry Sciences Laboratory, Corvallis, Oregon,
truffles (especially Tuber) and false truffles; and Greg Wright of Claremont,
California, southern Californian and southwestern mushrooms. Although these
people have been of great assistance in the preparation of this book, any errors
of fact are my responsibility, not theirs.
I am also very grateful to Prof. Isabelle Tavares of the University of California-
Berkeley, for helping me to locate, examine, and photograph various Gastero-
mycetes in the herbarium there; Barbara Waaland and the biology department of
the University of California-Santa Cruz, for providing laboratory facilities;
John Anderson, Charles Prentiss, and John Lane of the Santa Cruz City
Museum, for their longtime support of my studies; Michael Cabaniss of Santa
Cruz, for his fine illustrations; and the many people who helped make this book
as comprehensive as it is by graciously providing the photographs which I lacked:
Chuck Barrows, Alan Bessette, Nancy Burnett, Bill Everson, Michael Fogden,
Ray Gipson, Dan Harper, Richard Homola, Nancy Jarvis, Rick Kerrigan, Joel
Leivick, Keith Muscutt, Herb Saylor, Phil Sharp, Bob Short, Bob Tally, Bob
Winter, Greg Wright, and Joan Zeller.
Many people have played a vital role in the production of the book. I want to
thank Phil Wood and George Young of Ten Speed Press for underwriting the
entire project and having faith in it; Neil Cossman of ASAP Typography for
three years of extraordinary patience and cooperation while the text was being
tyepset and corrected, typeset and corercted, typeset and corrected; Hal Hershey
of Hal Hershey Book Design & Production and Brenton Beck of Fifth Street
Design, for their advice and expertise on the design and layout; and Ralph and
Mildred Buchsbaum of Boxwood Press, whose initial interest in the first edition
made the second edition possible.
Also deserving recognition are the many friends and strangers wno nave neipea
me along in my research. Some have provided me with food, drink, housing, and
companionship during my travels; others have supplied valuable information
or opinions on mushrooms or clues to their whereabouts. Some have even risked
their health or that of their loved ones in order to determine the edibility of
untested mushrooms. Many have braved torrential downpours, suffered severe
cases of poison oak, fought off swarms of mosquitoes and ticks, and trudged
miles out of their way to bring me rare and interesting (as well as uncommonly
delicious) mushrooms. To Ed Aguilar, Mine and Marc Doolittle, Joe and Nancy
Haydock, Mark Hildebrand, Ginny and Rosalie Hunt, Luen Miller, Craig
Mitchell, Ciro and Rose Marie Milazzo, Suzanne and Beanie Rainbow, Bob
Sellers, Sue Willis, Bob Winter, Reggie, Willie, Duke, Max, Butch, Hank, and
all the rest—thank you from the bottom of my basket!
I would also like to thank my parents, Harbans and Shirley Arora, for their
encouragement in all phases of this lengthy project; my legion of ex-housemates
for putting up with my habit of lapsing into Latin monologues at the dinner table
and tolerating a chronic case of “There’s too many fungi in the refrigerator!”;
all those who contributed to the first edition (without which the second would not
be possible); the many readers of the first edition whose letters have brought me
such joy and satisfaction (readers of the second edition, take note!); and any and
all contributors I have forgotten to mention (my propensity for forgetting the
names of important people is equaled only by my propensity for remembering
the names of obscure mushrooms).
Finally, I would like to express the deepest gratitude to my wife, Judith Scott
Mattoon, who, I am pleased to say, did not type a single word of the manuscript
nor provide countless hours of selfless assistance without which this book would
not have been possible. This book, in fact, would have been possible without her,
but our marriage would not. She has been a lovely and lively companion,
steadfast friend, and ruthless editor on those occasions when my literary
capacities deserted me. True, she’d rather forage for Agaricus bisporus in a
grocery store than Craterellus cornucopioides in the forest, but I haven’t given
up on her yet!
David Arora
Santa Cruz
1985
 CONTENTS
Fungophobia 1
What Is a Mushroom? 4
Mushrooms and the Environment 6
Names and Classification 8
Collecting Mushrooms 11
Identification and Terminology 14
How to Use the Keys 21
Questions About Mushrooms 23
Which Mushrooms Are Good to Eat? 23
What’s the Difference Between a Mushroom and a Toadstool? 25
When and Where Do Mushrooms Grow? 25
Can People Harm Mushrooms By Picking Them? 26
Can People Harm Themselves By Picking Mushrooms? 27
Do Other Animals Eat Mushrooms? 28
What Is the Nutritional Value of Mushrooms? 30
What Is the Medicinal Value of Mushrooms? 30
Can You Grow Wild Mushrooms? 30
Hey, Maaaaaaaaan, Do Any Psilocybin Mushrooms
Grow Around Here? 31
LBM’s: Little Brown Mushrooms 32
Habitats 34
Seventy Distinctive Mushrooms (A quick-reference check list) 48
Key to the Major Groups of Fleshy Fungi 52
Basidiomycotina (Basidiomycetes) 57
Hymenomycetes 57
Agaricales (Agarics or Gilled Mushrooms) 58
Russulaceae 63
Hygrophoraceae 103
Tricholomataceae 129
Entolomataceae 238
Pluteaceae 253
Amanitaceae 262
Lepiotaceae 293
Agaricaceae 310
Coprinaceae 341
Strophariaceae 367
Cortinariaceae 396
Bolbitiaceae 466
Paxillaceae 476
Gomphidiaceae 481
Boletaceae (Boletes) 488
 Aphyllophorales 548
Polyporaceae & Allies (Polypores and Bracket Fungi) 549
Stereaceae & Allies (Crust and Parchment Fungi) 604
Hydnaceae (Teeth Fungi) 611
Clavariaceae (Coral and Club Fungi) 630
Cantharellaceae (Chanterelles) 658
Tremellales & Allies (Jelly Fungi) 669
Gasteromycetes 676
Lycoperdales& Allies (Puffballs and Earthstars) 677
Tulostomatales (Stalked Puffballs) 715
Podaxales & Allies (Gastroid Agarics) 724
Hymenogastrales & Allies (False Truffles) 739
Phallales (Stinkhorns) 764
Nidulariales (Bird’s Nest Fungi) 778
Ascomycotina (Ascomycetes) 782
Discomycetes 783
Pezizales 783
Morchellaceae (Morels and Allies) 784
Helvellaceae (False Morels and Elfin Saddles) 796
Pezizaceae & Allies (Cup Fungi) 817
Tuberales (Truffles) 841
Helotiales (Earth Tongues) 865
Pyrenomycetes (Flask Fungi) 878
Mushroom Cookery 888
Mushroom Toxins 892
What It All Means (A Short Dictionary of Scientific Names) 899
Glossary 913
Bibliography: Suggested Readings and Primary References 919
General Index 926
Genus and Species Index 936
 FUNGOPHOBIA
BRING home what looks like a wild onion for dinner, and no one gives it a second
thought—despite the fact it might be a death camas you have, especially if you
didn’t bother to smell it. But bring home a wild mushroom for dinner, and watch
the faces of your friends crawl with various combinations of fear, anxiety,
loathing, and distrust! Appetites are suddenly and mysteriously misplaced, vague
announcements are hurriedly mumbled as to dinner engagements elsewhere, until
you’re finally left alone to “enjoy” your meal in total silence.
For there are few things that strike as much fear in your average American as
the mere mention of wild mushrooms or “toadstools.” Like snakes, slugs, worms,
and spiders, they’re regarded as unearthly and unworthy, despicable and inex¬
plicable—the vermin of the vegetable world. And yet, consider this: out of several
thousand different kinds of wild mushrooms in North America, only five or six
are deadly poisonous! And once you know what to look for, it’s about as difficult
to tell a deadly Amanita from a savory chanterelle as it is a lima bean from an
artichoke.
This irrational fear of fungi is by no means a universal trait. The media and
medical profession have done their part to perpetuate it, but they are certainly not
responsible for its origin. To a large extent, we inherited our fungophobia from
the British. William Delisle Hays, an astute Englishman writing in the 1800’s,
expressed it this way:
(All mushrooms) . . . are lumped together in one sweeping condemnation.
They are looked upon as vegetable vermin only made to be destroyed. No
English eye can see their beauties, their office is unknown, their varieties not
regarded. They are hardly allowed a place among nature’s lawful children,
but are considered something abnormal, worthless, and inexplicable. By
precept and example children are taught from earliest infancy to despise,
loathe, and avoid all kinds of “toadstools.” The individual who desires to
engage in the study of them must boldly face a good deal of scorn. He is
laughed at for his strange taste among the better classes, and is actually
regarded as a sort of idiot among the lower orders. No fad or hobby is
esteemed so contemptible as that of “fungus-hunter,” or “toadstool-eater.”
This popular sentiment, which we may coin the word “fungophobia” to
express, is very curious. If it were human—that is, universal—one would be
inclined to set it down as an instinct, and to revere it accordingly. But it is not
human—it is merely British. It is so deep and intense a prejudice that it
amounts to a national superstition . . .
It is a striking instance of the confused popular notions of fungi in England
that hardly any species have or ever had colloquial English names. They are
all “toadstools,” and therefore are thought unworthy of baptism. Can any¬
thing more fully demonstrate the existence of that deep-rooted prejudice
called here “fungophobia”? . . .
A century later, in America, Alan Snitow echoes similar sentiments in the
San Francisco Bay Guardian:
I wander through Bay Area woods and fields looking for mushrooms. When
I first started, I thought my new hobby offered final proof positive that I am
—well, a weird person. I didn’t know anyone else “into” mushrooms. My
2 FUNGOPHOBIA

friends’ puzzled looks seemed to confirm my view that I must suffer from a
form of repressed necrophilia, a perverse fascination with things rotted
and decayed . . .
(That some mushrooms can be eaten helps) . . . to justify the quest, espe¬
cially to friends who raise their eyebrows at the mention of fungi. If I am off
looking for mushrooms, they think I have a rational purpose to my actions.
Otherwise they would say: “He’s off walking around in the rain again.”
Rather than pitying looks at the soggy human who returns, they see a slightly
eccentric, but productive, worker, toiling to bring back something for the
evening’s table.

Mushrooms can be every bit as beautiful as birds, butterflies, shells, and flowers,
yet we never think to describe them in such flattering terms. When novelists or
poets want to conjure up an emotion of fear, loathing, total revulsion, and
imminent decay, they inevitably drag in the mushrooms and toadstools—
malignant instruments of death and disease that appear only in the dankest and
most abominable of situations. Witness Shelley:

And agarics and fungi, with mildew and mould

Started like mist from the wet ground cold
Pale, fleshy, as if the decaying dead
With a spirit of growth had been animated . . .

And Sir Arthur Conan Doyle, bygone creator of Sherlock Holmes:

... A sickly autumn shone upon the land. Wet and rotten leaves reeked and
festered under the foul haze. The fields were spotted with monstrous fungi of
a size and colour never matched before—scarlet and mauve and liver and
black—it was as though the sick earth had burst into foul pustules. Mildew
and lichen mottled the walls and with that filthy crop, death sprang also from
the watersoaked earth.

And D.H. Lawrence, that chronic belittler of the British bourgeoisie:

How beastly the bourgeois is

especially the male of the species—
Nicely groomed, like a mushroom
standing there so sleek and erect and eyeable—
and like a fungus, living on the remains of bygone life,
sucking his life out of the dead leaves of greater life than his own.
And even so, he’s stale, he’s been there too long,
Touch him, and you’ll find he’s all gone inside
just like an old mushroom, all wormy inside, and hollow
under a smooth skin and an upright appearance.
Full of seething, wormy, hollow feelings
rather nasty—
How beastly the bourgeois is!
Standing in their thousands, these appearances, in damp England
what a pity they can’t all be kicked over
like sickening toadstools, and left to melt back, swiftly
into the soil of England.
FUNGOPHOBIA 3

And that prim American poet Emily Dickinson:

Had nature any outcast face

Could she a son condemn
Had nature an Iscariot
That mushroom—it is him.

Has any group of organisms been so unjustly maligned? Actually, Dickenson’s

limp effort should come as no surprise, since she was a virtual recluse. Prejudice
is largely a measure of ignorance!
And yet, if you go to continental Europe, you’ll find that fungophobia is the
exception and not the rule. Most Europeans, especially those who live close to
the woods, know which mushrooms to pick and how to cook them. They bestow
upon each species an individual name and sell them in the markets. Many
Americans, on the other hand, are completely oblivious to the fact that there is
more than one fungus among us—those of recent European or Oriental ancestry
being notable exceptions.
The farther east you go in Europe, the more passionate is the love for mush¬
rooms. Which brings us to Russia. The Russians go absolutely bananas over
fungus. Mushrooming is a commonplace tradition there, not the hallowed turf of
the academic or connoisseur. Instead of talking about the weather, strangers
often engage in polite conversation about how the mushroom season is
progressing. And Russian children are raised on mushroom lore from earliest
infancy. Many family names are derived from fungi: Bribov, Borovikov,
Gruzdjov, Ryshikov, Opjonkin. Another one is Griboyedev, or “Mr. Mushroom-
eater.” The poet Majokovsky was a mushroom addict. (Poetry, like mushroom
hunting, is a great tradition there. A Russian poet draws 5,000 to a poetry reading
—here you’re lucky to draw 50.) Even Lenin is said to have been possessed by a
razh or “mushroom passion.”
In this country, it is only with the renewed interest in natural foods and the
desire to return to the earth (and what’s good for you) that mushrooms are being
noticed again. Mycological societies are sprouting up in the major cities. And
of course, business is capitalizing on the trend. Polka-dotted mushrooms have
appeared in startling profusion on curtains and calendars, pottery and stationery,
potholders and incense holders, bumper stickers and birthday cakes.
And yet, when it comes down to actually eating wild mushrooms, most
Americans are still afraid. Instead they opt for something more familiar and not
half as good, such as Grape Nuts or Malt Balls. Yet it stands to reason that if
mushroom-eating were an inherently dangerous activity, it could not exist to the
extent it does in Europe. And the mycological societies in America would be in
dire need of new members, their ranks depleted annually by the insidious “Mush¬
room Menace.” Like driving, swimming, walking, or breathing, mushroom¬
eating is only made dangerous by those who approach it frivolously.
If you treat mushrooms with discrimination and respect, you can learn to pick
your own edible wild mushrooms without fear of confusing them with poisonous
types—mushrooms which are nutritious, far more flavorful than the mass-
produced cultivated variety, and best of all, free! It does, however, require time
and effort—a willingness to plunge into the woods, to uncover their secrets, to
learn their characteristics, to penetrate their haunts. That’s what this book
is about.
4

WHAT IS A MUSHROOM?
FUNGI are neither plants nor animals. They don’t contain chlorophyll like
green plants, and as a result cannot manufacture their own food. In this respect
they resemble animals, because they feed themselves by digesting other organic
matter. However, they lack the nervous system, specialized organs, and mobility
characteristic of most animals. Furthermore, fungi reproduce by means of
microscopic reproductive units called spores. These are far simpler in structure
than seeds or eggs, and in fact, usually consist of only one cell.
The term mushroom is most often used to describe the reproductive structure
(fruiting body) of a fungus. In this sense a mushroom, like a potato or
persimmon, is not an organism, but a part of an organism. However, the term
“mushroom” can also mean any fungus which produces a fleshy fruiting body
(that is, one that has substance). By this definition, not all fungi qualify as
mushrooms. Athlete’s foot fungus, bread molds, water molds, yeasts, and
mildews are examples of fungi which do not form fleshy fruiting bodies. The
term “mushroom” can also be applied in a more restricted sense to those fleshy
fungi like the cultivated mushroom whose fruiting bodies bear spores on
radiating blades called gills.
Many fungi are exquisitely constructed, and their life cycles are among the
most complex to be found. It is not the purpose of this book to explore their
biology, but it is necessary to consider briefly how mushrooms grow and
reproduce. All of the mushrooms in this book belong to two subdivisions of the
true fungi. Most of them produce their spores on the exterior of microscopic
club-shaped cells called basidia (singular: basidlum), hence they are called
Basidiomycetes. A smaller number produce their spores inside microscopic
saclike mother cells called asci (singular: ascus), hence they are called
Ascomycetes.
The fruiting bodies of the Basidiomycetes and Ascomycetes vary greatly in
detail and design, but their function is always the same—they perpetuate their
species by disseminating spores. A typical gilled mushroom (the most common
type of fruiting body) is a straightforward structure consisting of a cap, gills,
and (usually) a stalk (see diagram). A protective covering called a veil may also
be present, and if so, will frequently form a ring (annulus) and/or a volva when
it ruptures. The parts of a gilled mushroom are discussed in more detail on pages
14-18, and fruiting bodies of a radically different structure, such as puffballs, are
illustrated and discussed in their respective chapters.

Spore formation. At left is a typical club-

shaped basidium, with four small stalks
(sterigmata) on which the spores form.
At right is an ascus, inside of which spores
(usually eight) form.

basidium ascus
 cap

annulus or ring
(partial veil remnants)

volva
(universal veil remnants)

Parts of a gilled mushroom. Mature Amanita at left has cap, stalk, gills, annulus, and volva. The
partial veil covers the gills when young and breaks to form a ring (annulus) on the stalk, while the
universal veil at first envelops the entire fruiting body and breaks to form a volva (sack, collar, or
series of concentric rings) at base of stalk. Development of fruiting body is shown on pp. 270-271.
At right is a mature Marasmius, which has neither annulus nor volva, but often has an umbo (knob)
on cap.

In a gilled mushroom, millions of spores are produced on basidia which line

the gills. These spores are subsequently discharged and carried by air currents to
new localities. Each is theoretically capable of germination, but only a small
percentage land in a favorable environment. Spores germinate by sending out a
germ tube which branches to form many threadlike cells called hyphae. When
two spores of different but compatible strains (or “sexes”) germinate in close
proximity to each other, their hyphae merge to form hyphae with two nuclei (one
from each parent). These hyphae grow rapidly, forming an intricate network of
filaments called the mycelium or spawn (see photo on p. 43). The mycelium is
the vegetative portion of the fungus. The tips of the mycelial hyphae liberate
enzymes which digest food to support growth.
Once the mycelium has established itself and built up an adequate food
reserve, it becomes capable of producing mushrooms. Under favorable
conditions (for most species this means damp but not soggy, cool but not cold),
some hyphae bundle together to form knots of tissue which gradually develop
into fruiting bodies. When these fruiting bodies are differentiated but not fully
developed (that is, after they have a cap and stalk but before the cap expands),
they are called buttons. The stalk then elongates, pushing the cap above the
surface of the ground (or other substrate). Finally, the cap opens and the veil (if
present) breaks, exposing the gills on which spores form. The mature fruiting
body is essentially a bundle of threadlike hyphae (each with two nuclei), but the
filaments in the bundle terminate in either spore-producing cells (basidia),
specialized sterile cells (cystidia), or unspecialized cells (basidioies, or in some
cases, paraphyses). (See illustration on p. 6.)
Though they lack the sexual organs of plants and animals, mushrooms
reproduce sexually, i.e., genes are recombined so that offspring are not
genetically identical to parents. Gene exchange takes place in the basidium (or in
Ascomycetes, a special cell called the ascogonium). The two parent nuclei fuse,
doubling the chromosome number, then divide twice while replicating their
chromosomes only once—thereby reducing the chromosome number to half

5
 developing spores

basidioles

c^stidia

Cross-section of a single gill

as seen under the micro¬
basidia
scope. The edge of the gill is
at the bottom of this section.

that after fusion. The four remaining nuclei migrate to the tip of the basidium,
and walls form behind them to produce four spores—two of each strain
(“sex”), each with one nucleus. With their subsequent discharge the life cycle
is completed.
The above life cycle is typical of most Basidiomycetes, except that it is
often complicated by the presence of more than two strains (just as our life cy¬
cle would be unimaginably complicated by the existence of more than two
sexes!). Also, some mushrooms are capable of forming spores asexually.

MUSHROOMS AND THE ENVIRONMENT

IT IS the “role” of fungi to break things down, to give things back. One of the
more obvious laws of nature is that existing life must die if new life is to flourish.
Stale air must go out the window if fresh air is to come in. If there were no vehicle
for the disposal of dead matter, there would soon be no need for one—we would
all be buried under a blanket of inert matter. Fungi, along with bacteria, are
precisely that vehicle. They are nature’s recyclers, the soil’s replenishers. Plants
deplete the soil by extracting minerals to manufacture their food. Animals, in
turn, devour plants. In feeding on dead (or occasionally living) matter, fungi and
bacteria reduce complex organic compounds to simpler building blocks, thereby
enabling plants to re-use them. Thus, in a very profound sense, fungi are life-
givers as well as destroyers. To associate them only with death and decay—as so
many people do—is to do them, as well as our own ability to perceive, an
injustice.
Fungi can be divided into three categories based on their relationship to their
substrate (immediate environment). Parasitic fungi feed on living organisms.
Most serious fungus pests (such as wheat rust) fall in this category, but relatively
few mushrooms are parasitic. Their ranks include Cordyceps species (on insects,
insect pupae, insect larvae, and other fungi); Asterophora and Hypomyces
species (on other mushrooms); various polypores (on trees); and Sparassis
crispa (on tree roots). Some, like the common honey mushroom, Armillariella
mellea, are parasitic under certain conditions and saprophytic (see below) under
others.
Saprophytic fungi subsist on dead or decaying matter (wood, humus, soil,
grass, dung, and other debris). When there is an even distribution of nutrients in
the environment, the mycelium of a terrestrial fungus may grow outward at the

6
A large fairy ring of Marasmius oreades.

same rate in all directions, periodically producing circles of mushrooms on its

outer fringes. These circles or arcs are called fairy rings, presumably because
people once thought fairies danced in them. Many mushrooms are capable of
forming fairy rings, including the aptly named fairy ring mushroom (Marasmius
oreades), which grows on lawns. Each year the fairy ring gets larger as the
mycelium grows outward, until something finally impedes its progress (usually a
lack of food), and the mycelium dies or breaks up into arcs. By measuring the
annual growth rate, it has been estimated that some fairy rings in the Midwest
prairies are six hundred years old!
Mycorrhizal fungi comprise the third category. They form a symbiotic or
mutually beneficial relationship with the rootlets of plants (mostly trees) called
mycorrhiza (from myeo, fungus, and rhiza, root). The mycelium forms a sheath
of hyphae around the rootlets of the host and an exchange of nutrients takes
place. The rootlets provide the fungus with moisture and organic compounds
(such as carbohydrates), while the fungus aids the roots in the absorption of
phosphorus, inorganic nitrogen, and other minerals, and apparently also
provides added resistance to certain diseases. As a rule, mycorrhizal fungi cannot
grow without their hosts, and studies have shown that trees deprived of their
mycorrhizal partners do not compete successfully with those that have their
normal complement. This is especially true in poor or exposed soils, where trees
need all the help they can get, and mycorrhizal fungi have proved invaluable
in reforestation projects.
Many mycorrhiza-formers are host-specific, i.e., they grow only with one kind
of tree. For instance, Suillus pungens grows principally with Monterey pine,
while Amanita rubescens is monogamous with live oak (in our area). A tree,
however, may have several mycorrhizal associates whose relationships with the
tree are qualitatively different (Alexander Smith reports finding over 50 species
of mycorrhizal mushroms under an isolated Douglas-fir). In other words, each
type of mushroom occupies a different ecological niche. Many factors are
encompassed by the concept of niche, some of which we don’t understand. A
niche is not so much an organism’s habitat as its “profession”—what it does for
a living, or the “role” it plays in its biological community. Each kind of mush-
8 MUSHROOMS AND THE ENVIRONMENT

room reproduces and germinates successfully within a certain humidity

and temperature spectrum. Each extracts particular nutrients from its
environment. Thus two species may occur in the same habitat, but occupy
different niches. One may be taking the lignin from a log, another, the cellulose.
Or one may be feeding on the heartwood, another on the sapwood. Or returning
to the mycorrhizal fungi, one may be supplying phosphorus to the tree, and
another, nitrogen. Succession also occurs—as one type of mushroom exhausts
its nutrient supply, another takes its place. A living tree may harbor certain types
of fungal growth. As soon as it dies, new species will appear. Eventually the
wood is reduced to fragments or powder, at which point still other mushrooms
take over, with growth habits better suited to the changed conditions.

NAMES AND CLASSIFICATION

NAMES, like automobiles, are largely vehicles of convenience. You can’t claim
to have a profound knowledge of human beings without knowing at least some
of them on an individual basis. Recognition is a prerequisite to getting to know
someone, and a name is helpful in associating that person with a unique set of
identifying characteristics, whether it be his big nose and hairy face, or her long
legs and swift smile. Rather than saying “the 6 ft. 7 in. acrobatic forward of the
Philadelphia 76ers,” we say “Julius Erving” or “Doctor J.” Instead of “you
know, that bright red mushroom with white spots that grows under pine,” we say
Amanita muscaria, or “fly agaric.”
Names can also be descriptive. For instance, the red-headed woodpecker has a
red head and pecks wood. What’s more, names can reflect common bonds. Your
last name identifies you as a member of a group with similar genes, and provides
a clue as to your origins. Your first name defines you as an individual entity
within that group.
Unfortunately, relatively few mushrooms have colloquial English names—a
tribute, as pointed out previously, to our fungophobic roots. In this book I have
used popular names where they exist, and in some cases have capriciously coined
common names, but to do so in every case would only create confusion, as there
is no assurance they would be accepted. Therefore, if you really want to get to
know mushrooms, it is necessary to know their scientific names. People usually
groan when they hear this, and to be sure, the long Latin names are intimidating.
But so, at first, is a can opener—it’s just a question of familiarity. In fact, you
may already have mastered some Latin (scientific) names without realizing it—
e.g., Eucalyptus, Rhododendron, Hippopotamus. Memorization is made easier
by learning the meanings of the names. For instance, Lactarius rubrilacteus
(rubri-red, lacteus-milk) exudes a red “milk” when cut. See “What It All
Means” on p. 899 for more details. Don’t get bogged down in pronounciation. It
doesn’t really matter how you say something as long as you communicate it.
Even taxonomists don’t agree on how some names should be pronounced!
As you begin to use scientific nomenclature, you’ll discover its many advan¬
tages. Common names do not necessarily reflect natural affinities. Hedge nettle
is not a nettle, and poison oak is by no means an oak. Likewise, the names
meadow mushroom, honey mushroom, matsutake, and horse mushroom
provide no clues as to which, if any, have common bonds or similar charac¬
teristics. Also, common names are not universal. For instance, Boletus edulis
has dozens of regional names, and memorizing all of them would be almost as
difficult as getting everyone to agree on one of them!
NAMES AND CLASSIFICATION 9

In contrast, scientific nomenclature transcends cultural and regional barriers.

It is used by naturalists and biologists throughout the world, and it is designed to
reflect natural relationships. It employs a binomial system in which each kind
of organism has two names. The second name, the species, is the kind of
organism; the first name, the genus (plural: genera) is a collection of species
with very similar traits. The species epithet is meaningless without the genus
name (or its abbreviation) attached (we never sign a check with only our first
name!). The names are mostly Latin because that language was universally
fashionable in learned circles when the binomial system was devised. The beauty
of the binomial system is that it indicates commonality while simultaneously
expressing singularity.* Amanita calyptrata (the coccora) and Amanita
phalloides (the death cap) are different species belonging to the same genus, just
as Canis lupis( wolf), Canis latrans (coyote), and Canisfamiliar is (dog) belong to
the same genus. Yet the common names of these organisms don’t indicate that
they are closely related!
Some kinds of similarities are clearly more fundamental than others.
Therefore, it was deemed necessary to erect a hierarchy of classification to
indicate the degree of commonality. Genus and species are only two levels in that
hierarchy. Just as species with common characteristics are grouped in a genus,
several genera with common features are grouped in a family (e.g., the red fox,
Vulpes fulva, belongs to the same family as the dog, but not to the same genus).
Families are in turn grouped in an order, orders in a class, classes in a
subdivision or division (or in the case of animals, in a phylum), and divisions
in a kingdom. At the other end of the scale, slightly different populations of a
single species are designated as subspecies, varieties, or forms.** Here is the
complete classification scheme for three mushrooms—the blewit, shaggy mane,
and inky cap.

Category Blewit Shaggy Mane Inky Cap

Kingdom Fungi Fungi Fungi
Division Eumycota Eumycota Eumycota
Subdivision Basidiomycotina Basidiomycotina Basidiomycotina
Class Hymenomycetes Hymenomycetes Hymenomycetes
Order Agaricales Agaricales Agaricales
Family Tricholomataceae Coprinaceae Coprinaceae
Genus Clitocybe Coprinus Coprinus
Species nuda comatus atramentarius

All three of these organisms are fungi—they possess neither photosynthetic

compounds nor seeds, and their vegetative phase is comprised of threadlike cells
(hyphae). Since none are ameba-like, they are not slime molds and hence
belong to the division Eumycota. All three produce their spores on microscopic
cells called basidia, hence they belong to the subdivision Basidiomycotina. All
forcibly discharge their spores from basidia which form on radiating blades
(gills), therefore they belong to the class Hymenomycetes and the order
Agaricales. However, the blewit has pinkish spores while the shaggy mane and
inky cap have black spores—which is one reason they are placed in different
families. Not only do the shaggy mane and inky cap belong to the same family
(the Coprinaceae), but they also belong to the same genus, Coprinus, because
*The singularity of a population of organisms; our personal names, on the other hand, indicate the singularity of
individual organisms.
**In this book the term “variety” is used in its ordinary English sense to mean a “type” of organism. Only when one
species is being discussed and “variety” is abbreviated (var.) is it applied in its scientific sense. The same goes for
“form” (f.).
10 NAMES AND CLASSIFICATION

both have gills that deliquesce (liquefy) at maturity. However, the shaggy mane
has a tall, shaggy, cylindrical cap while the inky cap has a smoother, broader,
oval or conical cap; consequently they are recognized as distinct species.
Note the suffixes at the levels above genus: —ceae denotes a family; —ales
indicates an order; and —cetes denotes a class or larger category.
It must be remembered, however, that this elaborate classification scheme is
contrived. It is our attempt at boxing and categorizing nature. There are
common gene pools and definite lines of evolution, but no such clearcut
categories exist. Thus, the definition and interpretation of species, genera,
families, etc., is largely a matter of opinion. Disputes invariably arise, many of
which have not been resolved. For instance, at the genus level and above, there
is the problem of deciding which similarities among fungi are fundamental
(indicators of common origin), and which are coincidental or superficial, or the
result of convergent evolution.* The microscope has been a tremendous help in
uncovering “hidden” similarities, but it has also exacerbated the confusion by
introducing a vast new set of criteria on which to pass judgment. The result is a
nomenclatural nightmare, from the upper echelons of the hierarchy right on
down to the species level.
Mycological literature is as riddled with contradictions as a Suillluspungens is
with maggots. Anyone who has used more than one mushroom book can testify
to the frustration of finding different names applied to the same fungus
(synonyms), or one name applied to several different fungi (homonyms). For
instance, Clitocybe nuda (the blewit) is better known as Lepista nuda, and was
formerly known as Tricholoma nudum.lt has been incorrectly called Tricho-
loma personatum, and in Europe is also known as Rhodopaxillus nudus! For an
even more confusing example, see the list of synonyms for Trametes occidentalis
on p. 550.
This confusion is partly the result of disagreement as to what exacly con¬
stitutes a “genus” or “species”—a difference in philosophy that is known in
taxonomic circles as the battle between the “lumpers” and “splitters.” The
“lumpers” are conservative in their approach. They interpret genera or species
broadly, allowing for a good deal of variation—in other words, they tend to stress
similarities between mushrooms rather than differences. “Splitters,” on the other
hand, are forever describing new genera and species based on the most minute—
but not necessarily insignificant—differences. Both approaches have their
advantages and drawbacks, and both are self-defeating when carried to an
extreme.**
The important thing to realize is that the system of classification used in this
book is by no means definitive. It represents an amalgamation of various inves¬
tigators’ views of the fleshy fungi, plus the overriding consideration of usefulness
to the amateur (since this book is designed for amateurs). Some of the names
used will undoubtedly be invalidated in the near future. A few have not been
validly published and are therefore placed in quotation marks. Synonyms have
been provided and homonyms elucidated. But the inherent advantages of the
binomial system of nomenclature will not be fully realized until stabilization is
achieved and one name is agreed upon for each kind of mushroom. In
exceptional cases like that of the blewit, it is perhaps easiest in the meantime to
use the common English name—if there is one.
*The fins and torpedo-shaped bodies of sharks and killer whales are independent adaptations to a similar en¬
vironment, not indicators of common origin. This phenomenon is called convergent evolution.

**One radical “lumper” I know recognizes only two kinds of mushrooms—the “pickers” and the “kickers” (those
that deserve to be picked, and those that deserve to be kicked!).
 11

COLLECTING MUSHROOMS
MUSHROOM hunting is not simply a matter of traipsing through the woods
after it rains. It is an art, a skill, a meditation, and a process. If you proceed at a
careful, deliberate rate, you’ll enjoy much more success than if you rush around
frantically picking whatever mushrooms you see, then stuff them in your basket,
bring the whole mess home and dump it on your table. Mushrooms collected
in this manner are likely to wind up in the garbage, unidentified and unappre¬
ciated.
Don’t just collect, but observe the mushrooms—and their surroundings. In
the process you’ll discover many other clandestine wonders you were previously
unaware of (see photographs on pp. 28-29). Be selective—pick only distinctive
species in good condition. Y ou enhance your chances of successful identification
immeasurably by collecting several specimens of each kind of mushroom. This is
absolutely essential, because you have no other means of assessing variation
within a given species. Since mushrooms decay rapidly and identification can
be a time-consuming process, don’t pick every kind you see. It’s better to fill your
basket with many good examples of a few distinctive species (in which case your
chances of identification are good) than with one or two specimens of many
species (in which case your chances of identification are very poor).
Don’t assume, however, that two mushrooms are the same species simply
because they’re growing together. Judge each on its own merits. If you’re
uncertain, assume for safety’s sake that they’re different, and treat them as such.
As you become more adept at observation, your ability to identify fleshy fungi
will “mushroom”—but only after a solid foundation or “mycelium” of ex¬
perience is laid.
It is far better to learn a few species well than a large number superficially. The
novelty of the “Easter egg” approach wears off quickly as its futility becomes
apparent. If possible, choose a specific quarry for each hunt. Suppose it’s
January and you’re going for a walk in a local live oak woodland. By using the
chart on pp. 48-51 and consulting the description of the blewit (Clitocybe nuda),
you find that the blewit is often abundant under oak in January. The next step is
to familiarize yourself with its fieldmarks: bluish-purple color, stocky stature,
absence of a veil, citrus odor, etc. Your mushroom hunt is thus transformed into
a blewit hunt. Of course, nothing stops you from gathering other interesting
fungi you encounter, but focusing your attention on the blewit insures that you’ll
learn something about blewits even if you don’t find any (namely, that the
locality and/or weather conditions were not conducive to its fruiting).
The desirability of this approach is underscored by the excitement it lends to
the hunt. It is much more gratifying to find something you are specifically trying
to find. And you’ll be more likely to remember what it looks like and where it
grows, so you can return to harvest more. By focusing on several species as the
season unfolds, you will develop a quicker and keener appreciation of what
grows where and what environmental factors they respond to. Many of the more
distinctive species occur throughout the world, so the knowledge you
accumulate will serve you elsewhere!
Always dig up unknown mushrooms so as not to miss the volva, if present. There’s no room for care¬
lessness, as shown here: Agaricus arvensis, at left, lacks a volva and is edible; Amanita ocreata, at
right, is furnished with a volva and is deadly poisonous!

EQUIPMENT
Foraging for fleshy fungi requires little in the way of sophisticated para¬
phernalia. The bare essentials are:
A rigid container for carrying the mushrooms. There’s no point in picking mushrooms
unless you transport them home in decent condition. A broad, shallow basket is best,but a
cardboard box or bucket will do unless it’s raining. Paper bags sag and the mushrooms
get crushed. DON’T USE PLASTIC BAGS! Mushrooms, like people, have to “breathe.”
Plastic bags trap moisture, making the mushrooms “sweat” and rot more rapidly.
Waxed paper is necessary when collecting mushrooms for identification. It provides
support for mushrooms within the basket and also keeps them separated. Never mix
unknown species together. Wrap each type separately and arrange them carefully in the
basket with the heavier ones on the bottom. Tall specimens like Agaricus and Amanita
species will bend unless placed upright (mushrooms exhibit negative geotropism: their
caps turn away from gravity so as to orient the gills downward). Small paper bags are
useful when harvesting familiar edible species, but again, don’t use plastic bags!
A knife or trowel is a must for digging up mushrooms or detaching them from trees. A
knife is also handy for cleaning edible species you are already familiar with, but always dig
up unknown mushrooms so as to ascertain whether or not a volva (sack) or “tap root” is
present. The telltale volva of the deadly Amanitas is usually buried in the ground!
A pencil and small notebook or index cards are useful for taking field notes and spore
prints.
Bread, cheese, and fruit are essential if you’re always hungry like I am. I make a practice
of stocking my basket generously. Then each time I put a mushroom in my basket, I’m
compelled to put something from the basket in my mouth! If you find some edible
Agaricus buttons, put them in a sandwich! Not known for wasting opportunities, the
French carry this tradition one step further—they bring wine, goblets, and a table cloth,
and pause for a picnic every half hour. The advantage of this strategy is obvious—you
needn’t find any mushrooms to have a good time!

12
EQUIPMENT 13

Binoculars are handy in open country (e.g., pastures). They enable you to distinguish at
a distance giant puffballs (Calvatia species) and horse mushrooms (Agaricus arvensis and
A. osecanus) from rocks and other assorted “pseudocarps.” And of course, they allow you
to watch birds and mammals as well.
A three- or four-pronged rake or cultivator and a small hand cultivator are necessary
if you want to find truffles and false truffles, unless you have the services of a truffle
hound or muzzled pig. The rakes will enable you to locate these elusive underground
fungi by sifting through the forest duff and scraping the topsoil beneath it. (But re¬
member: this practice can be unsightly as well as destructive, so don’t do it on a wide¬
spread basis, be discreet, and always cover up the holes you dig.) See the chapter on
truffles for more details.
Other optional equipment includes: a hand lens, compass, stick (for probing brambles
and “mushrumps”), a field guide (for leisurely use on a sunny day), small jars or vials (for
delicate specimens, such as Mycenas), a damp cloth or brush for cleaning edible species,
rainboots and other rain gear, gloves for frigid winter mornings, and photographic
equipment (usually too cumbersome except for special picture-taking expeditions).

FIELD NOTES
Is it growing on the ground or on a log? Is it near a tree? What kind(s)? Are
familiar types of mushrooms growing nearby? Which ones? If it’s growing on
wood, is the wood coniferous or hardwood? Living, recently felled, or in an
advanced stage of decay? If on the ground, is the humus layer deep? Is the ground
disturbed? Is there a road, trail, parking lot, or laundromat nearby?
You should automatically ask yourself these questions every time you find a
mushroom. Observation begins in the field. After all, in picking a mushroom
you are leaving behind the vegetative portion of the fungus. It is folly to depart
without some idea of the niche that fungus occupies in the larger scheme of
things. Field (or mental) notes should include:
Date, weather conditions, abundance (how many times you observed a
particular species), growth habit (solitary, scattered, gregarious, clustered,
in fairy rings, etc.), substrate (humus, soil, grass, moss, dung, wood, etc.),
vegetation (the kinds of trees and shrubs within 50 feet).
If growing on wood: stage of decomposition, type of wood (hardwood or
conifer), type of tree (if discernible), effects on the wood (see chapter on
polypores on p. 549).
If growing on dung: type of dung, stage of decomposition
If growing on ground: type of ground (disturbed, cultivated, hard-packed,
sandy, charred, etc.)
Don’t restrict your observations to specimens that you collect. After you’ve
gathered a representative sampling of a particular species, continue to note its
habitat each time you encounter it.
With terrestrial fungi, it is important to note all types of trees within 50 feet
because mycorrhizal species grow in association with rootlets that may be quite a
distance from the trunk of the host. Usually there are several kinds of trees in the
vicinity, and you have no way of knowing which (if any) is the mycorrhizal
associate. However, through repeated observation many possibilities can be
eliminated. For instance, if you find Suilluspseudobrevipes growing under pine,
you cannot conclude that there is a relationship between the two. But if you find
that Suillus pseudobrevipes always grows with pine and nowhere else, then an
intimate relationship of some kind can be inferred.
14

IDENTIFICATION AND TERMINOLOGY

IF YOU take the time to seek out mushrooms, it only makes sense to exercise the
extra care and trouble necessary to get them home in beautiful condition. Handle
them gingerly, don’t leave them in stuffy places like cars, and don’t shift them
unnecessarily from box to box. Always conduct your studies on fresh material,
preferably the day you pick them. Coprinus species digest themselves in a few hours,
and many types quickly lose their original color or are devoured overnight by
maggots. If you’re pressed for time, at least sort them out and separate the worm-
riddled specimens. Then refrigerate the ones you wish to save or spread them out in a
cool, dry place where they can “breathe.”
Now let’s assume you’ve taken field notes, brought several species home, and are
ready to study them. For the diligent and disciplined toadstool taxonomist, a
detailed written description of each species is a must. For practically everyone else,
compiling a written description is a tedious affair which tends to detract from the
enjoyment and spontaneity of the hunt. However, it is an ideal tool for learning how
to look at mushrooms critically, so try it at least a few times.
The basic terminology for identifying and describing gilled mushrooms is outlined
here. Fruiting bodies with a radically different structure, such as puffballs, are
illustrated and discussed in their respective chapters. Unfamiliar terms not
illustrated or defined here can be looked up in the glossary. Remember to base your
observations on as many specimens of each species as possible. The value of written
descriptions is enhanced when accompanied by sketches, photographs, and spore
prints of fresh material.

MACROSCOPIC CHARACTERISTICS
SIZE

Size is important for purposes of comparison. The terms large, medium, small, and
minute cannot be given absolute measurements, but they communicate a characteristic
size range that you will quickly learn to appreciate. The size of a fruiting body is
dependent on three major factors: age, amount of moisture available, and genes. Since
mushrooms grow very quickly, those that fruit during rainy weather are apt to be larger
than those that fruit during a dry spell—subject, of course, to genetic constraints.
The measurements given in descriptions and keys represent average size ranges; those
in parentheses indicate unusual dimensions, but do not take into account extremes due to
extraordinary conditions. The metric system is used, but a conversion rule is provided on
the back cover of this book. Also, it’s easy to remember that 1 inch equals approximately
2j/2 centimeters, or 2 inches equals 5 centimeters (or 50 millimeters).
COLOR
Color is one of the most noticeable features of any fungus, but is also one of the most
deceptive and variable. Unfortunately, beginners tend to attach undue importance to
color at the expense of more critical characteristics. This inevitably leads to
misidentification because many mushroom pigments are highly sensitive to
environmental influence. Direct sunlight or prolonged rain can bleach a mushroom
drastically, and I will never forget my experience following a prolonged rainy spell in
which practically every mushroom growing under redwood had a reddish cap!
(Apparently pigments from the redwood had dissolved in the drip-off and been absorbed
by the mushrooms.) None of these mushrooms would have keyed out properly unless this
phenomenon were taken into account! Color may also depend on age. An immature sea
MACROSCOPIC CHARACTERISTICS 15

gull is brown, while an adult is black and white. Likewise, an immature Hygrocybe conica
is red, orange, or yellow, but blackens as it ages.
Therefore, color should always be used in conjunction with other characteristics. The
most striking feature of Amanita muscaria is its bright red cap, but a slew of other mush¬
rooms are also bright red. Furthermore, Amanita muscaria has a yellow-capped and
white-capped form, and even the red-capped variety will fade to orange or even whitish. It
is the red cap combined with the presence of warts and a volva that render A. muscaria
distinct.
The color of the stalk is not as variable because it is sheltered by the cap, but the gill
color often changes as the spores mature. In fact, the disparity in gill color may be so great
that young and old specimens can be mistaken for different species unless intervening
stages are found.
Describing color is another problem. Color pictures are a great help, of course, but
even they can’t show the degree of variation within a given species. Standardized color
charts are available, but they represent an extra expense and pose problems of their own.
In this book, colors are described using familiar terms where possible, plus a few specific
ones (e.g., vinaceous) which are defined in the glossary.
COLOR CHANGES
The tissue of many mushrooms oxidizes when exposed to the air—that is, it undergoes
one or more color changes when bruised. This may occur instantaneously, like the blueing
of the tubes and flesh in Boletus erythropus, or slowly (over several hours), like the red¬
dening of the flesh in Amanita rubeseens. Bruising reactions should be looked for on the
surface of the cap, gills, and stalk, and on the cut flesh within the cap and stalk.
TEXTURE
The texture of the fruiting body is often significant—i.e., soft, watery, spongy, brittle,
tough, leathery, corky, woody, etc. The dried fruiting bodies of some types (notably
Marasmius) revive completely when moistened.

ODOR AND TASTE

It comes as a pleasant surprise to many people that mushrooms have distinctive tastes and
odors. For instance, Lactarius torminosus is excruciatingly hot (peppery), while Agaricus
subrufescens is sweet. Marasmius copelandi reeks of garlic and is often smelled before it is
seen, Clitocybe odora smells like anise, Russula fragrantissima like maraschino cherries, and
Phyllotopsis nidulans like sewer gas, while Armillaria ponderosa has a spicy odor—a
provocative compromise between red hots and dirty socks.
The problem with odor and taste is in the terminology. They are essentially chemical tests,
but each person’s “laboratory” is different. What smells like sauerkraut to one person smells
like sweet-and-sour sauce to another. Moreover, mushrooms with a “mild” odor may be very
pungent when crushed, or may only develop an odor at maturity, and odoriferous species may
be odorless if waterlogged. Consequently, odor and taste are not noted in this book except
where useful in identification.
Incidentally, almost any mushroom can be safely tasted by chewing on a small piece of the
cap and then spitting it out. However, it is not a good policy to sample unkown mushrooms,
particularly Amanitas. Do so only when it is called for in the keys or descriptions.
CAP
The cap (or pileus) is the structure that supports the spore-producing surface (gills, pores,
etc.). The skin of the cap is called the cuticle (or pellicle, if it peels easily). A viscid cap is
sticky when moist, often glutinous (slimy) when wet, and sometimes assumes a glossy
appearance as it dries. Debris stuck to the cap surface is a telltale sign that it was viscid when
moist. Also, if you press your lip or wet finger against the surface, it will cling or feel
slightly sticky. A hygrophanous cap appears watery (or even translucent) when moist and
opaque when dry, and fades dramatically as it loses moisture (Psathyrella candolleana, a
16 IDENTIFICATION AND TERMINOLOGY

common lawn lover, is a good example). A dry cap is neither viscid nor hygrophanous. It
often has a dull, unpolished appearance, but will naturally be moist or soggy in wet weather.
Other features to note are: size, shape (p. 17), color and color changes, surface
characteristics (whether smooth, scaly, granulose, fibrillose, warty, etc.), and margin (inrolled,
incurved, straight, or uplifted; and striate, translucent-striate, or not striate).

FLESH
Note color, color changes, texture, thickness, odor when crushed, and taste (don’t
swallow!).

GILLS
Gills (or lamellae) are the thin, radiating blades found on the underside of most
mushrooms, including the commercially cultivated variety. Features to note include: mode of
attachment to the stalk (p. 17), spacing, thickness, depth, forking pattern (if any), and color (in
both immature and mature specimens). Shorter gills (lamellulae) are often interspersed with
longer ones, but they don’t usually have taxonomic significance.

STALK
The stalk is the stemlike structure on which the cap is mounted. Its function is to thrust the
cap above the ground so the spores can be discharged into the air. Many wood-inhabiting
forms lack a stalk because the wood serves the same purpose. The technical term for stalk is
stipe, but I see no reason to clutter our vocabulary with yet another monosyllable starting
with st-. Hereafter, the terms stalk and stem are used interchangeably.
Features to note include: size, color and color changes, shape (p. 17), position (p. 17),
texture (fleshy or cartilaginous, hollow or solid or stuffed with a pith), surface characteristics
(fibrillose, scaly, smooth, etc ), viscidity, and presence or absence of an annulus and volva
(see below). Stalk width should be measured at the top unless otherwise indicated.

VEIL
A veil is a layer of specialized tissue that initially protects the developing mushroom and
then breaks up or collapses so the spores can be released. Some mushrooms have more
than one veil, others have only one veil, and many lack a veil altogether. A persistent veil
leaves visible remnants on the stalk and/or the cap after breaking; an evanescent veil
disappears and consequently can only be detected in the button stage. A membranous veil is
skinlike or kleenexlike and usually persistent; a fibrillose veil is hairy (composed of fine
fibers) and either disappears or forms a belt of collapsed hairs on the stalk; a Cortina is a
cobwebby fibrillose veil; a glutinous (slimy) veil is either evanescent or deposits a layer of
slime on the stalk and/or cap.
There are two basic types of veils: a partial veil (or inner veil) extends from the margin of
the cap to the stalk; it covers the gills (or pores) when young and frequently forms an annulus
(collar or ring) on the stalk. A universal veil (or outer veil), on the other hand, surrounds
most or all of the button mushroom. Sometimes it forms a volva (see below) after breaking;
sometimes it adheres to the underside of the partial veil (as in Agaricus arvensis); in other
instances it forms a “stocking” of scales on the stalk (as in Lepiota clypeolaria), and in still
others it completely disappears. Needless to say, not all veils fit conveniently into one or the
other of these two categories, and in this book the all-encompassing term veil is used for both
universal and partial veils except when the difference between the two is clearcut and of critical
importance (e.g., in Amanita).

ANNULUS
If an annulus (ring) is formed by the veil, note the color, texture (whether membranous or
fibrillose), shape (collarlike, skirtlike, or sheathlike), and position on the stalk (superior or
apical, median, inferior or basal).
MACROSCOPIC CHARACTERISTICS 17

SHAPE OF THE CAP

cylindrical

plane depressed funnel-shaped

ATTACHMENT OF THE GILLS TO THE STALK (as seen in longitudinal section)

free adnexed sinuate adnate decurrent

(not attached) (narrowly attached) (notched) (broadly attached) (running down
the stalk)

SHAPE OF THE STALK

tapering downward equal tapering upward enlarged below bulbous

(club-shaped)
POSITION OF THE STALK

off-center lateral absent

(eccentric) (sessile)
18 IDENTIFICATION AND TERMINOLOGY

VOLVA
In some mushrooms the universal veil is developed to such an extent that, upon breaking, it
leaves visible remains at the base of the stalk in the form of a sack, free collar, or series of
concentric scales or rings. These remains are called the volva. The different types of volvas are
illustrated under Amanita on p. 264. Volvariella, the stinkhorns, and certain stalked
puffballs also have a volva. If small pieces of volval (universal veil) tissue adhere to the
cap, they are called warts; a large piece of tissue is called a volval patch.

MYCELIUM
The mycelium is a loosely organized mass of threadlike cells (hyphae) which are invisible
to the naked eye except when they bundle together to form thicker strands (called
rhizomorphs). Close scrutiny of the forest humus will usually reveal the presence of
numerous mycelial strands, but they are virtually indistinguishable from each other without
fruiting bodies present. There are a few exceptions, like Chlorosplenium aeruginascens, an
Ascomycete which stains its substrate (a log or piece of wood) bluish-green. As a rule,
however, it is the fruiting body alone that provides clues to the identity of a fungus. This is one
reason why the classification of mushrooms has lagged behind that of plants—we’re
handicapped at the outset because our studies are restricted to only one aspect of the
organism, its “fruit.”

SPORE COLOR AND SPORE PRINTS

Spore color is an extremely useful character, particularly in the gilled mushrooms. Unlike
the color of the fruiting body, it is relatively constant for each species and not as susceptible
to environmental influence. Though individual spores can’t be seen with the naked eye, their
color in mass can be ascertained by taking a spore print. Just cut off the cap of any mature
mushroom and lay it gills down on a piece of white paper (covering it with a glass or bowl
helps protect it from air currents). In 2-6 hours you’ll usually have a spore print. Fruiting
bodies which are too young or too old, or too soggy or too dry, will not give spore prints, and
sterile specimens are sometimes encountered. Spore prints can also be obtained from boletes,
teeth fungi, chanterelles, coral fungi, and sometimes polypores.
Always take a spore print to determine the spore color of an unknown mushroom,
especially if you want to eat it. The color of the mature gills may indicate the spore color, but
not often enough to be completely trusted. If you are frustrated by the delay, carry white
index cards with you when you forage. Then each time you collect several individuals of the
same species, you can decapitate one, place it on a card, wrap it in waxed paper, and put it gills

Black spore prints of Panaeolus campanulatus.

MACROSCOPIC CHARACTERISTICS 19

down in the basket in such a way that it won’t be crushed. By the time you get home you’ll have
a spore print!
There are several short cuts for determining spore color, but I recommend them only for
experienced collectors because they are not completely reliable. When mushrooms grow in a
cluster, the lowermost caps will often be covered with spore dust from the upper ones (a wet
finger will remove it). Spores borne aloft by air currents will often coat the cap of any
mushroom, and falling spores are often trapped by the veil remnants (if present) or by the
stalk, especially if it is sticky.
Spore color is traditionally broken down into several broad categories, and assigning your
spore print to the correct category can be tricky at first. For instance, don’t expect “pink”
spores to be bright pink or “purple-brown” spores to be purple. “Pinkish” spores are really
closer to flesh-color, while “purple-brown” describes spores which could just as well be called
“oil-slick aubergine” (deep brown in mass and dark reddish-brown under the microscope).
Only through practice and comparison will you learn to assess the color of a spore print
correctly. The thickness of the spore print also affects the color (the heavier the deposit, the
darker it will be), as does the moisture content.
Always take a spore print on white paper (a colored background distorts color perception).
White spores will show up on white paper when viewed at an angle. Or you may wish to
position the cap so that half of it is on white paper and half on black. Special cards can be
designed for this purpose.

MICROSCOPIC CHARACTERISTICS
The microscope has had a great impact on the taxonomy of fungi, especially in the last fifty
years. However, the vast majority of people do not have access to a microscope, so
microscopic features are not stressed in this book. For those who do have a microscope, spore
characteristics (shape, size, and ornamentation) have been listed for each species, as spores are
the most easily seen microscopic cells. Spore size is measured in microns (or micrometers).
A micron is one thousandth of a millimeter. For each family or order of mushrooms, some
typical spores have been illustrated—not for the purpose of identification so much as to give
those without a microscope an idea of what they look like.
Other microscopic criteria used in the taxonomy of gilled mushrooms may be even more

ARRANGEMENT OF THE GILL HYPHAE (as seen in cross-section)

divergent convergent

STRUCTURE OF THE CAP CUTICLE CYSTIDIA

filamentous lance¬ flask- horned harpoon¬

shaped shaped like
20 MICROSCOPIC CHARACTERISTICS

significant than spore characters, but are not as easily observed. These criteria include:
orientation of the hyphae in the gill tissue, structure of the cap cuticle, and shape and size of
cystidia (specialized sterile cells) on the gills, stalk, and cap.
Spores can be examined by taking spore dust from a spore print and mounting it on a slide
with a drop of water and a cover slip. Cystidia, basidia, and gill hyphae are best seen by
making a thin cross-section of a gill with a razor blade and mounting it in a similar manner
(it takes practice!). The cap cuticle, cap tissue, and stalk are also best observed
in cross-section

CHEMICAL CHARACTERISTICS
The ways in which mushrooms react to different chemicals have also assumed great
importance in their classification. For instance, the genus Lyophyllum was erected to embrace
various white-spored agarics whose basidia contain granules that darken when heated in
aceto-carmine. Since most people are not equipped to conduct tests of this sort, they are not
stressed in this book. But there are two particularly useful chemicals that every serious
mushroom hunter should try to get: a 5-10% aqueous solution of potassium hydroxide
(KOH), and an iodine solution called Melzer’s reagent (see the glossary for formula). Spores
which stain bluish-gray to bluish-black in Melzer’s reagent are said to be amyloid. Spores
which turn brown or reddish-brown are dextrinoid. Many spores are neither amyloid nor
dextrinoid. The test is most useful on spores that are not deeply colored to begin with. It is
easily observed under the microscope, but can also be assessed by placing spore dust on a glass
slide (paper itself may be amyloid), treating it with Melzer’s reagent, then holding the slide
over a piece of white paper in order to easily see the color change, if any (the reaction
takes place within a few minutes). One of the ingredients in Melzer’s reagent, chloral hydrate,
is difficult to obtain, but other iodine solutions can be used in a pinch.

PRESERVING MUSHROOMS FOR FUTURE STUDY

Part of mushrooms’ mystique is their ephemerality—they’re literally here today and gone
tomorrow, and there is no adequate means of preserving their beauty. For scientific purposes,
however, they can be preserved by simply drying them out—they shrink and fade in the
process, but their anatomical (microscopic) features remain intact. Of course, for dried
material to have value it must be accompanied by a spore print and precise description of the
fresh fruiting bodies, plus a photograph if possible. Drying is best accomplished on a rack or
screen, using a light bulb or hot plate as a heat source. It is more important for the air to
circulate freely (thus carrying off moisture) than for the temperature to be extremely hot. For
this reason ovens are not suitable except in a pinch. Small or very fragile specimens may
“cook” when heated, and should instead be air-dried with the help of silica gel.

For ornamental purposes, mushrooms can be freeze-dried or encased in cubes ofplasticresin. Orthey
can be sliced and pressed in a book, like this Chroogomphus. Preserving mushrooms for consumption
is discussed in the chapter on mushroom cookery (p. 888).
 21

HOW TO USE THE KEYS

IF DESCRIPTIONS and photographs are the meat of this book, then keys are its
skeletal structure. Keys are tools designed to aid in the identification process. With
the exception of the two pictorial keys (pp. 52-55 and 61 -62), the keys in this book
are dichotomous. That is, they consist of a series of contrasting paired statements
(couplets). You are asked to decide which statement in a given couplet is applicable
to the mushroom in question. Having made your choice, you are then referred to the
number of another couplet, where you again make a choice. This process is repeated
until you are given the name of a mushroom (or group of mushrooms) and its
appropriate page number (if a page number is not given, it means that particular
mushroom is not treated or described beyond the key).
The dichotomous keys begin on p. 52 with “Key to the Major Groups of
Fleshy Fungi.” Under each major group of fungi you’ll find a key to families;
under each family, a key to genera; under each genus, a key to species. Keying will be
a laborious process at first. However, as you gain experience you can take shortcuts.
For instance, if you already know that your mushroom is an Agaricus, you needn’t
consult the keys to families and genera of gilled mushrooms. You can proceed
directly to the Agaricus key in order to determine the species.
The key to the proper use of a dichotomous key is an understanding of its
limitations. The following key to a banana, banana slug, hat, rabbit, and sea urchin
will admirably illustrate the assets and pitfalls of the dichotomous key. It’s designed
for the express use of creatures from the planet Fazoog, but I’m sure they won’t mind
if we use it here.
1. Object yellow.2
1. Object not yellow.4

2. Object more or less cylindrical .3

2. Object not cylindrical .hat

3. Object peeling easily . banana

3. Object not peeling easily . banana slug

4. Object purple and spiny . sea urchin

4. Not as above.5

5. Object moving of its own accord when poked, with four stumps or
projections on its underside . rabbit
5. Not as above.hat

Now let’s pretend you’re trying to identify a rabbit. You should always begin with
the first couplet, which in this case asks you to decide whether or not the object is
yellow. Presumably you’ll choose “not yellow.” Then, as indicated to the right of
“object not yellow,” you proceed to couplet #4 (thereby skipping #’s 2 and 3). Since
the object is not purple and spiny, you move on to #5. At this juncture you are
confronted with a more difficult decision. Does the object move of its own accord
when poked and have four “stumps” (legs)? If so, it is a rabbit, at least according to
the key.
But keys can be worthless unless accompanied by detailed descriptions. For
instance, if you tried to identify a tiger using this key, you would arrive at “rabbit.”
Unless a description of a rabbit is provided, you have no way of knowing whether it
22 HOW TO USE THE KEYS

is indeed a rabbit you have, or an entity not included in the key. Furthermore, if the
word “edible” or “harmless” appeared next to “rabbit,” you would find yourself in
serious trouble. Imagine the consequences of attempting to eat a tiger under the
mistaken impression that it is a rabbit (more likely the tiger would eat you!).
Similarly, a good key to mushrooms does not contain information on edibility.
Instead it refers you to a detailed description of the mushroom. Always compare the
mushroom you are trying to identify with the appropriate description as indicated in
the key (in a few cases, no page number is given, which means that the mushroom is
not treated or described beyond the key). Resist the urge to “fit” the mushroom to the
description or vice-versa. A major discrepancy between your mushroom and the
description has three possible explanations:
(1) IT IS THE MUSHROOMS’ FAULT. Mushrooms have not seen pictures or
descriptions of themselves, and do not know what they are “supposed” to look like
(even if they did know, why should they look the way they are supposed to—would
you?). We can only summarize what a given species usually looks like. A good key
will allow for a certain degree of variation. For instance, you will notice in the sample
key that “hat” appears twice, because some hats are yellow and others, thank God,
are not. The key does not account for other possibilities, however. Suppose you have
a dead rabbit. Since it does not “move of its own accord when poked,” it would be a
hat according to the key. Or supposing you find a peeled banana, or a blackened
banana (the kind my father likes to eat). It will not be yellow, and consequently will
not key out properly. A better key would account for these possibilities, but a key
cannot account for every possible variation without becoming hopelessly unwieldy.
At first you’ll have difficulty assessing just what is typical and what is not. The only
solution to this problem is to use several specimens of each type of mushroom—at
different stages of development if possible. You are then in a better position to decide
what is typical (i.e., if you have ten rabbits and one of them is dead, you conclude
there is something “wrong” with the dead one, and still use the key correctly.)
(2) IT IS YOUR FAULT. People often go astray because they misread a couplet
or inadvertently go to the wrong number, or exercise poor judgment. There are
several ways to minimize these mistakes. First and foremost, always read both
statements in a couplet before deciding which one is true. Second, if you are unsure
of your choice (as you undoubtedly will be at times), make a notation and choose the
one that seems most likely. If it later proves to be wrong, you can go back and try the
other choice. Remember the purpose of a key is not to identify, but to eliminate.
There are more than 1000 possibilities as to what any unknown mushroom can be. If
the keys eliminate 996 of these possibilities, they have done their job. It is then up to
you to carefully compare the descriptions of the remaining three or four species, and
decide which—if any—is the correct one. In this respect, keys are like mazes—if you
arrive at a dead end, you can always turn around and go back!
Another pitfall to be aware of is language. Watch for qualifiers such as and, if
usually, sometimes, generally, typically, when young, and at maturity; i.e., “often”
does not mean “always,” and “typically” means “most often.” Also watch out for the
statement “not as above,” as in the following excerpt from a dichotomous key to the
poems of an obscure but brilliant Santa Cruz poet.
1. Poem with a pointed social comment .2
1. Not as above.. .77
HOW TO USE THE KEYS 23

In this case, “not as above” means “social comment absent, or if present, then not
pointed.” However, if the first statement in the couplet said “social commment
present, often pointed,” then “not as above” would translate as “social comment
absent.”
(3) IT IS THE KEY’S FAULT. If you have followed the key correctly, and are
certain your mushroom(s) is not aberrant, then a discrepancy between the
mushroom and the description means either that the key is flawed (no key is perfect)
or that you have a mushroom that is not included in the keys (no mushroom book is
complete and none should pretend to be—our inventory of mushrooms is still in the
preliminary stages!). You may then either consult another book in hopes of finding
it, seek the counsel of a mushroom expert, discard it altogether, or give it an informal
name of your own.
In addition to aiding in identification, keys can be effective teaching devices. In
keying out a mushroom (instead of being told what it is or leafing through a bunch of
pictures), you are forced to judge critically and develop an eye for detail. By the time
you’ve keyed out a mushroom, you will have accumulated a considerable amount of
information on what it is, as well as what it is not. You learn things about it that
might otherwise go unnoticed. For instance, in keying out a banana slug using the
sample key, you must try to peel it—something you ordinarily would not do. You
would then learn that a banana slug does not peel easily. Another challenging
activity is to try devising keys yourself—to common mushrooms, household objects,
or people you know. You’ll discover that it’s not as easy as it looks!

QUESTIONS ABOUT MUSHROOMS

WHICH MUSHROOMS ARE GOOD TO EAT?

Fortunately, there is no easy answer to this question. If there was, everybody

would pick mushrooms and there wouldn’t be enough to go around!
The only way to determine the edibility of a mushroom is to eat it.
OK.
But who wants to risk their life for the sake of one lousy (or marvelous) meal?
Well, through just such a risky trial-and-error process we humans have
painstakingly accumulated a reliable body of information on the edibility of
many mushrooms. By systematically learning the identifying characteristics of
these mushrooms, you can take advantage of other people’s experiences—many
of them unfortunate—by eating only the safest and most savory ones while
avoiding the poisonous ones.
If possible, seek out the help of an experienced and knowledgeable mushroom
hunter.* There’s a subtle gap between the mushroom in the book and the
mushroom in the bush, and she or he can help you bridge that gap. If you don’t
have a mushroom mentor, proceed with caution. Just because you identify a
mushroom as an edible species doesn’t mean you should eat it. It is better to
collect it several times first, so that you become thoroughly familiar with it.
When you misidentify a rock or lizard, it doesn’t really matter, because you’re
not (hopefully) going to eat the thing. With mushrooms, it’s different. The

* Mushroom hunters can be experienced without being knowledgeable. A knowledgeable hunter knows the names
of the mushrooms she eats as well as those of all poisonous look-alikes.
Even relatively sophisticated devices such as this “Toadstool Tester” are unreliable. There is no
substitute for caution, experience, and a fundamental familiarity with the fleshy fungi.

cardinal rule is: Don’t eat any mushroom unless you are absolutely sure of its
identity! Or—WHEN IN DOUBT, THROW IT OUT! In other words, you’re
better off not eating an edible mushroom than eating a poisonous one, and it’s
more important not to eat a poisonous mushroom than to eat an edible one.
Empirical approaches such as the “silver coin” test are without exception pure
poppycock. The lethal Amanitas do not blacken silver (fortunately, silver coins
are a rarity nowadays, so this fallacious test is more difficult to carry out).
Mushrooms partially eaten by mammals or insects are not necessarily fit for
human consumption (one animal’s meat is another’s poison). Mushrooms that
smell and taste good are not necessarily edible (again, the deadly Amanitas are
said to be delicious.
Most insidious of all is the “intuitive” approach now in vogue. Intuition can
play a part in finding mushrooms, but not in determining their edibility. It is a
sad comment on our times that intuition is so often peddled as a substitute for
critical observation. If our intuitions were infallible insofar as mushrooms are
concerned, there would be no need for books on the subject. It takes
commitment and a good deal of conscious effort to identify mushrooms
correctly. The “intuitive” approach is appealing, I suspect, precisely because it
promises a maximum amount of satisfaction for a minimum amount of effort.

24
 25

WHAT’S THE DIFFERENCE BETWEEN A MUSHROOM

AND A TOADSTOOL?

“Toadstool” usually carries the connotation of being poisonous, but since

many people think that all wild mushrooms are poisonous, the two terms are
virtually interchangeable in popular usage. The word “toadstool” is perhaps a
testament to the old folk belief that toads gave warts to people who handled
them, and made mushrooms poisonous by sitting on them.

WHEN AND WHERE DO MUSHROOMS GROW?

Learning to recognize a mushroom should not be an end in itself, but a means
of getting to know the fungus. A field guide can teach you the physical features of
the fruiting body (the mushroom aspect of the organism), but you must discover
for yourself the traits of the fungal aspect—that is, its fruiting behavior and
ecological idiosyncrasies. This fungal aspect can only be appreciated by
spending a good deal of time in the woods and fields, and yet it is this aspect that
is ultimately the most rewarding—getting to know organisms you can’t see most
of the time on an intimate basis.
Fungally speaking, we are smiled upon most favorably. Our mild coastal
climate allows for a long mushrom season (late October through March), and at
least a few species can be found any month of the year (Agaricus augustus, for
instance, fruits prolifically during the dry summer months). However, the peak
months for most mushrooms are the wettest ones—November through
February. As you move north along the west coast, the season is progressively
earlier (September through November or December) and more compressed; as
you go south it is correspondingly later and more erratic. In the Rocky
Mountains, on the other hand, the best time is usually August, while in eastern
North America it is the summer and fall and in the Deep South it is the summer
and winter or even year-round.
Just as some wildflowers bloom in March and others in July, so each kind of
mushroom has its own biological clock. In our area the sulfur shelf (Laetiporus
sulphureus) fruits on eucaylptus stumps in September and October (before the
fall rains arrive), while the horn of plenty (Craterellus cornucopioides) seldom
shows up before Christmas, morels (Morchella species) appear in the spring, and
the blewit (Clitocybe nuda) fruits continuously throughout the mushroom
season.
Since rainfall and temperature are major determining factors, no two
mushroom seasons are quite alike. There are good mushroom years and bad
mushroom years, but most typical is a year which favors the fruiting of some
types at the expense of others. For instance, the warm winter rains of 1977-78
produced a bumper crop of Agaricus in our area, but Chroogomphus was
practically absent. The previous year, however, there was a preponderance of
Chroogomphus and only a smattering of Agaricus. It can be seen, then, that the
terms “common” and “rare” can be misleading. “Common” means “often
common,” while “rare” actually means “rarely common,” because some species
may be absent for many years and then, at the beckoning of some mysterious
signal, fruit suddenly in unprecedented quantity. As a result, you can search the
same locality for years and still find new species!
26 WHEN AND WHERE DO MUSHROOMS GROW?

To the beginner practically every mushroom is “rare.” Experienced hunters

usually find more mushrooms because they know exactly where to go. They
stake out secret mycelial “patches” of their favorite fungi which they visit
regularly and guard zealously. So unless it’s the peak of the season, you’re likely
to find little in the way of collectable delectables unless you know exactly what
you are looking for and where it is likely to grow. For instance, you don't go
looking for manzanita boletes (Leccinum manzanitae) under pine. If you do,
you’ll cpme home empty-handed, although without realizing it you may pass up
some delicious hedgehog mushrooms (Dentinum repandum), especially if the
area you search is overgrown with brambles and poison oak. The more you
search, the more “patches” you discover and the more you develop an intuitive
feel for when and where mushrooms grow. You learn to hunt as much with your
nose and fingers and belly as with your eyes. Eventually you can walk through
the woods when there are no mushrooms out, and successfully predict which
kinds will come up when it’s wetter.
And yet, a large part of mushroom hunting is timing, and a large part of timing
is luck. So much depends on where you happen to be at what time. If you’re too
early, the mushrooms will be invisible, still under the mulch. If you’re too late,
they’ll be “occupied” (riddled with maggots), already harvested, or too old to eat.
Make as many generalizations and specifications as you like, and mushrooms
will still defy them!
This element of uncertainty, though at times the cause of considerable
frustration, is also the reason for much of the excitement. No matter how
experienced and knowledgeable you are, you can never really be sure of what
you’ll find—you can only increase to some degree your chances of success. There
is the acute despair of not finding what you hoped for, the frenzied delight and
disbelief at finding more than you dreamed possible, the sure and comfortable
satisfaction of finding exactly what you “knew” would be there.

CAN PEOPLE HARM MUSHROOMS BY PICKING THEM?

Since mushrooms are the “fruit” of a fungus, picking them is like picking
apples or blackberries or figs—no harm is done providing you gather them
carefully and do not unduly disturb the environment. Nor does selective picking
interfere seriously with their ability to reproduce. Most mushrooms you find
have already shed spores (unlike flowers, which have yet to form seeds), and will
continue to shed them after they’re picked. Also, the parent mycelium is usually
perennial and will produce mushrooms periodically, and there is some evidence
to suggest that removing a few mushrooms can stimulate the mycelium to
produce more.
The last several years, however, have seen a tremendous increase in the amount
of wild mushrooms being harvested commercially—particularly in northern
California and the Pacific Northwest. These mushrooms are shipped to markets
and restaurants all over the country as well as abroad. As commercial picking is
only lucrative when large numbers of mushrooms are gathered, animosity is
developing between those who pick for money and those who pick for pleasure.
The commercial pickers and their distributors claim that commercial
harvesting provides valuable jobs in depressed areas (though the actual
CAN PEOPLE HARM MUSHROOMS BY PICKING THEM? 27

number is fairly small), and they augment this claim with the specious argument
that they are providing a service to the public, i.e., making wild mushrooms
available to a larger audience, or at least to more than just a select few. (The
“select few”, in this case, is almost any ambulatory person who wants to learn
about mushrooms; the “larger audience” is anyone who has extra money with
which to buy wild mushrooms, which are never cheap!)
The “select few,” on the other hand, profess concern about over-picking and
the possibility that certain edible species will be driven to extinction. However,
there is no hard evidence to suggest that the latter is happening—or about to
happen. I suspect what is really upsetting these mushroom hunters is that
their habitual stomping grounds are being ravaged by commercial exploi¬
tation. This is a perfectly legitimate concern, if you ask me, for there is
hard evidence that some—perhaps only a few—commercial pickers have not
only “raided” but completely “cleaned out” areas where mushroom hunters with
less time or mobility have been foraging for years. There ought to be enough
mushrooms out there to support both commercial and personal picking, but
mushroom hunting is one of many activities where a single selfish, inconsiderate
boor can spoil it for everyone else. So, you selfish inconsiderate boors—you
know who you are! (Or if you don’t, we do!)
Returning, then, to the original question—mushrooms are a renewable
resource, and the only sure way to eradicate a species is to eliminate its habitat
(which, unfortunately, is becoming a common practice as wild areas succumb to
pollution and the appetite of developers). So don’t feel guilty about picking
mushrooms, but don’t, on the other hand, pick more than you can use (COLOR
PLATES 79, 81). Respect the environment and be considerate of those who
follow in your footsteps—you may be one of them!

CAN PEOPLE HARM THEMSELVES

BY PICKING MUSHROOMS?
No. Handling wild mushrooms is not dangerous. You may want to wash your

Poison oak (left) and poison ivy are the banes of many a would-be mushroom hunter. The leaves grow
in threes, but are shed during the winter! If you’re allergic to them, shower thoroughly afterevery hunt
and put your clothes in the wash. Ticks (right) and boletivores (see p. 546) also cause problems.
Rattlesnakes are known to lurk near or under mushrooms, especially large boletes, and they do not
always take kindly to being disturbed. (Maybe that’s why boletivores carry long sticks!)

hands after prolonged contact with deadly Amanitas, but poisonous mush¬
rooms can cause harm only if ingested. There are, however, some corollary
dangers to mushroom hunting—see photos above and on previous page!

DO OTHER ANIMALS EAT MUSHROOMS?

Yes. As any mushroom hunter will tell you, many a marvelous meaty
mushroom has been reduced in a matter of hours to a writhing mass of beatific
maggots. These wiggly white “worms” with the black heads are the larvae of
fungus gnats (Mycetophiladae). Eggs are usually laid at the base of the stalk, and
the newly hatched maggots work their way up to the cap, gorging themselves

One of the unsung rewards of mushroom hunting is the discovery of a host of other fantastic clan¬
destine creatures. Some of these, like slugs, love to munch on mushrooms. Others, like mantises and
millipedes, use them for shelter or hiding places. Left: A banana slug grazing peacefully in a “fungal
jungle” of oyster mushrooms. Right: A praying mantis, up close and personal.
Close-up of a banana slug grazing on an oyster mushroom. Banana slugs can also be seen in Color
Plates 57 & 152, and at the bottom of p. 28.

along the way. Many other insects feed on fungi, including springtails and
various beetles, and flies like nothing better than to feast on stinkhorn slime.
Sowbugs seem to be very fond of Agaricus augustus. Slugs gormandize
mushrooms whenever possible (COLOR PLATES 57 & 152) and have special
“fungus-detectors” to aid them in the search. Tortoises have been known to
interrupt their imperturbable peregrinations to munch on a mushroom or two.
Rodents—particularly squirrels and chipmunks—ae confirmed fungophiles.
They’re fanatically fond of various boletes and agarics and those odoriferous
underground fungi called truffles and false truffles. (A study by Joseph Hall of
San Francisco State U niversity revealed that the summertime diet of the Kaibab
squirrel of Arizona consists almost entirely of fungi; during the summer captive
Kaibab squirrels rejected other food when mushrooms were made available.)
Pigs have a passion for true truffles. In Europe they are used to hunt them
down, but have to be muzzled so they don’t devour them (see p. 842)! Even the
ruminants—cattle, deer, etc.—occasionally indulge. I have seen cows grazing on
giant puffballs, and Siberian reindeer are addicted to Amanita muscaria (like
humans, they experience profound mental disturbances after eating it). They are
also said to be extremely fond of human urine, and is it coincidence that the
Siberians who ate A. muscaria also made a practice of drinking their own urine
in order to recycle the intoxicants? R. G. Wasson, in SOMA: The Divine
Mushroom of Immortality, quote Vladimir Jochelson, a Russian
anthropologist:
The reindeer have a keen sense of hearing and smell, but their sight is rather
poor. A man stopping to urinate in the open attracts reindeer from afar,
which, following the sense of smell, will run to the urine, hardly discerning
the man, and paying no attention to him. The position of a man standing up
in the open while urinating is rather critical when he becomes the object of
attention from reindeer coming down on him from all sides at full speed.

Some mushrooms, incidentally, are carnivorous! Cordyceps species feed on

insects, and it has recently been shown that several wood-rotting fungi (in¬
cluding the oyster mushroom) supplement their diet by digesting nematodes.

29
30

WHAT IS THE NUTRITIONAL VALUE OF MUSHROOMS?

Mushrooms are esteemed primarily for their flavor, but they can also be a
healthy supplement to your diet. Each type, of course, has a different chemical
composition, but in general their nutritive value compares favorably to that of
most vegetables. They are rich in the B vitamins (including choline, which acts as
a protective agent for your liver in case of mushroom poisoning), vitamin D, and
vitamin K. Some are also high in vitamin A (e.g, the chanterelle, Cantharellus
cibarius), and a few (e.g., Fistulina hepatica) contain vitamin C. Mushrooms are
also rich in minerals such as iron and copper and various trace elements.
Like fruits, vegetables, and human beings, mushrooms are mostly water (85-
95%). They have a low fat and carbohydrate content, and as a result, almost no
calories—unless, of course, they are cooked in oil or butter. Some types are high
in protein (especially Agaricus, Lepiota, and Calvatia species), and on a dry
weight basis Boletus edulis contains more protein than any common vegetable
except soybeans. However, some of this protein is indigestible, so mushrooms
are not a viable substitute for meat or other high-protein foods. Cooking mush¬
rooms maximizes their nutritive value by increasing their digestibility. Over¬
cooking them, however, removes some of their vitamins and most of the flavor.

WHAT IS THE MEDICINAL VALUE OF MUSHROOMS?

Fungi contain many unique and powerful substances—penicillin, for instance,
which was accidentally discovered in the bread mold Penicillium. Many mush¬
rooms contain antibiotic substances, particularly those that don’t decay quickly.
The Chinese prize several polypores as much as ginseng (e.g., Ganoderma
lucidum, Grifola umbellata) because of their alleged beneficial effects on health.
There is some evidence to suggest they may strengthen or stimulate the immune
system. The Japanese attribute similar properties to the shiitake (Lentinus
edodes). More research should be conducted to determine what healing proper¬
ties, if any, these and other fungi possess. It is a tribute to the general feeling of
our society that mushrooms are harmful at worst and worthless at best that very
little research of this kind is being done in this country.

CAN YOU GROW WILD MUSHROOMS?

Yes, but just how is beyond the scope of this book. Many wild mushrooms have
been successfully cultivated on a commercial basis—particularly in Japan, a
mushroom-loving country with relatively little wilderness in which to hunt them.
Some of these mushrooms are now being grown in the United States, e.g., the
shiitake (Lentinus edodes), enokitake (Flammulina velutipes), and oyster mush¬
room (Pleurotus ostreatus). In addition to the familiar cultivated mushroom
(Agaricus bisporus), so-called “magic mushrooms” such as Psilocybe cubensis
are also grown, albeit illicitly. Several species are better suited to homescale than
commercial production, including Clitocybe nuda, Coprinus comatus, Agaricus
subrufescens, Hericium spp., and Stropharia rugoso-annulata. Still others, such
as Boletus edulis, Cantharellus cibarius, and Morchella species, stubbornly
resist attempts to raise them. As a rule, mycorrhizal fungi are difficult to raise
because of their nutritional requirements (tree rootlets), while those that grow on
wood, compost, dung, or disturbed ground are relatively easy. For more infor¬
mation on how to grow mushrooms, see the bibliography.
Human beings, incidentally, are not the only ones to cultivate fungi, and were
Left: The shiitake, Lentinus edodes, was probably the first mushroom cultivated by humans. A native
of Japan, it is now grown in the United Statesand may establish itself in the wild. This cluster is fruiting
from a shiitake kit (a “log” of compressed sawdust). Right: Morels are among the many delicious
mushrooms that resist commercial cultivation. These are red morels (Morchellaelata group, p.790).

by no means the first. That honor belongs to certain ants and termites who raise
fungi in their nests. On almost any walk through a tropical rain forest you can
see long files of leaf-cutting ants carrying bits of leaves to their nests. These leaves
are not food for the ants, but food for fungi upon which the ants feed!

HEY, M A AA AA AAA A N, DO ANY PSILOCYBIN MUSHROOMS

GROW AROUND HERE?
I am asked this question more often than any other, and I must confess it
irritates me—not because I object to the use of hallucinogenic drugsper se—but
because of the attitude that usually—but not necessarily—accompanies their
use. Most of the people who ask this question would rather change their way of
looking at reality than face the difficult and discouraging task of transforming
reality itself. Hence they see mushrooms as means to their own ends. They go out
to the pastures and stuff their plastic bags with all the “LBM’s” (Little Brown
Mushrooms) they can find, then either pop the whole rotten mess into their
mouths (a dangerous practice!), or expect someone like me—whose time is
presumably less valuable than theirs—to sort them out. While clinging to the
moronic belief that they constitute a “counterculture,” they share our society’s
overriding urge for expediency. They make no attempt to learn about the
organisms they eat and it has always struck me as ironic that people with such a
low level of consciousness should be seeking “higher consciousness.”
Excerpts from Good Times (January, 1971) epitomize their line of thinking:
“Amanita Reality”; “us genetic revolutionaries the longhairs”; “circumcision of
the second charka”; “magic mushroom generation”; “I could relate the black
hole notion to the notion of total pollution in that either experience would seem
kind of freaky”; and “Amanita muscaria is total revolt, and the revolution is just
about won.” From this mindless drivel it is clear why Marge Piercy laments:
We grew up in Disneyland with ads for friends
and believed we could be made new by taking a pill.
We wanted instant revolution, where all we had to add
was a little smoke.
But there is no tribe who dance and then sit down
and wait for the crops to harvest themselves
and supper to roll over before the pot . . .
32 HEY MAAAAAAAAAN . . .

To answer the original question—yes, there are hallucinogenic mushrooms in

California and the rest of North America, and most of them contain psilocybin
and/or psilocin. But if you’re just looking for a new high, you really ought to
take up hang-gliding or bottle-throttling, or take your chances with what you get
on the street. If, on the other hand, you have a genuine interest in our co¬
inhabitants on this planet, and you wish to explore altered states of
consciousness, then the safest approach is to buy a responsible field guide to
hallucinogenic mushrooms and use it in conjunction with this book. Possession
of psilocybin and psilocin, incidentally, is prohibited by federal law (the
Comprehensive Drug Abuse and Control Act).

LBM’S: LITTLE BROWN MUSHROOMS

THE CAP is brown, the stem a shade browner, the gills browner still. This can be
said of nearly one half of all the mushrooms you find. On even the most casual
jaunt through the woods, you’ll find dozens and dozens of Little Brown Mush¬
rooms (COLOR PLATE 107) sprouting at your feet, and very likely under them
as well. The fact is, Little Brown Mushrooms (“LBM’s”) are so overwhelmingly
abundant and uncompromisingly undistinguished that it is more than just futile
for the beginner to attempt to identify them—it is downright foolish.
After spending a good 25% of my waking existence being downright foolish, I
have come to the painful but inescapable conclusion that the only possible
reason for there being more than one kind of Little Brown Mushroom is that
their “creator” has an inexplicable fondness for prospective professionals in
search of a profession, i.e., Little Brown Mushrooms provide an ever-expanding
plethora of pleasant possibilities for lengthy treatises with intriguing and
titillating titles such as, “A Preliminary Contribution toward a Partial
Monograph of the Section Ignobiles of Subgenus Obfustucantes, Genus
Immobilaria as it Occurs in Outer New Bunswick,” or “More Useless and
Uninteresting Agarics from Putrescent Point State Park.”
I wouldn’t begrudge this in the least were it not for the fact that this same
“creator” is unequivocally cruel when it comes to rank amateurs such as you and
I, who are not paid to peer down the narrow barrel of a microscope until we are
bug-eyed or get scholar’s thumb from flipping through all those worn-out stacks
of abstruse Ph.D. theses—and whose curiosity must consequently be swallowed
rather than satiated as we slam our big red mushroom book against the wall and
decapitate that unpretentious “LBJ” (Little Brown Job) that didn’t asked to be
picked and “demystified” in the first place, but which we went ahead and picked
anyway, and have ever since been attempting in vain to identify.
Though each new Little Brown Mushroom you find will bear a striking
resemblance to the last Little Brown Mushroom you threw away, don’t let this
deceive you into assuming they are identical. Par from it! Somehow, each one
finds a new and more minute way to be different, whether it be that the
incrustation of the pileal epidermis is cheeriose rather than pretzeloid, or that the
hymenial pleurocystidia contain mysterious particles with a refractive content
when mounted in a 10% aqueous solution of Pepto-Bismol. It is almost as if they
were deliberately challenging the taxonomist, who must tax his or her creative
powers to their utmost in order to uncover the differences.
An “LBM” (Little Brown Mushroom), also known as an “LBJ” (Little Brown Job).,

Actually, thanks to the diligence and expertise of the professionals, we are

slowly accumulating a large mass of knowledge on the Little Brown
Mushrooms. Not a completely coherent mass, mind you (in some ways it still
resembles a mess more than a mass), but six hours’ painstaking perusal of the
current literature will produce the name of at least one out of 50 of the featureless
little fellows instead of one out of 51, as it used to be. Part of the problem, I
suspect, is that new species are being designed and disseminated at
approximately the same rate that old ones are being defined and differentiated,
so that we will never attain the level of taxonomic command that we have of, say,
the Little Brown Lizards or the Big Brown Toads.
Incidentally, to qualify as a bonafide Little Brown Mushroom, you ought to
be little, and you most definitely have to be a mushroom, but you don’t
necessarily have to be brown, though it certainly simplifies things for you (by
making it more difficult for us) if you are. That is to say, there are any number of
boring buff Little Brown Mushrooms, wishy-washy white Little Brown
Mushrooms, and gratuitous gray Little Brown Mushrooms (to say nothing of
those whose color is so neutral or nebulous as to defy description), but there are
no breathtakingly blue Little Brown Mushrooms, and by and large Little Brown
Mushrooms are simply and unequivocally brown.
The point I’m trying to make, folks, is that it’s sheer folly to embark upon the
purchase of a field guide with the expectation of identifying each and every
fleshy fungal fructification you find. Please, for your sake, don’t expect the
“pegs” to fit neatly into the “holes.” The “holes”are made by human beings and
the “pegs” are not! Some “pegs” will fit a number of holes, some won’t fit
anywhere and must wait for a hole to be gouged out. A few will fit snugly into
one hole and one hole only, and these are the ones you should get to know. By
concentrating on these larger and/or more distinctive types (see the chart on
pp. 48-51), and proceeding at a measured, deliberate pace, you will experience a
mounting satisfaction at your ability to demystify, identify, appreciate—and
hopefully eat—some of the mushrooms you gather. Remember, what we know
of mushrooms (or anything, for that matter) is substantially more than what we
knew fifty years ago, but is still precious little compared to what we don’t know.
The bulk of this book deals with the larger, more easily recognized fleshy
fungi. However, in the interests of providing a broad overview, some of the more
noteworthy (or rather, less unnoteworthy) “LBM’s” have been included. Almost
every genus boasts a few, but the majority belong to Inocybe, Tubaria, Galerina,
Collybia, Psathyrella, Marasmius, Mycena, Pholiota, and Cortinarius.

33
An oak woodland, with clusters of honey mushrooms (Armiliariella mellea) in the foreground.
(Nancy Burnett)

HABITATS
DEVELOPING an awareness of biological communities is essential to any
nature study. In this chapter, some of the more common or distinctive
mushrooms are grouped according to habitat—the type of place where they’re
most likely to be found, or the type of tree with which they commonly grow. The
environmental aspect of collecting is frequently neglected by mushroom
hunters, who eagerly remove mushrooms from their place of growth without
taking note of their surroundings. In many instances the place of growth will
provide clues to the identity of the mushroom—and you’ll soon discover that
certain mushrooms always seem to grow together; that is, they are indicators of a
certain habitat or biological community.
Most of the habitats discussed here are named according to dominant
vegetation (oak, redwood, grass, etc.). Since only a few of our local trees are
significant in mushroom identification, it makes a lot more sense to learn those
trees before you learn the mushrooms, rather than trying to differentiate
between several hundred kinds of mushrooms before learning the basic trees.
Always bear in mind that the different habitats discussed here overlap, so that
in a given area there are usually several habitats present. For instance, a lawn with
a Monterey pine will feature grass-loving fungi as well as species mycorrhizal with
pine. And a road through the woods will offer many types of mycorrhizal or
humus-inhabiting mushrooms plus those that grow in disturbed ground (the
roadside). Be sure to check the list of cosmopolitan mushrooms (“Anywhere and
Everywhere”), for these will turn up in almost any habitat. Also remember that
mycorrhizal fungi do not necessarily grow under their host—they are associated
with the tree’s rootlets, which may be quite a distance away from the trunk. The
habitats described in the following pages include several widespread ones (e.g.,
lawns or disturbed ground) and several specific to our area (e.g., cypress and
redwood). Space does not permit an exhaustive discussion of forest trees outside
our area, but a few are briefly dealt with under the heading “Other Trees.”
34
 35

PINE (Pinus species)

Pine forests are favored by a melange of mycorrhizal mushrooms, plus multitudes of
minute, saprophytic Mycenas. When the needle carpet is dry (“potato chip conditions”*),
the larger mushrooms often hide underneath, manifesting themselves as low mounds or
“mushrumps.” Prized edible species include Boletus edulis, Dentinum repandum, Clito-
cybe nuda, Sparassis crispa, and Tricholoma flavovirens. Suillus and Chroogomphus
species often fruit together in enormous quantities, along with the green and orange
Lactarius deliciosus and the bright red or pink Russula rosacea. But the most spectacular
fungal feature of our pine forests is unquestionably Amanita muscaria, with its fiery red
cap studded with white “stars.”
Of course, from region to region the different kinds of pines feature different mycor¬
rhizal partners, and even in the same area there are usually some differences in the fungal
associates of each pine species. For instance, Suilluspseudobrevipes and Chroogomphus
pseudovinicolor grow with ponderosa pine in our area, Suillus pungens with Monterey
and knobcone pine, and Suillus fuscotomentosus with knobcone and ponderosa pine,
while Hygrophorus gliocyclus is abundant inland under digger pine, but also occurs
along the coast with Monterey pine, and Amanita caesarea and Lactarius indigo occur
with ponderosa pine in Arizona but are absent from California’s ponderosa pine forests.
Agaricus subrutilescens Cortinarius species Mycena species
Albatrellus flettii Cryptoporus volvatus Marasmius sp. (unidentified)
Amanita aspera Dentinum species Mycena species
Amanita caesarea Endoptychum depressum Naematoloma species
Amanita muscaria Elaphomyces species Phaeolus schweinitzii
Amanita pachycolea Gymnopilus sapineus Phellinus pini
Amanita pantherina Gymnopilus spectabilis group Pholiota species
Armiliaria ponderosa Hebeloma crustuliniforme Pluteus atromarginatus
Boletus barrowsii Hebeloma sinapizans group Rhizopogon species
Boletus edulis Helvella lacunosa Russula alutacea group
Boletus piperatus Hydnellum species Russula brevipes
Brauniellula nancyae Hygrophoropsis aurantiaca Russula emetica group
Callistosporium luteo-olivaceum Hygrophorus erubescens Russula rosacea
Chroogomphus species Hygrophorus gliocyclus Russula sororia group
Clavulina cristata group Hygrophorus hypothejus Sparassis crispa
Clitocybe deceptiva Hygrophorus purpurascens Suillus species
Clitocybe nuda Inocybe species Tricholoma flavovirens
Clitocybe sclerotoidea Inonotus tomentosus Tricholoma imbricatum
Clitocybe species Laccaria species Tricholoma pessundatum group
Collybia butyracea Lactarius deliciosus Tricholoma squarrulosum
Collybia dryophila Lactarius indigo Tricholoma terreum group
Cortinarius cinnamomeus group Lactarius rufus Tricholoma vaccinum
Cortinarius mucosus Lactarius subflammeus Tricholoma zelleri
Cortinarius obtusus group Lactarius vinaceorufescens Tricholomopsis rutilans
Cortinarius phoeniceus Lentinus ponder osus

DOUGLAS-FIR (Pseudotsuga species)

There are well over 1,000 kinds of mushrooms known to form mycorrhiza with Douglas-
fir, and the great Douglas-fir forests of the Pacific Northwest are among the best fungal
foraging grounds in the world. In our region, however, the Douglas-fir habitat is no
better than average. The most prominent mycorrhizal associates are Suillus, Gomphidius,
Lactarius, and Russula species. Russula xerampelina is among our few choice edibles
that grow principally with Douglas-fir. Laetiporus sulphureus is occasionally found on
logs and stumps, and Cantharellus cibarius and C. subalbidus are abundant under
Douglas-fir farther north. Tuber gibbosum, when you can find it, is also excellent.
*Potato chip conditions: when it’s so dry that it sounds like you’re walking on a bed of potato chips; equally unde¬
sirable are “tidepool conditions,” when the humus layer is so soggy it sounds like you’re traipsing across a bed of
sea anemones (squish squish squish).
36 DOUGLAS-FIR

The name “Douglas-fir” is hyphenated in this book because it is not really a fir. It is
easily told from redwood (with which it often occurs locally) by its browner bark, spurred
cones, and habit of dropping needles without twigs intact (redwood drops twigs with
the reddish needles intact).

Amanita pant her ina Lactarius vinaceorufescens Russulaplacita group

Cantharellus species Lepiota clypeolaria Russula xerampelina
Cortinarius species Mycena species Strobilurus trullisatus
Fomitopsis cajanderi Otidea species Suillus caerulescens
Fomitopsis pinicola Phaeolus schweinitzii Suillus lakei
Gomphidius species Phellinus pini Suillus ponderosus
Hygrophorus agathosmus Rhizopogon species Tricholoma terreum group
Inocybe species Russula gracilis group Truncocolumella citrina
Lactarius rubrilacteus Russula Integra group Tuber gibbosum

REDWOOD (Sequoia sempervirens)

Ironically, our largest tree supports a fungal phantasmagoria of dainty, fragile fungi,
but only a smattering of the large, fleshy types. Relatively few redwood-lovers are wood-
rotters, and even fewer (if any) are mycorrhizal—a tribute to the redwood’s unique
position among the conifers.
Caulorhiza (-Collybia) umbonata, with its conical or umbonate cap and long “tap
root,” is perhaps the most distinctive redwood-lover. Colorful waxy caps (Hygrocybe
species) and blue-black to purple-black Leptonias abound, but pickings for the table are
meager, at least south of San Francisco—Agaricus augustus (but usually near roads or
trails), Agaricus subrutilescens, Clitocybe deceptiva, and sometimes Cantharellus
cibarius. The largest inhabitant of our redwood forests is Leucopaxillus albissimus,
which often forms impressive fairy rings, but unfortunately, is not a good edible.

Agaricus augustus Entoloma madidum Mycena species

Agaricus hondensis Geoglossum species Nolanea species
Agaricus praeclaresquamosus Helvella species Polyporus hirtus
Agaricus subrutilescens Hygrocybe species Ramaria abietina
Boletus zelleri Hygrophoropsis aurantiaca Ramaria myceliosa
Camarophyllus species Lepiota species Ramariopsis kunzei
Cantharellus cibarius Leptonia species Stropharia ambigua
Caulorhiza umbonata Leucopaxillus albissimus Trichoglossum hirsutum
Clavaria vermicularis Macrotyphula juncea Tricholomopsis rutilans
Clitocybe species Microglossum viride Xeromphalina species

CYPRESS (Cupressus species)

Our cypress groves do not qualify as “woods” because nearly all of them were planted.
They have few (if any) mycorrhizal associates, but there are a number of fleshy fungi that
grow mainly under cypress. The harsh winds and salt spray to which coastal cypresses are
subjected make for unpredictable hunting, but under favorable conditions there is a large
burst of Lepiota and Agaricus species, many of them unclassified and/or endemic to
California. From a gastronomic standpoint the best cypress-lovers are Agaricus bisporus,
A. lilaceps, A. bernardii, Lepiota rachodes, and Clitocybe nuda. In southern Arizona,
incidentally, there are large forests of Arizona cypress, and it would be very interesting to
compare the fungal flora of those cypress tracts to that of coastal California.

Agaricus benesi Agaricus lilaceps Clitocybe nuda

Agaricus bernardii Agaricus pattersonae Geastrum species
Agaricus bisporus Agaricus perobscurus Hygrocybe species
Agaricus blandianus Agaricus species Lepiota rachodes
Agaricus californicus Agaricus xanthodermus Lepiota species
Agaricus fuscofibrillosus Battarrea phalloides Tyromyces basilaris
Agaricus fuscovelatus Camarophyllus species
 37

OAK (Quercus species)

Oaks are endowed with a rich array of fleshy fungi that differ drastically from the
conifer-lovers typical of northern California and the Pacific Northwest. In fact, oaks
appear to have more mycorrhizal partners than any other angiosperms (hardwoods), and
mushroom lovers are indeed fortunate that they are the dominant forest trees of the
central California coast. Oaks also boast the longest mushroom season of any local forest
type. The major fruiting, naturally, is in the fall and winter, but there is a characteristic
spring crop highlighted by Lactarius fragilis, L. rufulus, Clitocybe nuda, Tuber species,
and a trio of Amanitas— A. ocreata, A. rubescens, and A. velosa. Wood-inhabiting
bracket fungi are prominent year-round, including Trametes versicolor, Lenzites
betulina, Stereum species, and Tremella mesenterica. But to inveterate fungophiles, our
oak woodlands are synonymous with chanterelles and blewits (Cantharellus cibarius and
Clitocybe nuda)—the two not only grow together, they go together (in a variety of
delicious dishes). Other choice edibles include Boletus appendiculatus, B. regius, B.
barrowsii, Crater ellus cornucopioides, Lactarius fragilis, L. rufulus, Pleurotus ostreatus,
Amanita velosa, and Tricholomaportentosum. But beginners be careful—our most dan¬
gerous mushrooms, Amanitaphalloides and A. ocreata, are also associated with oak!
The following list of oak-lovers applies mainly to live oak (Q. agrifolia and close
relatives), the principal species in our area. Other kinds of oaks, mostly deciduous, occur
inland and offer a more modest selection of fleshy fungi, perhaps because they receive less
rainfall, or perhaps because they are deciduous.

Agaricus albolutescens Cortinarius regalis Lenzites betulina

Agaricus californicus Cortinarius sodagnitus group Mycena species
Agaricus hondensis Cortinarius species Omphalotus olivascens
Amanita baccata Craterellus cinereus Otidea species
Amanita constricta Craterellus cornucopioides Phellinus gilvus
Amanita magniverrucata Crepidotus species Phyllotopsis nidulans
Amanita ocreata Daldinia species Pleurotus ostreatus
Amanita pantherina Entoloma species Pluteus cervinus
Amanita phalloides Exidia glandulosa Pluteus lutescens
Amanita rubescens Genea species Polyporus decurrens
Amanita velosa Gyroporus castaneus Psathyrella hydrophila
Armillariella mellea Hebeloma crustuliniforme Psathyrella species
Boletus appendiculatus Hericium erinaceus Russula cyanoxantha
Boletus barrowsii Hericium ramosum Russula fragrantissima group
Boletus dryophilus Hygrophorus albicastaneus Russula maculata group
Boletus erythropus Hygrophorus eburneus Russula subnigricans group
Boletus flaviporus Hygrophorus roseibrunneus Schizophyllum commune
Boletus “marshii” Hygrophorus sordidus Stereum species
Boletus regius Hygrophorus species Trametes versicolor
Boletus satanas Hymenogaster species Tremella foliacea
Bulgaria inquinans Hysterangium species Tremella mesenterica
Cantharellus cibarius Inocybe sororia Tricholoma pessundatum group
Chlorociboria species Inocybe species Tricholoma portentosum
Clavariadelphus pistillaris Lactarius alnicola Tricholoma saponaceum
Clitocybe nebularis Lactarius argillaceifolius Tuber species
Clitocybe nuda Lactarius fragilis Tylopilus indecisus
Collybia dryophila Lactarius rufulus Xylaria hypoxylon
Cortinarius glaucopus group

TANOAK (Lithocarpus densiflorus)

Also known as tanbark oak, this close relative of the oaks forms dense stands with
madrone at higher elevations in the coastal mountains, and also grows in sheltered basins
with redwood. Among the prominent fungi of our tanoak woods are the lovely coral
mushrooms (Ramaria speciesj and the diminutive garlic mushroom (Marasmius
38 TANOAK

copelandi), which is often smelled before it is seen. The best edibles are Agaricus
silvicola, Armillaria ponderosa, Pleurotus ostreatus (on decaying logs), Boletus aereus,
Boletus appendiculatus, Russula cyanoxantha, and Craterellus cornucopioides.

Agaricus hondensis Cortinarius infractus Macrotyphula juncea

Agaricus silvicola group Cortinarius species Marasmius copelandi
Armillaria ponderosa Craterellus cornucopioides Phellinus gilvus
Boletus aereus Entoloma madidum Pleurotus ostreatus
Boletus appendiculatus Entoloma species Polyporus hirtus
Boletopsis subsquamosa Hydnellum caeruleum Ramaria species
Bulgaria inquinans Hydnum fuscoindicum Russula albonigra
Cantharellus subalbidus Hygrophorus russula Russula cyanoxantha
Clavariadelphus pistillaris Lactarius argillaceifolius Tricholoma saponaceum
Cortinarius collinitus group Lactarius subvillosus Tricholoma zelleri
Cortinarius cotoneus group

CHINQUAPIN (Castanopsis chrysophylla)

Also spelled “chinkapin,” this cousin of the chestnuts has chesnut-like burrs that
contain small edible nuts. Most of the local chinquapins are rather runty and ragged-
looking but farther north they attain heights of 75-100 feet. Chinquapins are also closely
allied to tanoak, and as might be expected, support a similar fungus flora, including
Ramaria species and Boletus aereus. Among the wood-inhabitors, three in particular
stand out: the sulfur tuft (Naematoloma fasiculare) and jack-o-lantern mushroom
(Omphalotus olivascens) because of their brilliance, and the beefsteak fungus (Fistulina
hepatica) because of its bizarreness. Boletus aereus is the best edible species.

Boletus aereus Marasmius copelandi Ramaria species

Entoloma species Naematoloma fasciculare Stereum species
Fistulina hepatica Omphalotus olivascens Trametes versicolor

MADRONE and MANZANITA (Arbutus menziesii and

Arctostaphylos species)
Madrone woods are my favorite foraging grounds. They boast their share of
gastronomic delights (e.g.. Amanita calyptrata, Craterellus cornucopioides, Leccinum
manzanitae), but their chief attraction is their beauty. Madrones don’t blot out the sun
like redwoods, and they’re more colorful (though not quite as venerable) as oaks. Every
year they shed their reddish bark in sheets, revealing smooth, yellow, musclebound limbs
beneath. In the summer the rags of stripped bark sizzle audibly in the sunlight as they curl
up like pencil shavings. In the fall, the stripped bark combines with the large leaves and
clusters of bittersweet orange-red berries to form a deep, incomparably rich, reddish-
black humus. In such splendid company, it’s hard to keep your mind on mushrooms!
Manzanitas are essentially miniature madrones. The most that can be said for foraging
in manzanita thickets is that it’s different—you spend the whole time crawling around on
your hands and knees wishing you’d taken up stamp collecting or basket weaving or
stayed home and watched the ball game.
It is interesting to note that a number of mushrooms normally associated with conifers
cross over to madrone (they just can’t resist!) and/or tanoak. These include Amanita
aspera, Amanita muscaria, Armillaria ponderosa, Tricholoma (-Armillaria) zelleri,
Tricholoma aurantium, T. flavovirens, Hygrophorus chrysodon, and H. eburneus.

Amanita aspera Boletus amygdalinus Cortinarius species

Amanita calyptrata Boletus flaviporus Craterellus cornucopioides
Armillaria ponderosa Cantharellus subalbidus Entoloma species
Boletus aereus Cortinarius balteatus Hydnum fuscoindicum
Watching bark peel can be as exciting as watching a bicycle rust, unless it’s madrone bark, shown here.

Hygrophorus chrysodon Omphalotus olivascens Tricholoma flavovirens

Hygrophorus eburneus Phellinus igniarius Tricholoma manzanitae
Leccinum manzanitae Russula cremoricolor Tricholoma saponaceum
Leccinum species Russula emetica group Tylopilus humilus
Lepiota atrodisca Tricholoma aurantium

EUCALYPTUS (Eucalyptus species)

This messy, aggressive intruder was originally brought to this country in the hope that
its noxious fumes would combat malaria. It grows quickly, and its greedy, shallow roots
inhibit the growth of many native plants and mushrooms (Leucopaxillus albissimus,
Coprinus plicatilis, Clitocybe nuda, and various Agaricus species are among the few
humus-inhabitors with the poor sense to tolerate it). However, eucalyptus makes fairly
good firewood and if you chop them down you will not only prevent them from barging
into someone’s bedroom during the next windstorm, but you’re also liable to get a nice
crop of succulent sulfur shelves (Laetiporus sulphureus) sprouting from the cut stumps!

Agaricus californicus Hymenogaster albus Leucopaxillus albissimus

Agaricus xanthodermus Hysterangium fuscum Marasmiellus candidus
Clitocybe nuda Laccaria species Marasmius plicatulus
Coprinus species Laetiporus sulphureus Psathyrella species
Hydnangium carneum Lepiota rubrotincta Setchelliogaster tenuipes

RIPARIAN WOODLAND
Streams and rivers being indisputably damp, many people assume they constitute an
ideal setting for mushrooms. The mixed hardwoods (alder, cottonwood, willow, maple,
etc.) of our stream valleys and ravines do indeed support a characteristic fungus flora, but
it is a surprisingly modest one that does not yield the bountiful harvests of our oak and
pine forests. Log-rotters abound (Pholiota, Psathyrella, Naematoloma, Armillariella,
Pleurotus, etc.), while the best edibles are Armillariella mellea and Pleurotus ostreatus in
the fall and winter, Verpa and Morchella species in the spring.
Armillariella mellea Marasmiellus candidus Pluteus species
Clathrus archeri Morchella species Psathyrella species
Clavaria vermicularis Mycena species Sarcoscypha coccinea
Coprinus micaceus Naematoloma species Stropharia ambigua
Disciotis venosa Pholiota species Verpa bohemica
Flammulina velutipes Pleurotus ostreatus Verpa conica

39
40 HABITATS

OTHER TREES
Only a few of our native trees have been discussed so far, but the remaining types are
not particularly significant from a mushroom identification standpoint. That is, the
mushrooms they harbor (either beneath or on them) are likely to be cosmopolitan (see the
“Anywhere and Everywhere” category). For instance, almost nothing fungal grows
under bay laurel, but the Ganoderma applanatum group (especially G. brownii) is
abundant on bay laurel as well as many other trees. Similarly, species of Laccaria
(omnipresent mushroom “weeds”) occur in droves under acacia, while the bush lupines
along the coast harbor nice crops of Flammulina velutipes and a brown-capped form of
Pleurotus ostreatus (both edible), and huckleberry is home to Cortinarius balteatus and
the C. cylindripes group.
A few trees of special interest to mushroom hunters that do not occur locally are
discussed briefly below, but space does not permit a detailed listing of their fungal
associates.

LARCH (Larix)—Larches, or tamaracks as they are sometimes called, are deciduous

northern conifers whose needles turn gold before dropping off. They feature a number
of unique mycorrhizal fungi, particularly boletes, plus the usual array of wood-rotters
and saprophytes. The following species favor larch over other conifers.

Gomphidius maculatus Fomitopsis officinalis Suillus cavipes

Hygrophorus speciosus Fuscoboletinus species Suillus grevillei

HEMLOCK (Tsuga)—Hemlocks are a common feature of the northern and montane

forests of North America. In the West they usually mingle with other conifers, and so
do their mycorrhizal mates. The following list includes only those species that seem
overly fond of hemlock. Among them is Boletus mirabilis, a choice edible.

Boletus mirabilis Ganoderma tsugae Ramaria species

Chroogomphus tomentosus Hysterangium separabile Rhizopogon rubeseens
Cortinarius subfoetidus Lactarius subpurpureus Russula xerampelina
Cortinarius species Pleurotus porrigens Tylopilus species
Elaphomyces granulatus

SPRUCE (Picea)—The Sitka spruce stands of coastal northern California and the
Pacific Northwest and the spruce forests of the North, Southwest, and Rocky Moun¬
tains are phenomenally rich in mycorrhizal mushrooms, particularly species of Russula
and Cortinarius and the delectable Boletus edulis. A sizable number of spruce addicts
have already been listed under pine; these species are excluded from the following list.
Agaricus smithii Clavariadelphus species Lactarius scrobiculatus
Albatrellus confluens Clitocybe odor a Leccinum species
Albatrellus ovinus Cortinarius species Macowanites americanus
Armiliaria albolanaripes Gomphidius glutinosus Phaeocollybia species
Armillaria caligata Gomphus species Phellodon species
Boletus pulcherrimus Hydnum species Polyozellus multiplex
Boletus rubripes Hygrophorus agathosmus Rhizopogon parksii
Calvatia fumosa Hygrophorus capreolarius Russula species
Calvatia subcretacea Hygrophorus olivaceoalbus Russula xerampelina
Cantharellus cibarius Hygrophorus pudorinus Tricholoma saponaceum
Catathelasma imperialis Lactarius alnicola Tricholoma virgatum
Catathelasma ventricosa Lactarius representaneus Tylopiluspseudoscaber group
Clavaria purpurea
OTHER TREES 41

FIR (Abies)—Firs, like hemlocks, usually occur in association with other conifers.
They feature many of the same fungi as their close relatives, the spruces. Some species,
however, seem to favor fir over other conifers. These include:

Boletus calopus Echinodontium tinctorium Nivatogastrium nubigenum

Boletus regius Gastroboletus turbinatus Oxyporus nobilissimus
Cortinarius species Hericium abietis Rozites caperata

JUNIPER (Juniperus)—Junipers favor arid habitats (at least in the West) and have
few if any mycorrhizal mates. Therefore, as might be expected, they offer a rather meager
selection of fleshy fungi: a few saprophytes (e.g., Geastrum and Tulostoma species) plus
some wood-rotters (e.g., Daedalea juniperinus, Gloeophyllum striatum, Truncospora
demidoffii). Several of the species listed under cypress and “Sand and Deserts” also
occur under juniper.

CEDAR and INCENSE CEDAR (Thuja and Libocedrus)—Cedars and incense cedars
are not “true” cedars. The “true” or Old World cedars are members of the pine family,
whereas American cedars have scales rather than needles and are more closely related
to juniper and cypress. Our “cedars” harbor several unique fungi (e.g., Tyromyces
amarus, which causes a heart rot of incense cedar), but they have few if any mycorrhizal
mates. Like redwood and cypress, they are blessed mostly with puny saprophytes that
grow best in a mushroom-poor environment (e.g., Leptonia and Hygrocybe species).

MAPLE (Acer)—See “Riparian Woodland.”

BIRCH (Betula)—As any northerner will tell you, birch has dozens of mycorrhizal
associates. In California it does not occur naturally, but planted trees in lawns
and parks often feature Lactarius torminosus, Lactarius pubescens, and/or Paxillus
involutus. Birch forests, on the other hand, are rich in other Lactarii as well as Russulas,
Amanitas, and tasty Leccinums. Several of the more distinctive birch lovers are:

Amanita muscaria Lactarius pubescens Leccinum species

Amanita virosa group Lactarius thejoga/us Paxillus involutus
Cortinarius armillatus Lactarius torminosus Phellinus igniarius
Cortinarius pholideus Leccinum scabrum Piptoporus betulinus
Tomes fomentarius

Lactarius pubescens (see comments under L. torminosus on p. 73), left, and Paxillus involutus
(see p. 477), right, commonly grow on lawns where birches have been planted. Both are poisonous,
at least raw, and both have a strongly inrolled cap margin when young.
Polyporus badius (p. 562) is a small polypore that commonly grows on dead aspen. Note dark cap,
dark stalk, and minute whitish pores.

ASPEN, POPLAR, and WILLOW (Populus and Salix)—In summer the aspen forests of
northern and western North America are excellent foraging grounds for fungi, and in the
fall they put on a dazzling display of leaf color (COLOR PLATE 25). Aspens are not close
relatives of birches, but resemble them in several ways. For instance, both kinds of trees
tend to have whitish bark and a northern distribution, and both feature many species of
Leccinum (all edible) as well as the usual panoply of Russulas and Cortinarii. Cotton¬
woods (poplars) are in the same genus (Populus) as aspens, but usually occur near water.
Among their mycorrhizal mates are the edible Tricholoma populinum; wood-rotters
include Pholiota destruens and sometimes massive clumps of delectable oyster mush¬
rooms (Pleurotus ostreatus). Willows (Salix) are closely related to cottonwoods and
often mingle with them; consequently their fungal flora is similar. A few of the more
distinctive and prominent species associated with aspen, poplar, and willow are:

Armiliaria albolanaripes Lactarius controversus Pleurotus ostreatus

Armiliaria species Leccinum species Polyporus badius
A rmiliaria straminea Morchella species Russula aeruginea
Clavicorona pyxidala Phellinus igniarius Stropharia kauffmanii
Cyptotrama chrysopeplum Pholiota aurivella group Tricholoma flavovirens
Flammulina velutipes Pholiota destruens Tricholoma leucophyllum
Helvetia leucopus Pholiota squarrosa Tricholoma populinum
Hericium ramosum

ALDER (Alnus)—Alders typically grow along streams (see “Riparian Woodland”)

or on logged-over or exposed areas with conifers. Characteristic alder addicts include:

Agaricus subrutilescens Entoloma rhodopolium Pleurotus ostreatus

Alpova diplophloeus Pleurotus dryinus Psathyrella species

DUNG AND MANURE

Cow patties—or almost any kind of dung—can be turned into miniature mushroom
gardens by keeping them in a humid environment. They will yield a fascinating succession
of dainty fungi, but few of them large enough to eat. The mycelium of a dung-inhabiting
fungus has a shorter life span than that of other fungi due to the transient nature of the
substrate. The spores are presumably ingested by cows (or other animals) with the grass
they eat, but pass through the digestive system unscathed, and into the dung, where they
germinate. In addition to the common dung addicts listed below, you’ll find a multitude
of minute Ascomycetes not treated in this book. Some of these are extremely specialized,
living only on lizard dung, frog feces, oppossum excrement, etc.

Agaricus bisporus Clitocybe tarda Peziza fimeti

Agrocybepediades group Conocybe species Peziza vesiculosa
Ascobolus species Coprinus species Psilocybe species
Bolbitius vitellinus Coprobia species Sphaerobolus stellatus
Cheilymenia species Cyathus stercoreus Stropharia semiglobata
Clitocybe nuda Panaeolus species Volvariella speciosa

42
Left: This “meadow muffin” (chunk of cow dung) shows extensive mycelial growth. The mushrooms
belong to the Agrocybe pediades group (p. 468). Right: This is one clump of Lentinusponderosus
(p. 143) that won’t be eaten!

PASTURES
Pastures and meadows offer a wide selection of easily identified edible mushrooms,
notably Agaricus campestris, A. cupreobrunneus, A. arvensis, A. crocodilinus, and the
A. osecanus group, plus Lepiota naucina and Calvatia species. Every fall there is a period
of about one month when our grazed pastures are chock-full of mushrooms, and
there is often another crop in the spring. The following list excludes mushrooms
associated with trees (such as Amanita velosa), which frequently fruit at the edges of
pastures, as well as those that grow on dung. It’s interesting to note that several species
which appear in pastures do not often grow on lawns (and vice versa).

Agaricus arvensis Astraeus species Clitocybe tarda

Agaricus campestris Bovista species Lepiota naucina
Agaricus crocodilinus Calvatia booniana Marasmius oreades
Agaricus cupreobrunneus Calvatia bovista Melanoleuca melaleuca group
Agaricus micromegathus Calvatia cyathiformis Nolanea sericea group
Agaricus osecanus group Calvatia gigantea group Panaeolus campanulatus group
Agaricus xanthodermus Clitocybe brunneocepha/a Psilocybe semilanceata
Agrocybe pediades group Clitocybe dealbata Stropharia species
Amanita sp. (unidentified) Clitocybe subconnexa group Vascellum pratense

LAWNS and GARDENS

To many people, mushroom hunting is synonymous with a day in the wild—
plunging into the dripping depths of a forest, lunging through manzanita thickets, or
meandering through puffball-pocked pastures. However, a mushroom hunt can and
should begin in the mundane confines of your own yard (assuming you have one), and
radiate outward gradually, like the mycelium of a fairy ring. After all, some of the most
delicious (Marasmius oreades, Agaricus campestris) or bizarre (Phallus impudicus)
mushrooms might be growing right outside your door!
Agaricus is the most prominent group of mushrooms in lawns and gardens, and more
than one species of Agaricus often grow together. A. campestris, A. bisporus, and A.
bitorquis are delicious, but care must be taken not to confuse them with A. californicus
and A. xanthodermus, which are poisonous to some people. Other choice morsels include

43
Don’t overlook your toolshed when looking for mushrooms! This hefty morel would have rotted
away uneaten if my wife hadn’t insisted that it was my turn to mow the lawn.

Lepiota rachodes, L. barssii, L. naucina, the omnipresent Marasmius oreades, and an

occasional morel (Morchella).
Lawns and gardens are actually two distinctly different habitats, but are treated
together here because they so frequently overlap. Volvariella speciosa and Agaricus
bisporus, for instance, grow in gardens but not on lawns, whereas Agaricus campestris
and Marasmius oreades grow on lawns but not in gardens. Mushrooms associated with
trees are excluded from the following list though they often appear in people’s yards.
Most yards also qualify as “disturbed ground,” and some may have dung or manure (in
the form of compost or mulch) present.

Agaricus arvensis Clathrus ruber Lysurus cruciatus

Agaricus augustus Clitocybe brunneocephala Lysurus mokusin
Agaricus bernardii Clitocybe dealbata Marasmius oreades
Agaricus bisporus Clitocybe nuda Melanoleuca species
Agaricus californicus Clitocybe tarda Morchella species
Agaricus campestris Conocybe lactea Naematoloma aurantiaca
Agaricus comtulus Conocybe tenera group Nolanea sericea
Agaricus cupreobrunneus Coprinus atramentarius Panaeolus foenisecii
Agaricus micromegathus Coprinus comatus Panaeolus subbalteatus group
Agaricus osecanus group Coprinus micaceus Phallus hadriani
Agaricus perobscurus Coprinus plicatilis Phallus impudicus
Agaricus subrufescens Coprinus species Pholiota species
Agaricus xanthodermus Gymnopilus species Pholiota terrestris
Agrocybepediades group Hohenbuehelia petaloides group Pluteus petasatus
Agrocybe praecox group Lepiota americana Psathyrella candolleana
Agrocybe sororia Lepiota barssii Psathyrella species
Bolbitius vitellinus Lepiota cepaestipes Psilocybe species
Bovista species Lepiota lutea Scleroderma species
Calocybe came a Lepiota naucina Stropharia species
Calvatia species Lepiota rachodes Vascellum pratense
Chlorophyllum molybdites Lepiota species Volvariella speciosa

INDOORS
In some cases a mushroom hunt can begin inside your house. The brown cup fungus,
Peziza domiciliana, is commonly found on walls, floors, carpets, tile, and plaster. Lepiota
lutea consistently pops up in flower pots, though it sometimes finds its way outdoors.

44
INDOORS 45

Several fungi are pests of structural timber (e.g., Serpula lacrymans), and a mysterious
Coprinus invaded the floor of my ’67 Chevy Nova. However, the majority of fungi found
indoors occur outside as well.

Coniophora puteana Lepiota cepaestipes Pluteus species

Conocybe species Lepiota lutea Poria species
Coprinus species Lepiota rachodes Serpula lacrymans
Galerina species Melanotus textilis Thelephora species
Gymnopilus species Mycena species Volvariella species
Hohenbuehelia petaloides group Peziza domiciliana

DISTURBED GROUND
“Disturbed” ground means roadsides, pathsides, gardens, vacant lots, building sites,
and the perimeters of parking lots. Of course, it overlaps with lawns and gardens (or
“cultivated ground”), but we do have a very distinct roadside fungus flora exemplified by
the shaggy mane (Coprinus comatus).
Just what makes certain mushrooms prefer (or even require) disturbed ground is a
mystery. Obviously, the process of disrupting, overturning, or paving the soil must release
certain otherwise inaccessible nutrients, and perhaps the moisture-absorbing properties
and/or pH of the soil are changed. An unsurpassed edible mushroom like Agaricus
augustus could be raised commercially if this puzzle were solved. Some mushrooms, such
as the Lyophyllum decastes group, seem to require disturbed ground whereas other
mushrooms only prefer it. Some show a liking for asphalt and will even poke up through
it—notably Coprinus comatus, Lyophyllum decastes, Agaricus bitorquis, Pisolithus
tinctorius, and Scleroderma geaster. Shaggy manes have even ruined tennis courts!
One asset of these fungi is that they’re easily spotted from the road (or trail).
Consequently, they are tailormade for people who have an extreme allergy to poison oak.
And many are worth stopping for!—Agaricus augustus, A. bitorquis, Coprinus comatus,
Lepiota naucina, Lepiota rachodes, and Lyophyllum decastes. The following list does
not include species that commonly occur in undisturbed as well as disturbed ground.

Agaricus augustus Astraeus species Lepiota rachodes

Agaricus bernardii Calvatia gigantea group Lyophyllum decastes group
Agaricus bitorquis Clitocybe dilatata Morchella species
Agaricus californicus Coprinus atramentarius Pholiota terrestris
Agaricus praeclaresquamosus Coprinus comatus Pisolithus tinctorius
Agaricus species Coprinus micaceus Scleroderma geaster
Agaricus xanthodermus Lepiota naucina Scleroderma species
Aleuria aurantia

BURNED AREAS
Many plants are adapted to growing in burned-over areas. The cones of the knobcone
pine, for instance, release their seeds only when subjected to extreme heat. Similary,
certain mushrooms fruit only in burnt ground, and others show a definite fondness for it.
After the eruptions of Mt. St. Helens, mushrooms were among the first organisms to
“colonize” the devasted slopes of the volcano, and the most reliable way to cultivate
morels is still to set fire to some woods and return the following spring! (This practice
actually had to be outlawed in Europe.)

Anthracobia species Peziza praetervisa Plicaria species

Ascobolus carbonarius Peziza proteana Psathyrella carbonicola
Coltricia species Peziza violacea Psathyrella species
Coprinus species Pholiota brunnescens Pulvinula species
Geopyxis species Pholiota carbonaria Pyronema ompha/odes
Lyophyllum species Pholiota fulvozonata Rhizina undulata
Morchella species Pholiota highlandensis Tarzetta species
Myxomphalia maura
46 HABITATS

SAND and DESERTS

Odd as it may seem, a number of mushrooms grow in sand or sandy soil, including the
delicious Lepiota rachodes (which I have found on Ano Nuevo Island) and various
morels (Morchella species). The deserts of the West support a fascinating but
exasperatingly elusive fungal flora in which the Gasteromycetes (puffballs and allies)
figure prominently. Deserts do not necessarily have sandy soil, but there is considerable
overlap in the two categories, so they are listed together here; those species which grow
exclusively in the desert are marked with an asterisk.
Astraeus hygrometricus Disciseda species Myriostoma coliforme
Battarrea species Geastrum species Phellorina strobilina*
Calvatia species Geopora species Podaxis pistillaris*
Carbomyces species* Longula texensis Schizostoma laceratum*
Chlamydopus meyenianus* Montagnea arenarius* Scleroderma species
Coprinus asterophorus* Morchella species Tulostoma species
Dictyocephalos attenuatus*

SNOWBANKS
The mountains of western North America feature a very characteristic collection of
spring fungi. Some are popularly known as “snowbank mushrooms” because they fruit
around the edges of melting snowbanks; others are more common shortly after the snow
has disappeared. The “snowbank mushrooms” and other spring fungi are more
dependent on snowmelt than rain for their moisture, but the latter can also have a
stimulating effect. Obviously, the higher you go, the later the snow melts, so that in some
areas the spring “snowbank” flora doesn’t appear until early or even mid-summer.
This is one category in which Ascomycetes are more preponderant than
Basidiomycetes. Morels (Morchella species) and false morels (Gyromitra and Helvella
species) are what make the spring fungus flora famous, but dozens of other species occur
also. Many of the larger agarics develop under the duff and show themselves only as low
mounds or “mushrumps” in the humus. The following list includes some typical
“snowbank mushrooms,” plus many that fruit within a month after the snow disappears.
The list does not include fungi with perennial fruiting bodies or species such as Boletus
edulis that may occur in the spring but are more common in the summer and fall.

Agaricus silvicola group Choiromyces alveolatus Geopora cooperi

Agrocybepraecox group Clitocybe albirhiza Guepiniopsis alpinus
Amanita calyptrata Clitocybe species Gyromitra esculent a
Amanita gemmata group Collybia a lb ip Hat a Gyromitra gig as
Armillaria albolanaripes Cortinarius species Helvella leucomelaena
Armillaria olida Cudonia monticola Hygrophorus goetzii
Auricularia auricula Discina species Hygrophorus marzuolus
Caloscypha fulgens Disciotis venosa Hygr ophorus purpurascens
Calvatia species Gautieria species Hygr ophorus sub alpinus

Lyophyllum montanum (p. 175) usually grows nearly melting snow in the spring.
SNOWBANKS 47

Lentinellus monlanus Nivatogastrium nubigenum Ramaria magnipes

Lyophyllum montanum Nolanea vema group Ramaria rasilispora
Lyophyllum species Peziza species Sarcosoma species
Melanoleuca species Phaeolus alboluteus Sarcosphaera crassa
Mitrula species Pholiota species Tyromyces leucospongia
Morchella species Plectania nannfeldtii Verpa bohemica
Mycena griseoviridis Plectania species Verpa conica
Mycena overholtzii Ramaria botrytis

SPECIALIZED HABITATS
Without specialization there would be far less diversity and little—if any—evolution.
All fungi are to some extent specialized, but in some cases the specialization is more
bizarre or extreme—or at least more apparent to us. For instance, it is said that certain
poorly known Laboulbeniomycetes grow only on the left anterior appendage (left front
foot) of their insect host!
Some kinds of fleshy fungi grow only in deep moss or in Sphagnum bogs. Others
grow exclusively on other mushrooms: Asterophora species, Claudopus parasiticus,
Collybia tuberosa, Collybia racemosa, Collybia cookei, Collybia cirrhata, Volvariella
surrecta, Boletus parasiticus, Psathyrella epimyces, and Hypomyces species. Cordyceps
species parasitize insects and certain truffles; and several Omphalina species are asso¬
ciated with algae. Many wood-rotters can grow on cones, but Auriscalpium vulgare
(as well as Baeospora myosura and several Strobilurus species) are restricted to cones.
The very specialized niche has helped Auriscalpium to survive, but if and when conifers
disappear from the earth (as they are slowly doing), so will Auriscalpium!

ANYWHERE and EVERYWHERE

No habitat list is complete without mentioning some of our cosmopolitan fungi or
“mushroom weeds.” Many of these are restricted to wooded areas, others (those marked
by an asterisk) are not. The best of the lot from an edibility standpoint are Armillariella
mellea, Clitocybe nuda, and Morchella species.

Agaricus californicus* Collybia dryophila Naematoloma fasciculare*

Agaricus hondensis Collybiafuscopurpurea group* Peziza species*
Agaricus praeclaresquamosus Cortinarius glaucopus group Phellinus pini
Agaricus silvicola group Cortinarius species Pholiota terrestris*
Agaricus subrutilescens Elaphomyces species Phvllotopsis nidulans
Agaricus xanthodermus* Fomitopsis pinicola Pluteus cervinus*
Aleuria aurantia* Galerina species* Poria species*
Amanita gemmata group Ganoderma applanatum group* Psathyrella candolleana*
Amanita pachycolea Geastrum species* Psathyrella species*
Amanita pantherina Helvella lacunosa* Rhodoeybe nuciolens
Amanita vaginata Inocybe geophylla Russula albonigra
Armillariella mellea* Inocybe sororia Russula brevipes
Boletus chrysenteron Inocybe species Russula sororia group
Boletus spadiceus Laccaria species Russula species
Boletus subtomentosus Lactarius chrysorheus Scleroderma cepa group*
Boletus truncatus Lactariusfragilis Scleroderma citrinum
Boletus zel/eri Lactarius vinaceorufescens Scleroderma species*
Cantharel/us cibarius Laetiporus sulphureus* Stereum species*
Cantharellus subalbidus Lepiota rachodes* Thelephora species*
Clavaria vermicularis* Leucopaxillus albissimus Trametes versicolor*
Clitocybe inversa Leucopaxillus amarus Trichaptum species*
Clitocybe nebular is Lycoperdon species* Tricholoma saponaceum
Clitocybe nuda* Melanoleuca melaleuca group* Tubaria species*
Clitocybe subconnexa group* Morchella species* Tyromyces species
Clitopilus prunulus* Mycena species* Volvariella speciosa*
48

SEVENTY DISTINCTIVE MUSHROOMS

(A quick-reference check list)

COMMON NAME SCIENTIFIC NAME EDIBILITY

□ Delicious Milk Cap Lactarius deliciosus edible

(p. 68; plate 2)
□ Bleeding Milk Cap Lactarius rubrilacteus edible
(p. 68; plates 3, 6)
□ Indigo Milk Cap Lactarius indigo edible
(p. 69; plate 4)
□ Candy Caps Lactarius fragilis, edible
(pp.. 80-82; plate 10) L. rufulus, L. camphoratus
□ Short-Stemmed Russula Russula brevipes edible
(P- 87)
□ Emetic Russula Russula emetica group poisonous
(P- 97)
□ Rosy Russula Russula rosacea poisonous
(p. 99; plate 13)
□ Witch’s Hat Hygrocybe conica doubtful
(p. 116; plate 19)
□ Ivory Waxy Cap Hygrophorus eburneus edible
(P- 119)
□ Oyster Mushroom Pleurotus ostreatus edible
(p. 134; plates 27, 28)
□ Jack-O-Lantern Mushrooms Omphalotus species poisonous
(pp. 146-148; plates 40, 41)
□ Blewit Clitocybe nuda edible
(p. 153; plate 32)
□ Man On Horseback Tricholoma flavovirens edible
(p. 179; plate 33)
□ White Matsutake Armillaria ponderosa edible
(p. 191; plate 37)
□ Honey Mushroom Armillariella mellea group edible
(p. 196; plates 39, 42)
□ Fairy Ring Mushroom Marasmius oreades edible
(p. 208; plates 38, 47)
□ Death Cap Amanita phalloides deadly
(p. 269; plate 50) poisonous
□ Destroying Angels Amanita ocreata, A. verna, deadly
(pp. 271-273; plate 53) A. virosa, A. bisporigera poisonous
□ Fly Agaric Amanita muse aria poisonous
(p. 282; plates 58, 59)
Armillariaponderosa, better known as the matsutake or white matsutake, is a mushroom well worth
knowing. Its odor is unique: something like a cross between “red hots” and dirty socks.

COMMON NAME SCIENTIFIC NAME EDIBILITY

□ Caesar’s Amanita Amanita caesarea group edible

(p. 284; plate 60)
□ Coccora Amanita calyptrata edible with
(p. 284; plates 61-63) caution
□ Grisette Amanita vaginata edible
(p. 288)
□ Green-Spored Parasol Chlorophyllum molybdites poisonous
(p. 295)
□ Shaggy Parasol Lepiota rachodes edible with
(p. 297; plates 69) caution
□ Parasol Mushroom Lepiota procera edible
(P- 298)
□ Meadow Mushroom Agaricus campestris edible
(p. 318; plate 71)
□ Yellow-Staining Agaricus Agaricus xanthodermus poisonous
(p. 329)
□ Horse Mushrooms Agaricus arvensis, edible
(pp. 332-334; plate 78, 79, 81) A. osecanus group
□ The Prince Agaricus augustus edible
(p. 337; front cover, plate 77)
□ Shaggy Mane Coprinus comatus edible
(p. 345; plate 85)
□ Sulfur Tuft Naematoloma fasciculare poisonous
(p. 382; plate 92)
□ Scaly Pholiota Pholiota squarrosa doubtful
(p. 389; plates 96, 97)
□ Violet Cortinarius Cortinarius violaceus edible
(p. 446; plate 109)
□ Pine Spikes Chroogomphus species edible
(pp. 484-487; plate 113)

49
Boletus aereus is one of many delectable boletes. These young specimens are easily recognized by
their white pore surface and dark cap that is covered with a fine whitish bloom.

COMMON NAME SCIENTIFIC NAME EDIBILITY

□ Slippery Jacks Suillus species edible

(pp. 498-505; plates 114-121)
□ Admirable Bolete Boletus mirabilis edible
(p. 521; plates 127, 129)
□ Butter Boletes Boletus appendiculatus, edible
(pp. 525-526; plates 133, 134) B. regius
□ Satan’s Bolete Boletus satanas poisonous
(p. 527; plate 137)
□ Apple Bolete Boletus frostii edible
(p. 528; plate 139)
□ White King Bolete Boletus barrowsii edible
(p. 529; plate 141)
□ King Bolete Boletus edulis edible
(p. 530; plates 142-144)
□ Queen Bolete Boletus aereus edible
(p. 531; plate 140)
□ Manzanita Bolete Leccinum manzanitae edible
(p. 539; plate 145)
□ Aspen Boletes Leceinum insigne & edible
(pp. 540-541; plate 147) close relatives
□ Old Man Of The Woods S trobilomyces floe copus edible
(p. 543)
□ Beefsteak Fungus Fistulina hepatiea edible
(p. 553; plate 152)
□ Hen Of The Woods Grifola frondosa, edible
(pp. 564-565) G. umbellata
□ Sulfur Shelf Laetiporus sulphureus edible with
(p. 572; plates 154, 155) caution
□ Artist’s Conk Ganoderma applanatum not edible
(p. 576) group

50
SEVENTY DISTINCTIVE MUSHROOMS 51

COMMON NAME SCIENTIFIC NAME EDIBILITY

□ Varnished Conks Ganoderma lucidum, not edible

(pp. 577-578; plate 156) G. tsugae, G. oregonense
□ Turkey Tail Trametes versicolor not edible
(p. 594; plate 158)
□ Hericiums Hericium species edible
(pp. 613-616; plates 163, 164)
□ Hedgehog Mushrooms Dentinum repandum, edible
(pp. 618-619; plates 161, 162) D. umbilicatum
□ Pink-Tipped Coral Mushroom Ramaria botrytis edible
(p. 656)
□ Cauliflower Mushrooms Sparassis species edible
(p. 657; plate 172)
□ Scaly Chanterelle Gomphus floccosus group doubtful
(p. 661; plate 174)
□ White Chanterelle Cantharellus subalbidus edible
(p. 662; plate 179)
□ Chanterelle Cantharellus cibarius edible
(p. 662; plates 175, 177, 178)
□ Horn Of Plenty Craterellus cornucopioides, edible
(pp. 666-668; plate 182) C. fallax
□ Witch’s Butter Tremella mesenteriea edible
(p. 673; plate 170)
□ Giant Puffballs Calvatia gigantea group, edible
(pp. 682-684; plates 184, 186) C. booniana
□ Sierran Puffball Calvatia sculpta edible
(p. 684)
□ Dead Man’s Foot Pisolithus tinctorius not edible
(p. 712; plates 185, 188, 189)
□ Stinkhoms Phallus impudicus & edible
(pp. 768-770; plates 193, 194) close relatives
□ Dog Stinkhorns Mutinus caninus, edible
(p. 771; plates 195, 196) M. elegans
□ Morel Morchella eseu/enta & edible
(p. 787; plate 203) close relatives
□ Black Morel Morchella data group edible
(p.790; plates 199, 202)
□ False Morel Gyromitra esculenta edible/
(p. 801; plate 206) poisonous
□ Fluted Black Elfin Saddle Hclvella lacunosa edible
(P-815)
□ Orange Peel Fungus Aleuria aurantia edible
(p. 837; plate 208)
52

KEY TO THE MAJOR GROUPS OF FLESHY FUNGI

1. Spores produced on mother cells called basidia; fruiting body variously shaped
(see pp. 52-54) .Basidiomycotina, p. 57
1. Spores produced inside mother cells called asci; fruiting body variously shaped
(see p. 55) .Ascomycotina, p. 782

BASIDIOMYCETES

Fruiting body with a cap and stalk, or just

a cap; spores borne on gills (radiating
blades) on underside of cap; spore print
obtainable (if spores are being produced)
. AGARICS (GILLED
MUSHROOMS), p. 58

Fruiting body with a cap and stalk,

usually vase-shaped or trumpetlike at
maturity, not gelatinous; fertile under¬
side of cap smooth, wrinkled, or with
decurrent veins or folds .
. CHANTERELLES, p 658

Fruiting body with a cap and stalk, fleshy

(not tough or woody); underside of cap
with a spongy, often separable layer of
tubes or pores; stalk more or less central;
usually on ground but occasionally on
wood . BOLETES, p. 488

longitudinal section
showing the tube layer

Fruiting body shelflike, bracketlike,

crustlike, or with a cap and stalk; usually
tough or woody but sometimes fleshy,
usually on wood but sometimes terres¬
trial; spores produced in a layer of tubes
or pores that usually line underside of cap.
Stalk when present usually off-center or
lateral (but sometimes central); tube
layer not normally peeling away easily
from cap . POLYPORES & BRACKET
FUNGI, p. 549
(but if fruiting body is black and char¬
coal-like with concentrically zoned flesh,
see Flask Fungi, p. 878)
MAJOR GROUPS OF FLESHY FUNGI 53

Fruiting body variously shaped (with or

without a cap), but always gelatinous or
very rubbery; usually growing on wood
..JELLY FUNGI, p. 669

Fruiting body crustlike or bracketlike

or occasionally with a cap and stalk,
usually tough, not gelatinous; fertile
surface smooth, wrinkled, veined, or
warted but lacking pores, tubes, or spines;
usually growing on wood . CRUST
& PARCHMENT FUNGI, p 604
(also see Morels, Elfin Saddles, & Cup
Fungi, p. 783)

Fruiting body bearing its spores on

downward-pointing spines or “teeth”;
spines either lining the underside of a cap
or suspended like icicles from a cushion
of tissue or a branched framework; stalk
present or absent; on ground or wood ..
. TEETH FUNGI, p. 611

Fruiting body erect, unbranched (club¬

like) or profusely branched from a
common base or “trunk” (coral-like);
cap absent; spores borne on the smooth
to slightly wrinkled surfaces of the
upright clubs or branches .
. CORAL & CLUB FUNGI, p. 630
(also see Earth Tongues, p. 865)

Fruiting body reminiscent of an un¬

opened, aborted, or deformed agaric,
consisting of a spore case or “cap” and
stalk; stalk long or very short, usually
percurrent (i.e., extending all the way
to top of fruiting body); spore mass or
fertile tissue usually composed of plates,
branching cavities, or contorted “gills”
which may or may not be exposed; spore
print not obtainable; found mainly in
deserts and mountains .
. GASTROID AGARICS, p. 724
54 MAJOR GROUPS OF FLESHY FUNGI

Fruiting body round to oval or pear-

shaped, or the outer skin splitting into
starlike rays; interior (spore mass) firm
when young but powdery or dusty at
maturity; spores borne inside a spore
case or numerous lentil-like capsules;
stalk absent or present only as a narrowed
sterile base or rootlike fibers; usually
growing above the ground .
PUFFBALLS & EARTHSTARS, p 677
longitudinal sections

Fruiting body with a puffball-like spore

case mounted on a well-developed,
clearly differentiated stalk; spore mass
(interior of spore case) dusty or powdery
at maturity; stalk terminating at base of
spore case (i.e., not percurrent); often
found in deserts, sand, or waste places ..
. STALKED PUFFBALLS, p. 715

Fruiting body minute (often less than

1 cm broad), shaped like a miniature
bird’s nest with one or more small “eggs”
inside (but top of “nest” often covered by
a layer of tissue when young) .
. BIRD’S NEST FUNGI, p. 778

Fruiting body emerging from an “egg”

whose skin forms a volva (sack) at base
of mature fruiting body; fruiting body
with a cap and stalk or just a stalk, or
with arms, tentacles, or a branched or
latticed framework; inside or outside
surfaces of fresh fruiting body coated
with a foul-smelling, greenish to brown
to blackish spore slime .
. STINKHORNS, p. 764

Fruiting body usually underground,

round to oval or knobby; interior (spore
mass) firm to spongy or gelatinous but
not often powdery, usually composed of
small chambers or cavities or occasionally
plates; columella (internal stalk) present
or absent .. FALSE TRUFFLES, p. 739
MAJOR GROUPS OF FLESHY FUNGI 55

ASCOMYCETES

Fruiting body usually underground,

round to oval or knobby; interior hollow
or with several large cavities or with
canals or interior solid and marbled with
sterile veins; texture of interior firm or
sometimes powdery but not gelatinous;
one hollow and
one solid but marbled
columella (internal stalk) only rarely
present . TRUFFLES, p. 841

Fruiting body growing on insects, spiders,

or truffles or engulfing other mushrooms
in a pimpled, powdery, or lumpy layer of
tissue or growing on wood; if on wood
then dark brown to black (or black
beneath white spore powder) and tough,
hard, or charcoal-like .
. FLASK FUNGI, p. 878

Fruiting body disc-shaped (flat) to cup¬

shaped, vaselike, or earlike (with or
without a stalk) or fruiting body with
a stalk and clearly differentiated cap; cap
when present cup-shaped to saddle-
shaped, irregularly lobed, brainlike,
thimble-like, or pitted .
MORELS, ELFIN SADDLES, & CUP
FUNGI, p.783
(also see Earth Tongues, p. 865)

Fruiting body usually erect, simple, and

unbranched (clublike), but usually with
a differentiated (often flattened or
swollen) fertile “head” or even a small,
rounded or wrinkled cap; fruiting body
usually small, if with a small cap then
usually tough or gelatinous .
. EARTH TONGUES, p 865
(also see Morels, Elfin Saddles, & Cup
Fungi, p. 783)
The shaggy mane, Coprinus comatus (p. 345), is one of many Basidiomycetes. It is a familiar sight
along roads and parking lots in cool wet weather.

56
 57

Basidiomycetes

BASIDIOMY COTIN A
THE Basidiomycetes are a large group (subdivision) of fungi that bear spores on (but not
inside) specialized cells called basidia. Two major classes of Basidiomycetes produce
fleshy fruiting bodies or “mushrooms”—the Hymenomycetes and the Gasteromycetes,
keyed below. Other classes of Basidiomycetes, such as the rusts and smuts, are not treated
in this book.
Key to the Basidiomycetes
1. Basidia and spores borne externally (on the exposed surfaces of gills, tubes, spines, branches,
lobes, etc.); spores forcibly discharged at maturity, i.e., a spore print often (but not always)
obtainable; fruiting body with a cap and stalk, or clublike, or branched, orbracketlike, or crust¬
like (without a stalk or sometimes without a cap) or lobed or bloblike, etc.2
1. Basidia and spores borne internally (inside the fruiting body or inside a spore case or small cap¬
sules); spores not forcibly discharged, thus a spore print unobtainable Gasteromycetes, p. 676
2. Fruiting body at first egglike with a gelatinous interior, the outer skin then breaking and an
unbranched (often phallic), branched, or lattice-like fruiting body emerging; spores contained
in a greenish to brown or black, foul-smelling slime that coats all or part of the fresh fruiting
body (but slime dispersed by flies or rain); gills and tubes absent .. . Gasteromycetes, p. 676
2. Not as above .Hymenomycetes, below

HYMENOMYCETES
IN this large divison of the Basidiomycetes the spore-bearing cells (basidia) form a layer
or palisade (hymenium) on an exposed surface or surfaces of the fruiting body, and the
spores are forcibly discharged at maturity. Terrestrial forms are usually furnished with a
cap and stalk, but many of the wood-inhabiting types are shelflike, hooflike, bracketlike,
crustlike, or bloblike. In the more primitive forms the hymenium is smooth to slightly
wrinkled or warted, while in the more specialized (advanced?) types it takes the form of
gills, tubes, spines, or upright branches.
The Hymenomycetes are traditionally divided into two orders: the Agaricales include
the gilled mushrooms (agarics) and fleshy tube fungi (boletes), while the Aphyllophorales
are an artificial group embracing the polypores, crust fungi, coral fungi, chanterelles,
and teeth fungi. A third group, the jelly fungi (Tremellales, Dacrymycetales, and Auri-
culariales), are treated in this book as Hymenomycetes, although some taxonomists
consider them to be only distantly related.
In the absence of a detailed fossil record, there is a great deal of speculation as to
whether the Hymenomycetes evolved from the Gasteromycetes or vice-versa. It may
actually be that evolution has flowed both ways. Spore dispersal is enhanced if the spores
are actively (forcibly) discharged as in the Hymenomycetes, but the enclosed fruiting body
of the Gasteromycetes is advantageous to spore development because it affords greater
protection against the environment. Thus, many Hymenomycetes have adapted to
inhospitable (cold, hot, or arid) environments by fruiting underground and/or never
exposing their hymenium—thus becoming “gastroid” and losing their ability to discharge
spores in the process. On the other hand, there may be lines of evolution leading from
primitive Gasteromycetes to certain Hymenomycetes through the incremental develop¬
ment of a hymenium, stalk, and cap.

♦Since most fungi decay quickly, they are far less apt to be fossilized than plants.
58 HYMENOMYCETES

Key to the Hymenomycetes

1. Underside of cap with hundreds of pores (tube mouths) which may be obvious or very minute;
tube layer visible when fruiting body is cut longitudinally (perpendicularly) .2
1. Not as above; pores and tubes absent . 3
2. Fruiting body fleshy, rapidly decaying, usually but not always terrestrial; stalk typically central
or nearly so; tubes usually—but not always—easily separable from cap (and often from each
other); veil present or absent .Boletaceae (a family in the Agaricales), p. 488
2. Fruiting body usually (but not always!) tough in age and usually (but not always!) growing
on wood; stalk absent or lateral to off-center, or if central then tube layer not easily separable
from cap; veil absent .Aphyllophorales, p. 548
3. Underside of cap with radiating blades (gills) .Agaricales, below
3. Gills absent (but spines, warts, folds, veins, or wrinkles may be present) .4
4. Fruiting body gelatinous (jellylike) or very rubbery; usually (but not always) growing on wood;
basidia partitioned or forked (under the microscope) .Tremellales & Allies, p. 669
4. Not as above .Aphyllophorales, p. 548

Agarics (Gilled Mushrooms)

AGARICALES
THE agarics, or gilled mushrooms, bear their spores on radiating blades or plates called
gills. They are by far the most familiar, numerous, and complex group of fleshy fungi, and
are thought to have arisen from several different ancestors. All of the gilled mushrooms
were originally grouped in a single massive genus, Agaricus, but in the 19th century a
Swedish naturalist, Elias Fries (the “father” of mushroom taxonomy), divided Agaricus
into a number of smaller genera based on easily ascertained features such as attachment
of the gills, texture of the stalk, presence versus absence of a veil, and color of the spores.
In the ensuing years, agaricologists have tried to achieve a more “natural” classification
(i.e., one that reflects common ancestry rather than superficial similarities), and in so doing
have come to rely heavily on microscopic and chemical characters, such as the shape and
arrangement of the cells in the gill tissue and cap cuticle and the shape, ornamentation, and
amyloidity of the spores. Over one hundred genera of gilled mushrooms are now recog¬
nized, including most of Fries’ original genera, albeit often in modified form (for example,
Agaricus (not to be confused with the term “agaric”), in its modern sense, is restricted to
the cultivated mushroom and its close relatives). The “Friesian” system, however, has
remained popular among amateurs or “toadstool-testers” because of its simplicity.
Since modern agaric taxonomy is based on so many seemingly esoteric characters, and
since most people have neither a microscope, nor the requisite chemical reagents, nor the
necessary training to correctly use a microscope and reagents if they had them, a com¬
promise between the modern and Friesian systems has been used in this book, to wit: most
of the modern genera and families have been recognized, but the characters used to dis¬
tinguish them are largely Friesian (macroscopic). The Agaricales have been divided into
fourteen families, which are keyed out here. The boletes constitute a fifteenth family, but
are treated separately. In order to use the key it is necessary to know the spore color—
preferably by taking a spore print. It is also necessary to have some patience—or at least
to realize that subjectivity (e.g., waxy versus non-waxy gills) and fallibility are the ineluc¬
table byproducts of any key that relies on macroscopic (field) characters, when the critical
distinctions between the different entities are in fact microscopic.
AGARICALES 59

A pictorial chart of gilled mushroom genera has also been provided on pp. 61-62,
thereby circumventing the family category. It is by no means infallible, but as already
pointed out, neither is the dichotomous key, nor for that matter, are you and I.
Key to the Agaricales
l. Gills deformed or contorted or branched to form large cavities, sometimes never exposed; spores
not forcibly discharged, hence spore print not obtainable; found mostly in deserts and
mountains .(see Podaxales & Allies, p. 724)
1. Not as above; spores forcibly discharged, hence a spore print obtainable if spores are being
produced; gills exposed at maturity; common and widespread.2
2. Spore print white to buff, yellow, yellow-orange, or lilac-tinged . 3
2. Spore print some other color (pinkish, salmon, yellow-brown, brown, rusty-orange, rusty-
brown, chocolate-brown, purplish, greenish, black, etc.) . 10
3. Universal veil enveloping young specimens and forming a volva at base of stalk when it ruptures
and/ or leaving numerous remnants (warts or flat patches) on cap .... Amanitaceae, p. 262
3. Neither volva nor warts present (but cap and stalk may have scales or fibrils) .4
4. Gills typically free and veil present; veil usually forming an annulus (ring) on stalk, or if not then
stalk typically scaly or slimy below the veil . 5
4. Not as above; veil absent, or if present then gills normally attached to stalk .6
5. Cap slimy or viscid when moist; stalk sometimes viscid also; gills tissue divergent (under the
microscope); not common .Amanitaceae, p. 262
5. Cap dry or only slightly viscid when moist; stalk dry; gill tissue not divergent; common .
.Lepiotaceae, p. 293
6. Gills decurrent and foldlike (at least when young), i.e., gills thick, blunt, shallow, and usually
forked or with cross-veins .(see Cantharellaceae, p. 658)
6. Not as above; gills usually platelike or bladelike .. 7
7. Gills and/ or flesh exuding a latex (milk or juice) when broken; stalk typically more than 3 mm
thick; spores with amyloid warts or ridges .Russulaceae, p. 63
7. Not as above . 8
8. Fruiting body brittle and rather rigid, the stalk snapping open cleanly like a piece of chalk (i.e.,
without fibrous context); cap usually plane to depressed at maturity; stalk typically at least
3 mm thick; veil absent; usually but not always terrestrial; cap and stalk tissue typically con¬
taining nests of sphaerocysts; spores with amyloid warts or ridges .Russulaceae, p. 63
8. Not with above features . 9
9. Gills soft and clean, with a waxy appearance or texture; cap often brightly colored; stalk more
or less central; usually terrestrial; basidia long and narrow (at least six times as long as the
spores); spores smooth .Hygrophoraceae, p. 103
9. Not as above; gills not normally waxy; stalk central to lateral or absent; on ground or wood
.Tricholomataceae, p. 129
10. Spore print pinkish to flesh-colored, salmon, pinkish-cinnamon, or sordid reddish . 11
10. Spore print some shade of orange, brown (including cinnamon-brown), green, purple, gray, or
black . 16
11. Universal veil present, usually forming a saclike volva at base of stalk .... Pluteaceae, p. 253
11. Not as above; volva typically absent . 12
12. Gills typically free at maturity; growing on wood (but wood sometimes buried!) or sawdust or
sometimes in lignin-rich humus; spores not angular .Pluteaceae, p. 253
12. Not as above . 13
13. Odor strong (fishy or like cucumber) and stalk velvety or arising from an underground, often
hollow “tuber” or found on wood and fruiting body pinkish to yellow-orange with a reticulate
(veined) cap surface; spores not angular; not common .Tricholomataceae, p. 129
13. Not as above; common . 14
14. Spore print pale pinkish, pinkish-buff, or pinkish-cream; gills sometimes purple or purple-tinged
when fresh; spores not angular .Tricholomataceae, p. 129
14. Spore print usually deeper in color (flesh-colored to sordid reddish, salmon, etc.); gills not often
purple; spores usually angular in end and/or side view .15
60 AGARICALES

15. Usually growing in clusters; flesh fragile; spores not angular; not common, at least in the western
states . Coprinaceae, p. 341
15. Not as above; spores typically angular in end and/or side view; very common and widely
distributed . Entolomataceae, p. 238
16. Spore print greenish to grayish-olive . 17
16. Not as above . 19
17. Gills adnate to decurrent .Gomphidiaceae, p. 481
17. Gills free or only slightly attached . 18
18. Gills blood-red to reddish to dark brown .Agaricaceae, p. 310
18. Gills white to dull greenish or grayish .. Lepiotaceae, p. 293
19. Spore print purple-brown to purple-gray, purple-black, smoky-gray, black, chocolate-brown,
or deep brown . 20
19. Spore print rusty-orange to rusty-brown, cinnamon-brown, yellow-brown, dull brown, bright
brown, cigar-brown, etc. (but may appear darker in heavy deposits!). 26
20. Gills usually decurrent (but sometimes adnate); spore print smoky-olive to smoky-gray to black;
associated with conifers .Gomphidiaceae, p. 481
20. Not as above; gills free to adnexed, adnate, or occasionally decurrent . 21
21. Gills and/ or cap auto-digesting (i.e., turning into an inky black mass) at maturity; spore print
black . Coprinaceae, p. 341
21. Not as above . 22
22. Veil present, usually forming an annulus (ring) on stalk; gills free or nearly free at maturity,
whitish to pinkish when young but becoming chocolate-brown or darker in age; cap not deeply
striate; spore print chocolate-brown .Agaricaceae, p. 310
22. Not as above . 23
23. Cap brightly colored (yellow, green, orange, red, etc.) .Strophariaceae, p. 367
23. Cap dull (some shade of brown, buff, gray, white, etc.) .24
24. Gills decurrent; cap margin inrolled when young; spore print reddish-brown or cocoa-brown
. Paxillaceae, p. 476
24. Not as above . 25
25. Cap usually viscid when moist (but may dry out!); fruiting body fragile or not fragile; cap cuticle
typically filamentous (under microscope).Strophariaceae, p. 367
25. Cap usually not viscid; fruiting body usually quite fragile; cap cuticle typically cellular (see
illustrations on p. 19) . Coprinaceae, p. 341
26. Growing on other mushrooms . Tricholomataceae, p. 129
26. Not as above . . 27
27. Stalk absent or rudimentary; usually growing shelflike on wood . 28
27. Stalk present; on wood or ground . . 30
28. Fruiting body very tough, leathery, or corky .(see Polyporaceae & Allies, p. 549)
28. Not as above; fruiting body fleshy . 29
29. Gills forked or with cross-veins or even with pores near base of cap . Paxillaceae, p. 476
29. Not as above . Cortinariaceae, p. 396
30. Fruiting body small, fragile, often withering quickly; cap usually oval to conical or bell-shaped;
usually growing in grass, dung, or gardens; cap cuticle typically cellular (see illustrations on
p. 19) .Bolbitiaceae, p. 466
30. Not with above features . 31
31. Gills typically decurrent, often forked (or even forming pores near stalk)and often peeling easily
from cap; margin of cap usually inrolled when young; veil absent; spore print yellowish to
brown; stalk fleshy . Paxillaceae, p. 476
31. Not as above; gills occasionally decurrent but usually not; veil present or absent; spore print
variously colored (rusty-orange, rusty-brown, dull brown, etc.); very common .
.. Cortinariaceae, p. 396
 61
PICTORIAL KEY TO GILLED MUSHROOMS
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62

Spore print white to buff, pale pinkish to Spore print brown to rusty-brown, gray, purple-
Characteristics pinkish-buff, yellowish, yellow- reddish to yellow-brown, cinnamon-brown, brown, or deep
orange, greenish, or tinged lilac flesh-color or rusty-orange brown
PICTORIAL KEY TO GILLED MUSHROOMS
 63

spores

RUSSULACEAE
THIS family differs fundamentally from all other gilled mushrooms in the anatomy of
the cap and stem tissue: nests or rosettes of large, roundish cells called sphaerocysts are
interspersed with the usual filamentous hyphae, giving the mushroom a characteristic
brittle, granular texture. In addition, the spores are conspicuously ornamented with
strongly amyloid warts, spines, and/or ridges. These unique features suggest that the
Russulaceae evolved independently from other agarics—perhaps from underground
ancestors. Some toadstool taxonomists consequently place them in a separate order,
the Russulales, rather than in the Agaricales.
There are two large genera in the Russulaceae: Lactarius, which exudes a milk or juice
(latex) when broken, and Russula, which does not. Though they’re often characterized as a
“white-spored” family, their spore color ranges from white to yellow, buff, or ochraceous.
Lactarius and Russula are mycorrhizal and often fruit together in spectacular abun¬
dance. From an edibility standpoint they are good for beginners because few, if any, of the
mild-tasting species are dangerous. Those species with an acrid or bitter taste are best
avoided since many are poisonous or indigestible, at least raw. To determine whether or
not a species is acrid, chew on a small piece of the cap and gills for a minute, then spit it
out. If it is acrid, you’ll experience a burning sensation on your tongue—in fact, you may
have to wash out your mouth to get rid of the taste! If you’re not sure whether it’s acrid,
then it’s probably “mild.” Be careful—some species really pack a wallop!
Key to the Russulaceae
1. Fresh fruiting body usually exuding a latex (milk or juice) when broken (the latex is best seen
by cutting the gills near the stalk)* .Lactarius, p. 64
1. Latex absent* .Russula, p. 83

*In dry or very wet weather, Lactarius species with a scanty latex may exhibit none at all. The following
characteristics will help distinguish such specimens from Russula:
If the fruiting body has greenish stains and colored flesh, it is probably a Lactarius.
If the cap is brightly colored and has white flesh, it is probably a Russula.
If the cap margin is striate, it is probably a Russula.
If the gills, stalk, or flesh are some color besides white or yellow when fresh, it may well be a
Lactarius. (Some Russulas, however, discolor reddish, gray, brown, or black in age.)
If the stalk is scrobiculate (pitted with darker spots), it is a Lactarius.
If the cap margin is bearded, it is a Lactarius.
If none of the above are applicable, try Russula first, then Lactarius.

The latex which distinguishes Lactarius from Russula is best seen by cutting the mushroom near the
juncture of the gills and stalk. In many species, such as L. argillaceifolius var. megacarpus, shown
here, the latex is milky white, but in some species it is brightly colored and in others practically absent.

■
64

LACTARIUS (Milk Caps)

Typically terrestrial, mostly medium-sized woodland mushrooms. CAP usually depressed at
maturity, usually with an mrolled or incurved margin when young, sometimes concentrically zoned.
Flesh crisp, brittle, usually exuding a latex when broken; taste mild or acrid. GILLS attached
(usually adnate to decurrent), variously colored. STALK typically rigid, brittle, snapping open
cleanly like chalk; central, often scrobiculate. VEIL and VOLVA absent. SPORE PRINT white to
buff or yellowish. Spores with amyloid warts and/or ridges. Cap tissue nearly always with nests of
sphaerocysts.

LACTARIUS species are called milk caps because they exude a milky or juicy fluid
(latex) when broken (see Color Plate 6 and photo on p. 63). In dry or very wet weather
they may lack a distinct latex (especially if old), but their crisp or brittle flesh distinguishes
them from all but the Russulas (see footnote at bottom of p. 63 for a more detailed
comparison). A few Mycenas possess a latex, but they strike a radically different pose:
a conical to bell-shaped cap perched on a long, thin, fragile stalk.
The most important features to note on any Lactarius are the color of the latex, and
the color changes the latex undergoes or produces on surrounding tissue when exposed
to the air. The colors should always be noted on fresh material. The latex may react with
the air rapidly (within 45 seconds) as in L. vinaceorufescens, or slowly (within 5 hours)
as in L. deliciosus, or not at all, as in L.fragilis. The taste—whether mild or acrid (peppery)
—is also significant, as are microscopic features such as the structure of the cap cuticle
and pattern of ornamentation on the spores. It is also helpful to note whether or not the
stalk is scrobiculate (pitted with darker spots).
Lactarius contains a number of delectable collectables. The candy cap (L.fragilis) is the
tastiest of the local species, besides being one of the most abundant. The green-staining
milk caps (L. deliciosus, L. indigo, et al) are edible and popular, and in eastern North
America, L. volemus, L. corrugis, and L. hygrophoroides are very good. Many species,
on the other hand, are poisonous or indigestible. As a rule, the peppery-tasting ones
should be avoided as well as those with yellow- or purple-staining latex. In Europe, some of
the acrid species are eaten after pickling or parboiling, but it hardly seems worth the effort
or risk to do so.
Lactarius is a very large and complex genus, with over 200 species described from North
America. It is especially diverse in eastern—and more specifically southeastern—North
America. In fact, the milk caps are roughly to eastern hardwood forests what the
Russulas are to the coniferous forests of the West—so exceedingly diverse and ubiquitous
as to seem like “mushroom weeds.” In California, however, there are fewer than 50 re¬
corded species of Lactarius, and in our area there are only about a dozen. As a result,
they are much easier to identify than their bewildering brethren, the Russulas. Like the
Russulas, they are mycorrhizal, mostly terrestrial woodland fungi, but often appear at the
edges of woods or on tree-studded lawns. Many are specific to certain hosts—e.g., L.
torminosus grows with birch. In California the milk caps fruit from late summer through
the winter and sometimes rival the Russulas in abundance.
The publication in 1979 of The North American Species of Lactarius byL. R. Heslerand
Alexander Smith has resulted in a large number of name changes. For instance, the edible
species widely known as L. sanguifluus is now L. ruhrilacteus and our L. trivialis has
become L. argillaceifolius. Those of you who find it hard enough to master one scientific
name for a mushroom without being responsible for two or three can salvage some solace
in the fact that while the names have changed, the mushrooms haven’t. In other words,
so long as you know how to recognize a particular species, it hardly matters what you call
it—unless you are living with a taxonomist! Twenty-one species of Lactarius are de¬
scribed here and many others are keyed out and/ or mentioned.
LACTARIUS 65

Key to Lactarius
l. Latex colored (red, orange, blue, brown, etc.), but often scanty or even absent; fruiting body
often (but not always) staining greenish in age or where bruised (slowly, within 5 hours) . 2
1. Latex white to buff or clear (but may stain yellow or some other color when exposed!) .... 9
2. Latex blue; entire fruiting body blue to bluish-gray (or greenish-stained) . . . L. indigo, p. 69
2. Fruiting body not blue (may be greenish-stained), or if partly blue then latex not blue .... 3
3. Latex carrot-orange to salmon-orange, yellow-orange, or rusty-orange (but may darken to
red after exposure to air) . 4
3. Not as above; latex red, purplish, brown, etc., or if orange in base of stalk then bluish in cap 5
4. Cap whitish when fresh; found in southeastern U.S. . . L. salmoneus (see L. deliciosus, p. 68)
4. Not as above; widespread and common .L. deliciosus & others, p. 68
5. Latex dark red to purplish; fruiting body never bluish (but often greenish-stained) . 6
5. Latex purple-brown to reddish-brown to brown or yellow-brown, or bluish in cap and red
or orange in stalk; fruiting body sometimes with blue tints (especially young cap) . 7
6. Associated with western conifers (mainly Douglas-fir and pine) .8
6. Associated with eastern conifers (mainly hemlock) L. subpurpureus (see L. rubrilacteus, p. 68)
7. Latex when present yellowish to brown or the color of grasshopper juice; found in eastern North
America and the Southwest .L. chelidonium (see L. indigo, p. 69)
7. Not as above.L. paradoxus & others (see L. indigo, p. 69)
8. Cap whitish to pinkish-brown; found in Southwest .. L. barrowsii (see L. rubrilacteus, p. 68)
8. Not as above; cap reddish-brown to orangish or tan.L. rubrilacteus, p. 68
9. Latex quickly becoming yellow upon exposure to air (usually within 45 seconds) . 10
9. Not as above (but latex may slowly stain yellow or stain wounded tissue yellow) . 15
10. Cap pinkish to pinkish-cinnamon to reddish or vinaceous L. vinaceorufescens & others, p. 74
10. Cap differently colored (white to yellowish, ochre, orange, etc.) . 11
11. Cap white to yellowish to dark ochre; margin of cap bearded with woolly hairs (at least when
young) or if not then associated with northern or mountain conifers.12
11. Not with above features . 13
12. Cap pale yellow to dark ochre; stalk usually scrobiculate. L. scrobiculatus, p. 73
12. Cap whitish when young (but often pale yellowish or yellow-stained in age); stalk not scrobi¬
culate or somewhat so only at maturity .L. resimus (see L. scrobiculatus, p. 73)
13. Cap orangish; associated with conifers or birch in northern and western North America . 53
13. Not as above; either differently colored or associated with hardwoods such as oak . 14
14. Cap orange to bright yellow-orange .L. croceus (see L. vinaceorufescens, p. 74)
14. Not as above .L. chrysorheus & others (see L. vinaceorufescens, p. 74)
15. Cap (and often stalk) with a velvety texture or appearance (use hand lens if unsure); cap gray
to smoky-brown to dark brown or black, not viscid and not very large . 16
15. Not as above (if velvety, then differently colored—white, orange, red-brown, etc.) . 17
16. Gills close to crowded; associated with western conifers.L.fallax, p. 77
16. Not with above features .L. lignyotus & others (see L.fallax, p. 77)
17. Latex drying purplish or stained wounded areas purplish to dull lilac (often slowly); cap not
whitish, or if whitish then also viscid or slimy when moist . 18
17. Not as above . 21
18. Cap distinctly hairy or fibrillose at or near the margin; cap yellow, ochre, buff, or dull orange
.L. representaneus & others, p. 75
18. Not as above; margin of cap naked . 19
19. Cap pale yellow; cap and stalk viscid when moist L. aspideoides {see L. representaneus, p. 75)
19. Not with above features . 20
20. Cap and stalk white and slimy, at least when young and moist; common under conifers in the
Pacific Northwest .L. pallescens (see L. uvidus group, p. 75)
20. Not as above; cap usually buff, gray, brown, purplish, etc. . . L. uvidus group & others, p. 75
21. Cap dark green to sordid greenish- or olive-brown (at least at center), sometimes honey-colored
at margin; stalk similarly colored; latex white; taste acrid; cap staining magenta in KOH 22
21. Not with above features . 23
66 RUSSULACEAE

22. Cap and stalk dark or dull greenish; associated with hardwoods (especially oak) in eastern
North America . L. atroviridis (see L. olivaceoumbrinus, p. 70)
22. Cap and stalk usually browner; found mainly with conifers L. olivaceoumbrinus & others, p. 70
23. Margin of cap distinctly bearded with woolly hairs, at least when young; cap sometimes whitish,
but usually pale pinkish to pinkish-orange (at least at center); taste very acrid; associated
with birch or sometimes aspen .L. torminosus & others, p. 73
23. Not as above; if margin bearded then cap differently colored or habitat different . 24
24. Gills well-spaced; cap whitish to pale grayish- or pinkish-tan or yellowish; latex and wounded
areas staining rosy-salmon to rusty-orange (especially gills and stalk base); found under
hardwoods in eastern North America (especially common in Southeast) L. subplinthogalus
24. Not as above . 25
25. Cap pinkish-gray to purplish-gray to vinaceous-gray or vinaceous-buff (or brown tinged with
these colors), not viscid; odor distinctly coconut-like; found mainly with birch (often in
parks) or alder; northern . L. glyciosmus
25. Not as above . 26
26. Gills pinkish to pale pinkish; cap viscid when moist, white or with pinkish to lavender stains;
taste acrid; associated with aspen, poplar, and willow .L. controversus, p. 70
26. Not as above (cap if whitish usually not viscid) . 27
27. Margin of cap with cottony tissue (at least when young); cap usually whitish when young, but
may discolor yellowish, tan, etc. in age .L. deceptivus & others (see L. piperatus, p. 71)
27. Margin of cap naked or bearded with hairs but not as above . 28
28. Cap predominantly whitish (but may discolor in age), not viscid; taste distinctly acrid (some¬
times extremely so!); found mostly with hardwoods in eastern North America. 29
28. Not as above (if cap whitish, then taste not acrid) . 31
29. Latex staining wounded tissue pinkish; especially common in Southeast.L. subvernalis
29. Not as above . 30
30. Cap and stalk minutely velvety; gills close to well-spaced L. subvellereus (see L. piper atus, p. 71)
30. Cap and stalk not velvety; gills crowded to very crowded.L. piperatus & others, p. 71
31. Cap whitish to olive-buff (often with darker and paler areas); latex white, scanty, staining
tissue reddish to pinkish-gray or pale yellow; taste not acrid; favoring pine L. pallidiolivaceus
31. Not as above . 32
32. Latex white and very copious, staining wounded tissue brown (often slowly); taste mild; cap
and/ or stalk minutely velvety, never viscid; cap white to yellow, orange, red-brown, etc. (never
grayish or olive); common under hardwoods in eastern North America. 33
32. Not as above (but may have some of above features) . 35
33. Cap and stalk whitish to buff or slightly yellowish .L. luteolus (see L. volemus, p. 78)
33. Cap and stalk tawny to orange-brown, reddish-brown, or brown . 34
34. Cap brown to reddish-brown or rusty-brown, often wrinkled; stalk typically 1-3 cm thick; odor
usually mild .L. corrugis (see L. volemus, p. 78)
34. Cap tawny, orange-brown, rusty-brown, or sometimes yellowish, not normally wrinkled;
stalk typically less than 1.5 cm thick; odor often fishy in age .L. volemus, p. 78
35. Cap with various shades of yellow, pale buff, ochre, copper, orange, or pinkish-orange, often
(but not always) concentrically zoned; taste very acrid (peppery or burning). 36
35. Not with above features (but may have some of them) . 40
36. Cap with various shades of yellow, ochre, dark ochre, buff, etc. 37
36. Cap oranger, pinker, or more coppery than above . 38
37. Cap margin with fibrils which are brown to grayish in age L.payettensis(see L. alnicola, p. 71)
37. Not as above .L. alnicola & others, p. 71
38. Margin of cap bearded when young L. subvillosus & L. psammicola (see L. torminosus, p. 73)
38. Not as above; margin of cap naked even when young . 39
39. Associated with northern conifers .L. olympianus (see L. alnicola, p. 71)
39. Associated with southern hardwoods (mainly oak).L. yazooensis (see L. alnicola, p. 71)
40. Cap cinnamon to butterscotch-brown or buffy-brown; cap and stalk slimy or viscid when moist;
taste acrid; common in eastern North America (rare or absent in West).L. affinis
40. Not with above features . 41
LACTARIUS 67

41. Cap and stalk viscid or slimy when moist; cap dark brown to smoky-brown to charcoal-gray
(without orangish, reddish, yellowish, or purplish shades), but may fade in age; associated
with conifers . 42
41. Not as above; cap viscid or dry; stalk not viscid and/or fruiting body differently colored . 43
42. Stalk more or less tan or paler .L. kauffmanii (see L. pseudomucidus, p. 77)
42. Stalk darker or grayer (colored like cap or slightly paler) . . L. pseudomucidus & others, p. 77
43. Taste distinctly to excruciatingly acrid (sometimes latently so); cap pinkish-tan to reddish-tan
to reddish-brown or brick-red (at least in age) or sometimes orangish . 44
43. Not as above; cap differently colored and/or taste mild to only slightly acrid . 48
44. Found under manzanita; cap small and viscid when moist L. manzanitae {see L. rufus, p. 79)
44. Not with above features .45
45. Associated with hardwoods in eastern North America . 46
45. Not as above; usually but not always associated with conifers . 47
46. Cap 6-20 cm or more broad, pinkish-tan to reddish-tan to brick-red (but whitish when young
or where covered by humus), not zoned concentrically; gills white to buff.L. allardii
46. Cap 5-15 cm broad, orange-brown to reddish-brown, often zoned concentrically; mature gills
reddish-brown to dull purple-brown .L. peckii
47. Cap dull or dark reddish to brick-red, vinaceous, or dark brown . . . . L. rufus & others, p. 79
47. Cap oranger or more brightly colored than above. 48
48. Odor distinctly fragrant (sweet), at least when dried or cooked (and often when fresh) ... 49
48. Odor not fragrant or sweet, even when dried or cooked . 52
49. Cap viscid, orange-brown to caramel-colored; odor coconut-like when dried; known only
from California .L. cocosiolens (see L. subflammeus, p. 79)
49. Not with above features . 50
50. Latex clear (colorless); cap 4-15 cm broad; associated with northern conifers and especially
common in boggy areas . ..L. helvus(-L. aquifluus)
50. Latex white or watery white; cap 2-10 cm broad; widespread in many habitats .51
51. Stalk solid or stuffed (but sometimes with a central hollow in age), typically 0.5-3 cm thick; cap
usually reddish but sometimes orangish; known only from California .L. rufulus, p. 82
51. Stalk often (not always!) hollow or partly hollow, especially in age, and usually slender (0.4-1
or occasionally 1.5 cm thick); cap usually burnt orange but sometimes redder, browner, or
tawnier; widely distributed (including California) .L. fragilis & others, p. 80
52. Cap small and olive-brown to dark grayish-brown or dark brown when young (but often tawny
to cinnamon in age); common under alder in N. Calif, and Pacific Northwest L. occidentalis
52. Not with above features . 53
53. Cap orange to reddish, pinkish, reddish-brown, or liver-colored . 54
53. Cap grayish to violet-gray, dingy brown, olive brown, or dark brown . 59
54. Cap viscid or slightly viscid when moist . 55
54. Cap not viscid, often with a dull (matte) appearance . 56
55. Cap scarlet to orange, orange-brown, or tawny .L. subflammeus & others, p. 79
55. Cap pinkish to reddish to reddish-brown or liver-colored .... L. subviscidus & others, p. 80
56. Cap and stalk minutely velvety, orange to orange-brown; gills widely spaced, pallid; latex
very copious; common under eastern hardwoods L. hygrophoroides (see L. volemus, p. 78)
56. Not as above . 57
57. Associated with oak; mainly but not exclusively southern . 58
57. Not as above; northern . L. alpinus, L. thejogalus, L. oculatus & others (see L. rufulus, p. 82)
58. Found in California .L. rufulus & others, p. 82
58. Found in eastern North America.L. subserifluus & others (see L. rufulus, p. 82)
59. Cap medium-sized to large (4 cm or more broad), at least slightly viscid when moist .60
59. Not as above; cap small and violet-gray to gray or ochre-gray, usually with small scales at least
in age; found in eastern North America, often on or near rotten wood .L. griseus
60. Latex often discoloring injured tissue slowly; gills often dingy yellowish or yellow-brown in old
age; found with hardwoods (mainly oak) L. argillaceifolius var. megacarpus & others, p. 76
60. Not as above .L. circellatus & others (see L. argillaceifolius var. megacarpus, p. 76)
68 RUSSULACEAE

Lactarius deliciosus (Delicious Milk Cap) Color Plate 2

CAP 5-16 cm broad, broadly convex or with a depressed center and an inrolled margin
when young, becoming depressed or shallowly funnel-shaped; surface viscid when moist
but soon dry, smooth, often zoned; color variable: dull orange to carrot-orange or orange-
brown, sometimes blotched with or entirely green; fading in age or dry weather to
brownish, gray, dull greenish-gray, or even yellowish; margin inrolled when young. Flesh
thick, brittle, orange to yellowish or greenish; taste mild or slightly bitter. LATEX very
scanty, bright carrot-orange (but in some forms slowly staining dark red when exposed),
eventually staining wounded or aged tissue greenish (within 5 hours). GILLS typically
bright to dull orange, but varying to yellowish or orange-buff, greenish where wounded;
adnate to decurrent, close. STALK 2-7 cm long, 1-2.5 cm thick, equal or narrowed at base,
soon dry, sometimes scrobiculate; rigid, hollow inage, frequently maggot-riddled, colored
like cap or paler. SPORE PRINT creamy yellowish-buff; spores 7-11 * 6-8 microns,
broadly elliptical to nearly round, with amyloid ridges.
HABITAT: Scattered to gregarious or in troops under conifers (pine, spruce, etc.),
common and widely distributed. It is abundant in our coastal pine forests in the late fall
and winter, generally after the major crop of look-alike L. rubrilacteus.
EDIBILITY: Edible, but not necessarily delicious. Several varieties or forms occur and
some are apparently better than others. Special treatment is required to overcome the
grainy texture and latent bitterness. Its abundance and distinctiveness, however, make
experimentation worthwhile. Some sources recommend slow cooking (e.g., baked in a
casserole), others insist it should be cooked rapidly in a frying pan with very little butter.
It is popular in Europe, and the Russians are fanatical about it—especially salted.
COMMENTS: This variable, cosmopolitan fungus strikes a discordant but colorful
note with its unlikely combination of pistachio-green and carrot-orange—even more so
when found in the company of the bright red Russula rosacea, as is so often the case in our
area. The carrot-colored latex separates it from L. rubrilacteus, and confusion with other
mushrooms is unlikely. The latex is so scanty as to often be non-existent, but the brittle
flesh and greenish stains on the cap, stalk, and/ or gills will identify it. The fruiting bodies
persist for a long time and are often completely hollowed out by maggots. Several varieties
of L. deliciosus have been described based on slight differences in color, staining reactions,
and microscopic characteristics; one variety tends to have an areolate (scaly-cracked) cap
in age. The Greeks were apparently fond of L. deliciosus, for it is depicted on a fresco from
Herculaneum (buried in 79 A.D.). Other species: L. thyinos, of northern bogs, has carrot-
orange latex but stains red, not green; in other respects it is quite similar. L. salmoneus is
a smallish but distinctive southeastern species with a whitish cap, bright orange gills,
flesh, and stalk (when fresh) and orange to salmon-orange latex. L. pseudodeliciosus is
buff or dingy colored with yellow-orange to rusty-orange latex. All of these are edible and
the latter two stain greenish in age or where wounded.

Lactarius rubrilacteus (Bleeding Milk Cap) Color Plates 3,6

CAP 4-14 cm broad, broadly convex with a depressed center and inrolled margin when
young, depressed or shallowly funnel-shaped in age; surface viscid when moist, smooth,
reddish-brown to orange, orange-brown, or tan, or often concentrically zoned with these
colors; duller and greenish-stained in age. Flesh thick, brittle, brownish to buff, reddish,
etc.; taste mild or slightly bitter. LATEX scanty, dark red (but occasionally orange-red
in old specimens), slowly staining wounded areas greenish. GILLS adnate to slightly
decurrent, close, reddish or dull purplish-red, or tan with a dark reddish sheen; greenish
where wounded. STALK 2-6 cm long, 1 -2.5 cm thick, equal or narrowed below, firm, rigid,
LACTARIUS 69

hollow, colored like cap or paler, sometimes scrobiculate. SPORE PRINT pale yellowish
or buff; spores7.5-10 * 6-8 microns, broadly elliptical to nearly round, withamyloid ridges.
HABITAT: Scattered or in large troops under conifers throughout the West; associated
in our area with Douglas-fir and abundant in the fall and early winter. Where pines pre¬
dominate it is largely supplanted by L. deliciosus, and where pine and Douglas-fir grow
together, the two milk caps often mingle. In Europe, L. sanguifluus (which may be the
same as L. rubrilacteus) is mycorrhizal with pine.
EDIBILITY: Edible, but not pleasing to everyone because of its granular texture; usually
better, however, than L. deliciosus (see comments on edibility of that species).
COMMENTS: The dark red latex is the telltale trait of this handsome fungus, which for
many years has been known to fungophiles as L. sanguifluus. In dry or very wet weather
the latex may be absent, but in these conditions the greenish stains are usually quite
pronounced. In fact, as with L. deliciosus, weathered fruiting bodies may be entirely
green, and mature specimens often have tiny, aborted green “buttons” at their bases. Be
sure not to confuse this species with the similarly colored L. vinaceorufescens and
L. chrysorheus—they often grow with L. rubrilacteus, but have a white latex that quickly
turns yellow. Other species: L. barrowsii has dark red latex, but its cap is much paler
(whitish to pinkish-brown) and it occurs mainly with ponderosa pine. Another similar
edible species, L. subpurpureus of eastern North America, has wine-red latex and a wine-
red to silvery cap, and favors hemlock. See also L. paradoxus (under L. indigo).

Lactarius indigo (Indigo Milk Cap) Color Plate 4

CAP 4-15 cm broad, convex or centrally depressed with an inrolled margin when young,
usually depressed in age; surface smooth, viscid when moist, then dry; indigo-blue when
fresh but fading to grayish- or silvery-blue, sometimes with greenish stains; often zoned
concentrically. Flesh pallid to bluish, brittle, slowly staining greenish; taste mild to slightly
bitter-acrid. LATEX indigo-blue (bright dark blue), scanty, slowly staining wounded
tissue greenish. GILLS adnate to slightly decurrent, close, indigo-blue becoming paler in
age. STALK 2-6 cm long, 1-2.5 cm thick, indigo-blue to silvery-or grayish-blue, equal or
narrowed at base, rigid, hollow in age, soon dry. SPORE PRINT creamy-yellowish;
spores 7-9 * 5.5-7.5 microns, elliptical to nearly round, with amyloid warts and ridges.
HABITAT: Scattered to gregarious in summer and fall, mostly in oak and pine woods;
found throughout southern and eastern North America, but most common along the Gulf
Coast and in Mexico. I have found it in Arizona under ponderosa pine. If it should turn up
in southern California it would certainly be a most welcome addition to our fungus flora.
EDIBILITY: Edible and very good—superior, at least, to the other greenish-staining
milk caps, such as L. deliciosus. In Mexico it is sometimes sold in farmer’s markets.
COMMENTS: The overall blue to blue-gray color and bright blue latex make this one of
the safest and most memorable of all agarics. No other milk cap has blue latex, let alone
a bluish fruiting body. Other species: L. paradoxus, common under pine in eastern North
America (especially the S outh), often has a bluish-tinged cap when young, but has reddish-
brown to purple-brown latex and gills. Another species, L. chelidonium, has a yellowish
to dingy yellow-brown to bluish-gray cap and yellowish to brown latex “the color of
grasshopper juice.” It occurs commonly in the eastern United States and Southwest. L.
hemicyaneus is a medium-sized to large species which usually has bluish flesh in the cap
and orange to red-orange flesh in the base of the stalk. It may possibly occur in southern
California, according to Greg Wright and Paul Harding. All of the above species are
edible despite their tendency to stain green in age.
70 RUSSULACEAE

L actarius olivaceoumbrinus (Toadskin Milk Cap)

CAP 3-12 cm broad, convex with an inrolled margin becoming plane or shallowly
depressed; surface viscid when moist, typically a mixture of dark olive, sordid olive-
brown, and olive-buff, often spotted or zoned concentrically, but zones fading in age. Flesh
thick, dingy olive, brittle, taste very acrid. LATEX copious, white, becoming greenish-gray
(sometimes very slowly). GILLS adnate to decurrent, crowded, pallid becoming spotted or
colored greenish or olive-gray. STALK 4-8 cm long, 1-3 cm thick; solid, becoming hollow
in age; viscid when wet, then dry; rigid, colored more or less like cap; usually scrobiculate.
SPORE PRINT pale buff; spores 7-10 x 6-9 microns, elliptical to nearly round, with
amyloid warts and ridges. Cap surface staining magenta in KOH or ammonia.
HABITAT: Solitary or scattered or in small groups on ground under conifers in late
summer and fall, Pacific Northwest and northern California, occasional.
EDIBILITY: Unknown. The sordid appearance and acrid taste are major deterrents.
COMMENTS: This species and its close relatives are among our most distinctive milk
caps—easily recognized by their overall dingy greenish to murky olive-brown color and
copious, acrid white latex. I know very few people who would call them beautiful, but
that’s precisely what they are—beautiful—in a grotesque sort of way that only a thoroughly
jaded fancier of the fleshy fungi can appreciate. (For some reason, they remind me of toads
—see Color Plate 1.) Other species: L. sordidus is a similar species with a somewhat
browner cap and smaller spores; it also occurs in the Pacific Northwest as well as in eastern
North America. L. atroviridis (COLOR PLATE 1) has a mottled dark greenish cap and
stalk and grows under hardwoods (especially oak) in eastern North America. Neither is
edible. The nameL. necator(& European species) was previously applied to this group.

Lactarius controversus (Poplar Milk Cap)

CAP (5) 7-20 cm broad, broadly convex becoming depressed; surface viscid when moist,
then dry, white, or whitish with lavender to pinkish stains; margin at first inrolled. Flesh
thick, firm, brittle, white; taste slowly acrid(burning). LATEX white, unchanging. GILLS
adnate to slightly decurrent, crowded, narrow, pinkish to creamy-pink. STALK 2.5-7 cm
long, 1-3 cm thick, equal or tapered downward, white, sometimes spotted; hollow in age;
rigid. SPORE PRINT creamy to pale pinkish; spores 6-7.5 * 4.5-5 microns, elliptical, with
amyloid warts and ridges.

Lactarius controversus has pinkish gills, white latex, acrid taste, and inrolled cap margin when young.
LACTARIUS 71

HABITAT: Scattered to gregarious under aspen, poplar, and willow; widely distributed.
It can be found in the aspen forests of the Sierra Nevada in the late summer and fall. I
have seen large fruitings in New Mexico in the late summer.
EDIBILITY: To be avoided due to the acrid taste.
COMMENTS: This handsome milk cap is one of several large, whitish species with white,
unchanging latex and a very peppery taste. The viscid cap when moist and pinkish-tinted
gills plus the association with aspen, poplar, and willow distinguish it from the other
large white Lactarii (see L. piperatus), while the presence of a latex places it in Lactarius.

Lactariuspiperatus (Peppery White Milk Cap)

CAP (4) 6-16 cm broad, broadly convex and usually depressed centrally, becoming broadly
vase-shaped in age; surface more or less smooth, dry, unzoned, white to creamy-white,
but often developing dingy buff or tan stains in age; unpolished; margin naked. Flesh thick,
crisp, brittle; odor mild, taste extremely acrid. LATEX white, copious, unchanging or
drying yellowish (or in one variety staining wounded tissue dingy greenish). GILLS adnate
to decurrent, narrow and very crowded, white to creamy, often forked. STALK 2-8 cm
long, 1-3 cm thick, equal or tapered toward base, dry, white, smooth or with a whitish
bloom, not scrobiculate. SPORE PRINT white; spores 4.5-7 * 5-5.5 microns, elliptical
to nearly round with inconspicuous amyloid warts and ridges.
HABITAT: Solitary to widely scattered or gregarious on ground under hardwoods,
common in the summer in eastern North America. It has also been reported from the west
coast, but I have not seen it west of Minnesota.
EDIBILITY: Not recommended. In Russia and Scandinavia it is eaten, along with other
acrid milk caps, after parboiling or pickling. However, it is rather difficult to digest and
may even be poisonous if not properly prepared.
COMMENTS: The dull dry white cap, very crowded narrow gills, extremely peppery taste,
and copious white “milk” form a distinctive set of characters. It has much the aspect of
Russula brevipes, but of course that species lacks a latex. There are several similar,
medium-sized to large whitish, acrid milk caps with white latex, including: L. neuhoffii,
with larger spores and only moderately crowded gills that are sometimes pale pinkish-buff
at maturity; L. subvellereus, with minutely velvety cap and stalk, and close to well-spaced
gills; andL. tomentoso-marginatus andL. deceptivus, with a whitish cap that becomes dull
tan or brownish in age and has conspicuous cottony tissue on the margin when young. All
of these are common in eastern North America under hardwoods; the latter also grows
with conifers. None have the slightly viscid cap and pinkish gills of L. controversus. Other
species: L. pseudodeceptivus mimics L. deceptivus, but grows with northwestern conifers.

Lactarius alnicola (Golden Milk Cap) Color Plate 5

CAP 5-14 (20) cm broad, convex with a central depression and conspicuously inrolled
margin when young, depressed or broadly funnel-shaped in age; surface viscid when
moist, usually zoned concentrically with various shades of ochre and pale yellow, but
sometimes nearly evenly colored; margin naked or slightly hairy but not bearded. Flesh
thick, brittle, crisp, whitish; taste very acrid. LATEX white, unchanging or very slowly
yellowing or staining wounded tissue yellowish. GILLS crowded, adnate to decurrent,
whitish when young becoming buff or ochraceous-toned in age; wounded areas stained
yellowish to yellow-brown. STALK 2-6 cm long, 1.5-3 cm thick, equal or with narrowed
base, hard, usually hollow in age; pallid or tinged cap color, often (but not always)
scrobiculate (with darker yellow to ochraceous spots). SPORE PRINT whitish to
yellowish; spores 7.5-10 * 6-8 microns, elliptical, with amyloid warts and ridges.
Lactarius alnicola has a golden zoned cap and peppery white latex. This is our oak-loving variety,
which has a latently acrid taste and bald cap margin. See color plate for conifer-loving form.

HABITAT: Scattered to gregarious under spruce and other conifers (despite the species
epithet, which implies alder) in the Rocky Mountains and Pacific Northwest in the late
summer and fall, and also abundant (a slightly different form—see comments) in our live
oak woodlands in the fall and winter. Still another variant occurs in coastal sand dunes
with willow and bush lupine.
EDIBILITY: Not edible. The excruciatingly peppery taste is a formidable deterrent.
COMMENTS: The overall yellowish-ochre color and zoned, non-bearded cap plus
the frequently scrobiculate stalk, acrid taste, and white latex typify a very confusing group
of milk caps which have traditionally passed under the European names of L. insulsus and
L. zonarius. There is still doubt, however, as to just what those species are. The common
member of this group in central and southern California differs from typical L. alnicola in
its mycorrhizal mate (oak), somewhat smaller size and paler cap (yellowish-buff),
frequently short, off-center stem, and latently acrid taste. Its latex usually stains
surrounding tissue dingy yellow but does not turn yellow itself. Forms are encountered,
however, in which the latex does yellow slowly, and forms in which it discolors tissue buff or
grayish-buff, thus muddling the picture so thoroughly that I begin to yearnfor the baseball
season. (Ah, the simplicity, security, and symmetry of baseball, where each participant has
a fixed name, number, and position, and a unique and indisputable set of statistics and
characteristics.) Other species with a more or less zoned, non-bearded cap and acrid white
latex include: L. olympianus, a common western conifer-lover, typically with an oranger
cap and non-scrobiculate stalk; and two oak-loving easterners,L. psammicolaf glaber,
with a yellowish cap and latex that stains wounded tissue buff to pinkish-cinnamon, and
L. yazooensis, a southern species with an orangish cap. Similar species that are bearded
are listed under L. torminosus and L. scrobiculatus. They include L. subvillosus, a com¬
mon local species whose cap is often naked in age, but pinker or oranger than that of L.
alnicola. Also worth mentioning is L. payettensis, a western conifer-lover with white
scanty latex that yellows slowly and brownish to grayish fibrils on the cap margin. N one of
the above species should be eaten, at least until they are better known.

72
LACTARIUS 73

Lactarius torminosus (Bearded Milk Cap) Color Plate 7

CAP 4-12 cm broad, convex with a central depression and strongly inrolled margin when
young, shallowly depressed in age; surface viscid when moist, yellowish-buff to pinkish-
buff to whitish, the center usually pinkish to pinkish-orange when fresh, or sometimes pale
pinkish to pinkish-orange throughout; margin bearded with a dense white mat of soft,
woolly hairs (while inrolled) that may mimic a veil; hairs sparse or even absent inage. Flesh
thick, firm, brittle, white or tinged pinkish; taste very acrid. LATEX often scanty, white
and typically unchanging (but staining gills yellowish in var. nordmanensis). GILLS white
to yellowish-tan or developing a pinkish tinge, crowded, narrow, adnate to slightly
decurrent. STALK 2-7 cm long, 0.5-1.5 cm thick, rigid, equal or with a narrowed base,
firm, dry, often hollow in age; colored like cap or paler, sometimes with dingy ochre spots.
SPORE PRINT creamy-white; spores 7-10 x 6-8 microns, elliptical, with amyloid ridges.
HABITAT: Scattered to gregarious under or near birch or occasionally aspen; common
and very widely distributed. In our area this species and L. pubescens(see comments) are
common after the first fall rains on lawns with planted birch trees (often in the company of
Paxillus involutus). The mycelium is probably imported on the roots of birch saplings.

EDIBILITY: Not recommended, as it is indigestible or even poisonous unless thoroughly

cooked. However, in Russia and Scandinavia it is collected in large quantities and pickled,
and in Norway it is roasted and added to coffee.
COMMENTS: The bearded, pinkish-tinged cap, white latex, acrid taste, and growth with
birch are the trademarks of this attractive Lactarius. Buttons with an inrolled margin are
reminiscent of Paxillus involutus (another birch-lover), but paler in color. L. pubescens is
practically identical to L. torminosus but has smaller spores(6-8 microns long) and is more
apt to have a whitish cap (see photo on p. 41). It is common with birch in parksand gardens
as well as in forests, and is often mistaken for L. torminosus. One collection I made had
greenish bands on the gills and stalk just like in Russula brevipes var. acrior. Other similar
species: L. subvillosus is common in our tanoak-madrone woodlands. Its pinkish-orange
to orange cap is bearded when young but often naked in age, leading to confusion with L.
alnicola, which has a yellower cap. L. psammicola has an orangish, zoned, bearded cap
but grows under eastern hardwoods. For bearded yellow-stainers, see L. scrobiculatus.

Lactarius scrobiculatus (Scrobiculate Milk Cap)

CAP 4-15 (20) cm broad, convex to plane with the center usually depressed, in age often
vase-shaped; surface viscid when moist, smooth or scaly in age, unzoned or faintly zoned,
pale yellow to yellowish to bright ochre, the center sometimes darker; margin inrolled
when young and typically bearded with hairs (but hairs absent or inconspicuous in var.
montanus and var. pubescens). Flesh thick, firm, brittle, whitish but staining yellow when
exposed; taste variable: strongly acrid to mild. LATEX copious or scanty, white, quickly
staining yellow when exposed. GILLS adnate to slightly decurrent, whitish to pale or dull
yellow, close. STALK 3-11 cm long, (1) 2-4 (5) cm thick, equal or narrowed at base, firm,
dry, hollow in age; white to yellowish, usually scrobiculate (i.e., pitted with large, glazed,
darker or brighter yellow to honey-colored spots). SPORE PRINT whitish to creamy or
yellowish; spores 6-10 x 5-7.5 microns, broadly elliptical, with amyloid warts and ridges.
HABITAT: Solitary, scattered, or in groups under northern and mountain conifers;
widely distributed. Its various varieties are common in the summer and fall in the Sierra
Nevada, northern California, and other parts of the West, but absent in our area.
EDIBILITY: Not recommended; yellow-staining milk caps should be avoided.
COMMENTS: This robust but variable milk cap can have a mild to strongly peppery
(acrid) taste and prominently bearded to virtually bald cap margin. However, it can
Lactarius vinaceorufescens (see description below) has white latex that quickly stains yellow when
exposed. It is variable in other respects and intergrades with L. chrysorheus, another yellow-stainer.

be recognized by its overall pale yellow to dark ochre color, scrobiculate stalk, and white
latex that quickly turns yellow or stains exposed tissue yellow. The latter feature distin¬
guishes it from L. alnicola, which stains yellow slowly or not at all. Other species: L.
resimus is a bearded, yellow-staining species whose stalk is scrobiculate only in age if at
all. Its cap is white when young but often pale yellowish in age. It favors birch, aspen,
alder, manzanita, and conifers. Also see L. payettensis (under L. alnicola).

Lactarius vinaceorufescens (Yellow-Staining Milk Cap)

CAP 3-7 (9) cm broad, broadly convex with incurved margin, then plane or depressed;
surface dry to slightly viscid, smooth, cinnamon-buff to pinkish, reddish-cinnamon, or
dark reddish, often darkening to vinaceous-brown in old age; often faintly zoned or with
darker watery spots; margin not bearded. Flesh brittle, staining yellow when cut; taste mild
to somewhat bitter or acrid. LATEX white, quickly turning sulfur-yellow when exposed
(within 45 seconds). GILLS adnate to slightly decurrent, close, pallid when young, then
tinged cap color and eventually aging or staining dark reddish. ST ALK 3-7 cm long, 1 -2.5
cm thick, equal, pallid or colored like cap but usually paler, often stained dark reddish in
age; smooth, not scrobiculate; base often with hairs. SPORE PRINT white to yellowish;
spores 6.5-9 * 6-7 microns, broadly elliptical to round, with amyloid warts and ridges.
HABITAT: Scattered or in groups or troops under both hardwoods and conifers; widely
distributed. It is often abundant in our area from fall through early spring, particularly
with manzanita, oak, Douglas-fir, and pine (often mingling with L.fragilis, L. rubrilacteus,
Russula emetica, and R. cremoricolor). L. chrysorheus (see comments) is also common.

EDIBILITY: Reportedly poisonous—all yellow-staining milk caps should be avoided.

COMMENTS: This ubiquitous milk cap superficially resembles L. rubrilacteus, L. rufus,
L. subviscidus, L. fragilis, and L. rufulus, but is easily distinguished by the prompt
yellowing of the latex when exposed to the air. The rate at which it yellows depends on the
moisture content and age of the mushroom, but it normally occurs in 5-30 seconds. L.
vinaceorufescens has been confused in the past withL. chrysorheus, widespread and also
common in our area, which has a somewhat paler (pallid to yellowish-cinnamon to pale
pinkish), often zoned cap and gills and stalk that do not discolor as much in age. Other

74
LACTARIUS 75

species with yellow-staining latex include: L. maculatipes and L. croceus, found under
hardwoods in eastern North America, the first with a whitish to creamy-yellow, often
spotted cap and slimy stalk when fresh, the latter with a bright saffron to yellow-orange or
orange, viscid cap; and L. xanthogalactus of California, probably the same as L. chry-
sorheus or L. vinaceorufescens. For other yellow-staining species, see L. scrobiculatus.

L actarius uvidus group (Purple-Staining Milk Cap)

CAP 3-10 (12) cm broad, broadly convex to plane or shallowly depressed; surface smooth,
viscid to slimy when moist, pallid becoming grayish, lilac-gray, or pale lavender-brown,
sometimes obscurely zoned; margin naked. Flesh thick, white, staining lilac slowly when
wounded; taste mild to slowly bitter or acrid. LATEX white or creamy, staining wounded
areas lilac or dull purple. GILLS adnate to slightly decurrent, close, white to yellowish;
wounded areas staining purplish, then eventually dingy tan. STALK 3-8 cm long, 1-2 cm
thick, more or less equal, smooth, viscid when moist but soon dry, rigid, pallid or sometimes
tinged cap color, often ochraceous-stained toward base. SPORE PRINT yellowish-white;
spores 8-12 ><7-8 microns, elliptical, with amyloid warts and ridges. Cap surface staining
green in KOH (potassium hydroxide).
HABITAT: Solitary, scattered, or in small groups in woods; widely distributed. In our
area this species and its close relatives (see comments) are fairly common in mixed woods
in the late fall and winter, but seldom fruit in quantity.
EDIBILITY: Said to be poisonous. All purple-staining milk caps should be avoided.
COMMENTS: The tendency of the latex to stain wounded tissue purple typifies this
species (see photo on p. 897) and its close relatives. A stouter, conifer-loving version,//.
uvidus var. montanus, has a darker, somewhat browner, nearly dry cap and resinous taste.
Other purple-stainers include: L. pallescens, with a slimy or viscid whitish cap and stalk,
common under conifers in northern California and the Pacific Northwest; L. califor-
niensis, an acrid-tasting species whose purplish to brownish-gray cap often has yellowish
tinges or stains; L. cascadensis, a large northwestern species with dull brownish gills in age,
favoring swampy areas (under alder, etc.); L. maculatus of eastern North America,
with a distinctly zoned cap and acrid taste; and L. subpalustris, larger, with a grayish to
dingy brown, often spotted cap that does not stain green in KOH, plus a mild taste. For
purple-staining species with a yellowish to buff cap, see L. representaneus.

Lactarius representaneus (Purple-Staining Bearded Milk Cap)

CAP (4) 6-20 cm broad, broadly convex with a central depression and inrolled margin,
becoming broadly depressed or vase-shaped in age; surface viscid when moist, sometimes
zoned concentrically, pale yellow to golden-yellow to orange-buff, often developing rusty
and sometimes purple stains; smooth at center but coarsely hairy (fibrillose) toward the
margin, which is bearded with woolly yellowish hairs when young. Flesh thick, brittle,
white, staining dull lilac or purplish slowly where cut or bruised; taste slowly bitter or acrid.
LATEX copious, white or creamy, drying or slowly staining wounded tissue dull purple or
lilac. GILLS adnate to decurrent, close, buff to dull ochre or orange-spotted, with lilac
or purplish stains where bruised. STALK 4-12 cm long, 1-3 (4.5) cm thick, equal orthicker
below, hard, whitish or more often colored like cap, usually prominently scrobiculate,
sometimes lilac-stained; hollow or stuffed. SPORE PRINT whitish to yellowish; spores
8-12 x 6.5-9 microns, broadly elliptical, with amyloid warts and ridges.
HABITAT: Solitary to scattered or often gregarious onground under northern conifers—
especially spruce and fir; widely distributed but not found in our area. I have seen it under
Engelmann spruce in Colorado and New Mexico in August, and under Sitka spruce in
northern California in November. It is said to be abundant in Alaska.
76 RUSSULACEAE

EDIBILITY: Not recommended. The taste is not appealing and it may be poisonous.
COMMENTS: This often large, impressive milk cap is easily identified by its yellowish
cap with the hairy margin, and purple-staining latex. It resembles L. scrobiculatus and
L. alnicola in appearance, but those species have unchanging or yellow-staining latex.
Other purple-staining species (see L. uvidus) are differently colored and do not have a
bearded cap. Similar species include: L. speciosus, with a dull buff to tan cap and hairy
margin, common under hardwoods in eastern North America (especially the South); and
L. aspideoides, with a viscid, pale yellow cap that is not bearded. Neither should be eaten.

Lactarius argillaceifolius var. megacarpus (Vulgar Milk Cap)

CAP 7-18 (27) cm broad, broadly convex-depressed with an inrolled margin when young,
plane to depressed in age; surface viscid, drab grayish, violet-gray, or sometimes brownish
tinted with violet, often fading in age to dingy tan or grayish-buff; sometimes obscurely
zoned or with rusty-ochraceous spots. Flesh thick, pallid; taste mild or somewhat acrid.
LATEX creamy-white, unchanging, but slowly staining wounded areas dingy brownish.
GILLS pallid becoming dingy yellowish or buff, often darker (dingy yellow-brown) in old
age, slowly staining brownish or grayish where bruised; adnate to slightly decurrent, close.
ST ALK 6-15 cm long, 2-6 cm thick, firm, rigid, slightly viscid or dry; white to buff or tinted
cap color, at times with ochre or yellow-brown stains. SPORE PRINT whitish to buff;
spores 7-11 * 6-9 microns, elliptical to nearly round, with amyloid warts and ridges.
HABITAT: Solitary, scattered, or in small groups in humus under hardwoods (live oak,
tanoak, madrone), fairly common in our area in the late fall and winter. It is known only
from the Pacific Coast, but variety argillaceifolius {see comments) is widespread.
EDIBILITY: Possibly poisonous; to be avoided.
COMMENTS: The robust dimensions, dingy tan to grayish color, and white latex are the
distinguishing features of this undistinguished mushroom. It is our largest and least
attractive Lactarius. The taste is rather variable, but generally the latex is at least somewhat
acrid while the flesh may be mild. The violet tints that sometimes pervade the cap are
reminiscent of L. uvidus (with which it sometimes grows), but that species stains purple
instead of dingy brown. L. argillaceifolius var. argillaceifolius is smaller and browner than
var. megacarpus, and has a tendency to become dingy brown or yellow-brown overall in
age. It is a characteristic summer mushroom in the hardwood forests of eastern North
America. Other species: L. caespitosus is similar, but has a gray to olive-brown cap and
grows under western conifers; L. vietus is also similar but favors birch and northern coni-

Lactarius argillaceifolius var. megacarpus can be told by its relatively large size, dingy color, and
white latex that slowly stains the gills brownish. For a close-up of the latex, see photo on p. 63.
Lactariuspseudomucidus, a conifer loving milk cap with a dark cap and stem that are slimy when wet.

fers and often has a lilac- or vinaceous-tinged cap;L. cinereus has a gray to olive-gray cap,
non-staining gills and stalk apex often tinged pale pinkish, and grows with beech; L. cir-
cellatus favors birch and northern conifers, but has non-staining gills and a brownish-gray
to bluish-gray or lavender-tinged cap, and zebra-like amyloid stripes on the spores. For
years L. argillaceifolius has been called//, trivialis, a similar slimy-stalked species whose
gills discolor scarcely if at all; it occurs in the Pacific Northwest, usually under conifers.

Lactarius pseudomucidus (Slimy Milk Cap)

CAP 3-8 (10) cm broad, broadly convex becoming plane or depressed; surface smooth,
viscid (with a thick layer of slime when moist), evenly colored blackish-brown to dark gray;
not zoned; margin naked. Flesh grayish, thin, fragile, taste slowly acrid. LATEX white,
unchanging, but may stain gills tan to brownish, gray, or olive-gray. GILLS adnate to
decurrent, white or tinged gray, sometimes discolored (brownish, etc.) in age. STALK
4-10 cm long, 0.5-1 cm thick, fragile, smooth, without spots, evenly colored like cap or paler
(gray); very slimy when wet, usually thicker below. SPORE PRINT white; spores 7-9 * 6-7
microns, broadly elliptical, with amyloid ridges.
HABITAT: Scattered to gregarious under conifers in late summer and fall; known only
from northern California and the Pacific Northwest, common.
EDIBILITY: Unknown. Only the most ardent slippery jack lover would fail to be
deterred by the copious layer of slime coating the cap and stem.
COMMENTS: The dark grayish-brown cap and stalk under a thick layer of slime (when
fresh) and the white latex are usually enough to distinguish this glutinous woodland
mushroom. It is slimmer and slimier than either L. argillaceifolius or L. trivialis, and
darker besides. It has been called L. mucidus, but that species, which is commoner in
eastern North America, has a pale cap margin and only slightly viscid stalk. Another
look-alike, L. glutigriseus, has a slightly viscid stalk that is paler at the apex and base.
L. kauffmanii, also frequent in the Pacific Northwest and sometimes found with L.
pseudomucidus, is similar but usually larger and stouter, with a distinctly tan or paler
(not gray) stalk. Also see the species listed under L. argillaceifolius var. megacarpus.

Lactarius fallax (Velvety Milk Cap) Color Plate 9

CAP 2.5-7 (9) cm broad, convex to plane (often with an umbo) when young, often shallow¬
ly depressed in age; surface dry and more or less velvety, often wrinkled toward the center;
evenly dark brown to nearly black, not zoned; margin often scalloped. Flesh rather thin,
brittle, whitish, taste mild or slightly acrid. LATEX white, usually copious, slowly stain¬
ing wounded tissue dull reddish. GILLS adnate to slightly decurrent, narrow, crowded,
white tocreamy-buff(butedgesdarkbrowninoneform).STALK2.5-7 cmlong,0.5-1.5 cm
78 RUSSULACEAE

thick, rather slender, more or less equal, colored like cap or somewhat paler; dry and
unpolished or velvety. SPORE PRINT yellowish; spores 9-12 * 8-11 microns, more or
less round, with amyloid warts and ridges.
HABITAT: Scattered or in small groups under conifers (especially fir) or occasionally on
rotting wood; fairly common in the late summer and fall in the Pacific Northwest and
northern California. I have yet to find it in our area.
EDIBILITY: Unknown.
COMMENTS: This species is one of several milk caps with a brown to nearly black,
velvety cap. Though not good edibles, they are notable for their beauty, and therefore likely
to attract the attention of even the casual collector. L.fallax is the most common member
of the group in the West, and is easily distinguished from the similarly colored L. pseudo-
mucidus, by its dry, velvety rather than smooth, slimy cap and stem. There are several
closely related species found mainly in eastern North America, including: L. lignyotus,
with broader, more widely-spaced gills; L. gerardii, with very well-spaced (distant) gills;
L. fuliginellus, a hardwood-lover with close gills; and L. fumosus, with close gills and
a paler (smoky-brown) cap. None of these are worth eating.

Lactarius volemus (Weeping Milk Cap; Bradley)

CAP 5-10 (13) cm broad, convex becoming plane or depressed, or sometimes umbonate;
surface dry, minutely velvety to nearly smooth, golden-tawny to orange-brown or rusty-
orange (or pale yellowish in var. flavus). Flesh thick, firm but brittle, whitish; odor often
becoming fishy in age; taste mild. LATEX white or creamy, very abundant, slowly staining
brown or staining wounded tissue brown. GILLS close, white to creamy or sometimes pale
tan, adnate to slightly decurrent. STALK 4-12 cm long, 0.8-1.2 (2) cm thick, equal or
tapered downward, orange-brown to tawny (colored like cap or slightly paler), dry,
unpolished, minutely velvety to nearly smooth; not scrobiculate. SPORE PRINT whitish;
spores 7.5-10 x 7.5-9 microns, round or nearly round, with amyloid ridges (reticulate).
HABITAT: Widely scattered to gregarious on ground under hardwoods (especially oak)
or in mixed woods; common in the summer and early fall in eastern North America.
EDIBILITY: Edible and delicious when properly cooked! Like most milk caps it has a
slightly granular texture that displeases some people, but the flavor is excellent; slow
cooking is best. L. corrugis and L. hygrophoroides (see comments below) are equally
good if not better. All three species are usually free of maggots.
COMMENTS: This species is one of a trio of tasty milk caps (see below) that rank among
the best edible mushrooms of eastern North America. To my knowledge they are not found
in the West, but are to be looked for in southern Arizona, where several so-called “eastern”
mushrooms occur (e.g., Strobilomyces floccopus). L. volemus is easily recognized by its
dry, tawny to orange-brown cap and stalk which are often minutely velvety, its mild taste,
and extremely copious white milk that slowly stains brown. The latex is so copious that
the slightest nick of the gills will cause them to “weep,” i.e., exude a stream of milky
droplets. The fishy odor that often develops after it is picked or as it is cooked is also
distinctive, but in no way reflects or affects its flavor. Also common under hardwoods in
eastern North America are three similar species with copious white latex, mild taste, and
minutely velvety cap and/ or stalk: L. hygrophoroides is a beautiful, similarly colored or
oranger species with well-spaced gills, a mild odor, and latex that does not stain brown;
L. corrugis (COLOR PLATE 8) is larger and more robust, with a frequently wrinkled,
dark brown to reddish-brown to rusty-brown cap, whitish to slightly yellowish or
cinnamon-tinged gills, brown-staining latex, and a (typically) mild odor;L. luteolus has
a white to buff or yellowish cap, brown-staining latex, and a frequently fishy odor. All
of the above species are edible. See p. 79 for a photograph of L. volemus.
Left: Lactarius volemus, a delectable species with a dry cap and copious white latex. Note how stature
is more slender than that of the closely related L. corrugis (Color Plate 8). Right: Lactarius sub-
flammeus has a viscid orange cap and white latex. Note the rather long, slender stem.

Lactarius subflammeus (Orange Milk Cap)

CAP 2-6 (7) cm broad, convex becoming plane or shallowly depressed; surface smooth,
viscid, scarlet when young soon fading to bright orange and eventually dull orange; not
zoned concentrically; margin naked. Flesh thin, fragile; odor mild, taste slowly acrid.
LATEX white, unchanging. GILLS adnate to decurrent, fairly close, whitish or colored
like cap but paler. STALK 4-9 cm long, 0.5-1.5 cm thick, usually rather long and thicker
toward the base; hollow, rigid but fragile, colored like cap; not viscid. SPORE PRINT
whitish; spores 7.5-9 * 6.5-7.5 microns, elliptical, with amyloid ridges and warts.
HABITAT: Scattered to gregarious under pine, spruce, and other conifers; found in
western North America in the late summer and fall and especially common in northern
California and the Pacific Northwest. I have not seen it south of San Francisco.
EDIBILITY: Unknown; best avoided until better known.
COMMENTS: The bright orange viscid cap, stalk usually longer than the width of the
cap, and white unchanging latex typify several similar species that have traditionally
passed under the name L. aurantiacus. They can be told from the candy cap (L.fragilis)
by their viscid cap and brighter color, plus their mild odor. L. luculentus is a very similar
species with a mild to bitterish taste, an orange to tawny cap that is only slightly viscid,
and gills that may stain brownish; it is also common in northern California and the Pacific
Northwest. L. cocosiolens, discovered in California by Andrew Methven, has a viscid
orange-brown to caramel-colored cap, mild taste, and copious white latex, and smells like
coconut when it dries. L. substriatus is one of several look-alikes with yellow-staining latex.

Lactarius rufus (Red Hot Milk Cap)

CAP 4-12 cm broad, broadly convex becoming plane or depressed; surface usually not
viscid; smooth, dark brick-red to bay-red or reddish-brown, not zoned; margin naked.
Flesh dingy reddish, rather fragile; odor mild, taste strongly—but often latently—acrid.
LATEX white, unchanging. GILLS crowded, adnate to slightly decurrent, whitish when
young, flushed reddish in old age. STALK 4-11 cm long, 1-1.5 cm thick, equal, rigid but
rather fragile, stuffed or hollow, dry, more or less colored like cap. SPORE PRINT pale
yellowish; spores 7.5-11 x 5-7.5 microns, elliptical, with amyloid warts and ridges.
HABITAT: Scattered to gregarious or in troops under conifers; widely distributed. It is
common in northern California and the Pacific Northwest from late summer through early
winter, but I have not seen it south of San Francisco. It is also abundant in northern
sphagnum bogs. Like L. deliciosus, its favorite mycorrhizal mates are pine and spruce.

79
80 RUSSULACEAE

EDIBILITY: Not recommended. Like several other peppery milk caps, it is harvested
and canned commercially in Scandinavia. However, North American variants have not
been thoroughly tested and it may be poisonous raw.
COMMENTS: This species occurs in droves under northern conifers. It can be told from
other reddish milk caps by its red-hot taste, unchanging white latex, and fondness for
conifers. Fresh young specimens are among the most acrid of all mushrooms. When you
taste one, take only a small bite, and remember: the burning sensation can be delayed!
Other species: L. manzanitae has a small (2-5 cm), viscid, orangish to coppery-red or
brick-red cap, strongly acrid taste, and unchanging latex; it occurs under manzanita in
California. L. hysginus var. americanus has a brown to dark vinaceous-brown, viscid
cap and viscid stalk; it grows under conifers. For similar species with a milder taste, see
L. subviscidus, L. subflammeus, and L. rufulus.

Lactarius subviscidus
CAP 1-4 (5) cm broad, shallowly depressed becoming broadly or more deeply depressed in
age; surface smooth, viscid to thinly slimy when wet, dark reddish or reddish-brown
to brick-red or paler (pinkish) in age, not normally zoned; margin sometimes faintly striate.
Flesh thin, tinged cap color, fragile; odor mild, taste mild to slightly acrid. LATEX white,
rather scanty, unchanging. GILLS close, adnate to decurrent, pinkish-buff to pinkish-
cinnamon, or darker in age. STALK 2-5 cm long, 4-8 mm thick, more or less equal, usually
hollow; smooth, not viscid, colored like cap or gills. SPORE PRINT white to yellowish;
spores 8-10 * 7-8 microns, broadly elliptical, with amyloid ridges.
HABITAT: Scattered to gregarious on ground or rotten wood under conifers (spruce,
pine, etc.) in the Pacific Northwest and California. It is fairly common in our area in the
fall and early winter under pine, Douglas-fir, and manzanita.
EDIBILITY: Unknown.
COMMENTS: This small species looks like a candy cap (L.fragilis or L. rufulus), but the
thinly viscid cap (often with debris stuck to it), mild odor, and white (rather than watery
white) latex distinguish it. The latex does not stain yellow as in L. vinaceorufescens, but
may slowly stain white paper yellow (overnight). The cap is not orange as in L. subflam¬
meus and not as acrid (peppery) as L. manzanitae (see L. rufus). Similar species include:
L. riparius, common under conifers, alder, and willow in seepage areas in the Sierra
Nevada, with a brownish-red to brick-red cap that is slightly viscid when moist, mild
taste, and copious unchanging white or watery-white latex; L. atrobadius, with a viscid
liver-colored to blackish-red cap and non-yellowing latex; and L. hepaticus, with a viscid
or dry cap and yellowing latex. For similar species with dry caps, see L. rufulus.

Lactarius fragilis (Candy Cap) Color Plate 10

CAP 2-7 cm broad, broadly convex or plane or sometimes with an umbo, becoming
slightly to broadly depressed in age; surface dry (never viscid!), often uneven or somewhat
wrinkled; usually colored burnt orange to cinnamon but sometimes reddish-brown;
sometimes darker toward center but not zoned concentrically; margin often wavy or
frilled. Flesh thin, fragile, tinged cap color; taste mild or slightly bitter; odor faintly fragrant
or pungent becoming strongly aromatic (like maple syrup) upon drying. LATEX white or
watery-white, unchanging, often scanty or absent. GILLS adnate becoming decurrent,
close, pale pinkish-cinnamon or tinged cap color (or sometimes more yellowish),
darkening to more or less capcolor in age, or dusted whitish with spores. STALK(2)4-9 cm
long, 0.4-1 (1.5) cm thick, usually rather slender; fragile, more or less equal, colored like
Lactarius fragilis. These mature specimens are rather regular in shape, but it is not uncommon for
the caps to have frilled edges. Color ranges from burnt orange(see color plate) to cinnamon to reddish-
brown and the stalk is usually hollow in age. Fragrant odor is also distinctive.

cap; often hollow in age, with hairs at base. SPORE PRINT white to pale yellowish;
spores 6-9 microns, more or less round, with amyloid warts and ridges (reticulate). Cap
and stalk tissue containing numerous nests of sphaerocysts.
HABITAT: Widely scattered to densely gregarious or in small clumps on ground in
woods (often along trails and in other damp places, sometimes on wood). It is apparently
confined to the Pacific Coast and Southeast, but the very similar L. camphoratus (see
comments) occurs elsewhere. In our area it is normally abundant from the late fall through
early spring, especially under oak but also with other trees (pine, Douglas-fir, etc.).

EDIBILITY: Edible and one of the very best of our late season mushrooms. Fresh speci¬
mens can be sauteed and used like any other mushroom; when chopped up and slowly
dried, however, their flavor becomes sweet and they are great in pancakes, cookies, on
cinnamon toast with sesame seeds, or even in ice cream! Fortunately, they are plentiful
enough to merit collecting in spite of their small size, but be sure of your identification!
A few poor souls, alas, are allergic to them.
COMMENTS: The most remarkable feature of candy caps is the sweet, persistent fra¬
grance they develop when cooked or dried. If just a few candy caps are sauteed, their odor
will permeate the entire house and linger for days. Stranger still, when eaten in quantity
they imbue one’s perspiration and bodily exudates with the unmistakable aroma of maple
syrup—thereby provoking some puzzled stares! Their odor has also been likened to that
of butterscotch, fenugreek, burnt sugar, and sweet clover. Fresh specimens may have
only a slight odor, in which case the mild taste, watery-white unchanging latex, and overall
burnt orange or “ferruginous” color separate them from most other mushrooms (but see
below). In dry weather the latex is often absent, leading to confusion with Laccaria laccata,
Clitocybe inversa, Rhodocybe nuciolens, Collybia and Cortinarius species, and various
other similarly colored mushrooms. Fortunately, the odor is usually quite pronounced
under those conditions. Be sure the latex does not stain yellow—L. vinaceorufescens often
mingles with it and can be practically the same color.
The typical variety of L. fragilis occurs in the Southeast and is slightly tawnier. The
western version, L. fragilis var. rubidus, may actually be a distinct species. For years it has
passed as L. camphoratus. That species, however, is found in northern coniferous and
hardwood forests and has a slightly redder, frequently nippled (umbonate) cap and non-
reticulate spores. Still another candy cap, L. rufulus, occurs in California under oak,
but tends to be larger, is often redder, usually has a thicker and firmer stem, and contains
few if any sphaerocysts in its tissue(see description of L. rufulus). For similar species with
a mild odor, see L. subviscidus and comments under L. rufulus.
81
Lactarius rufulus. This west coast native resembles L.fragilis, but differs microscopically and usually
has a solid, thicker stem (as shown here) and larger cap. Color is variable but usually reddish.

Lactarius rufulus (Rufous Candy Cap)

CAP 3-10 cm broad, broadly convex to plane or slightly umbonate, becoming wavy or
depressed; surface often uneven or somewhat wrinkled, not viscid, color variable: usually
dark red to reddish-brown or brick-colored, but sometimes oranger; evenly colored or
with darker areas; margin incurved at first. Flesh firm, whitish to buff or tinged cap color,
brittle; taste mild to faintly acrid, odor fragrant to mild or faintly pungent. LATEX white
or watery white, unchanging, often scanty or even absent. GILLS adnate to decurrent,
fairly close, pallid to cinnamon-buff, pinkish-buff, or tinged cap color (or yellower), often
darker red or reddish-brown in age, at least on the margins. STALK 4-12 cm long, (0.5)
0.8-3 cm thick, equal or slightly tapered, colored more or less like cap, firm, usually solid
or stuffed but sometimes partially hollow in age. SPORE PRINT creamy to yellowish;
spores 7-9 microns, round or nearly round, with amyloid warts and ridges(at least partially
reticulate). Nests of sphaerocysts scanty to completely absent in cap and stalk tissue.
HABITAT: Scattered to densely gregarious or tufted under oak, late fall through spring;
known only from California. In my experience it is not as common as L.fragilis in regions
of high rainfall (e.g., coastal central and northern California), but more common than
L.fragilis in drier areas (e.g., inland and southern California).
EDIBILITY: Edible and good, but not quite as fragrant or tasty as L.fragilis.
COMMENTS: This oak-loving candy cap tends to be redder and more robust than L.
fragilis, and often has a solid stem. The critical difference is microscope, however: the
absence or near absence of swollen cells (sphaerocysts) in the cap and stem tissue is highly
unusual for a Lactariusl L. rufulus can also be confused with L. rufus, but that species has
an acrid taste and favors conifers, while L. vinaceorufescens has yellowing latex.
There is a whole slew of very similar, reddish-brown to dull rusty-orange milk caps with
a mild to somewhat bitter taste and little or no odor. These are difficult to differentiate
without a microscope, and far too numerous to ennumerate. As a group they have passed
under the nameL. subdulcis, but of course are not the “true” L. subdulcis of Europe (if they
were, we would probably be calling them something else!). These species include: L. sub-
serifluus, with clear latex and orange-brown cap, the oak-loving eastern counterpart of
L. rufulus (it also lacks sphaerocysts in the cap tissue); L. thiersii, also lacking sphaero¬
cysts but much smaller (cap 1 -3 cm), known only from California;/,, oculatus, with a red-
brown to pinkish, often nippled,dry to slightly viscid cap, found in eastern Sphagnum bogs
and under conifers;/,, alpinus var. mitis, one of several species common under alder and
northern conifers or in seepage areas;/,. hepaticus(see comments under L. subviscidus)\
and L. thejogalus, with somewhat yellowing latex, occurring in swarms under birch.
L. herpeticus should also be mentioned. For similar viscid-capped species, see L. subvisci-
dus. None of these species should be eaten until better known.
 83

RUSSULA (The Russulas)

Medium-sized to large, mostly terrestrial woodland mushrooms. CAP plane or depressed at
maturity, viscid or dry, skin often peeling easily, margin often striate. Flesh brittle, usually white;
taste mild or acrid. Latex absent. GILLS attached or free, brittle, usually white to yellow, at least
when young (but may discolor in age). STALK central, rigid, brittle, breaking open cleanly like
chalk. VEILand VOLVA absent. SPORE PRINT white to yellow, yellow-orange, or ochraceous.
Spores with amyloid warts and/ or ridges. Cap tissue (and usually other tissue) containing nests or
rosettes of sphaerocysts.

RUSSULA is distinct by virtue of its brittle (dry or granular) flesh, rigid fruiting body,
white to yellow-orange spore print, and warty amyloid spores. “Brittle,” of course, is a
subjective term, but the crisper Russula species will snap audibly when broken, and in
both the crisp and fragile types the stalk is fleshy and snaps open cleanly like a piece of
chalk—i.e., there is no fibrous tissue present as in most fleshy-stemmed mushrooms.
Lactarius species also snap open like chalk, but usually exude a latex when broken, while
Leucopaxillus is somewhat brittle but has a tough, fibrous stem that does not break like
chalk and often has white mycelium at the base. In addition to their brittle texture, the
Russulas have a characteristic appearance that, though difficult to describe, makes them
one of the easiest groups to recognize. The cap is plane to depressed at maturity and often
broader than the length of the stem—with exceptions, of course.
Though Russula as such is a very clearcut group, mastering their identification is almost
to be despaired of, for Russulas come in a bewildering panoply of reds, purples, pinks,
yellows, oranges, browns, greens, and whites that is at once their most attractive and
deceptive feature. The color pigments are unusually sensitive to environmental and genetic
caprice, so that no two mushrooms look quite alike. Identification is made even more
difficult by the fact that many species produce only a few fruiting bodies per mycelium—
making it hard to gauge accurately the degree of color variation. The result is that many
species are still poorly known or unclassified, while synonyms and homonyms abound.
.Ktmw/tf-researchers have resorted to examining the spore ornamentation of different
species with an electron microscope in an effort to establish fixed criteria for each species
and dispel some of the confusion.

Russula xerampelina is one of the few Russulas worth gathering for the table. Note the shape, which is
fairly typical of the genus Russula, and note how the stalk fractures like a piece of chalk.
84 RUSSULACEAE

The task of getting to know the Russulas is expedited by assigning them to groups. For
instance, the section Compactae includes several large, hard-stemmed species with
gills that usually alternate long and short (R. albonigra and R. densifolia are good
examples). Another group, centered around R. fragrantissima and R. sororia, has a
yellow-brown to brown, conspicuously striate cap and a strongly fragrant to unpleasant
odor. A third group, mainly northern in distribution, has flesh which turns grayish when
exposed (e.g, R. claroflava). The remaining Russulas can then be divided into four large,
artificial groups based on spore color and taste: to wit, spores white and taste mild (e.g., R.
cyanoxantha), spores white and taste acrid (e.g., R. emetica), spores yellow and taste acrid
(e.g., R. rosacea), and spores yellow and taste mild (e.g., R. xerampelina).
Russulas are mycorrhizal with a broad range of hardwoods and conifers, and are almost
always terrestrial. They grow not only in woods, but at the edges of pastures, in brushy
areas, and on lawns near trees. They are among the most omnipresent mushrooms of our
western coniferous forests, but are often concealed by leaves and needles and evident only
as low mounds (“mushrumps”) in the humus.
In eastern North America the Russulas, like the milk caps (Lactarius), are often
prominent during muggy weather when other mushrooms are relatively scarce, while in
the Pacific Northwest and Rocky Mountains their season more closely coincides with that
of other fungi. In our area Russulas run rampant throughout the mushroom season,
subject, of course, to the whim of the weather. A metagrobolizing melange of species (R.
albonigra, R. xerampelina, R. cyanoxantha, et al) usually erupts after the first fall rains,
followed by another burst (spearheaded by R. emetica) in the winter.
Russulas are among the most maligned of all mushrooms. Even veteran mushroom
hunters treat them mercilessly—throwing them over their shoulder or crushing them
underfoot with disparaging remarks like “Oh, it’s a JAR” (“Just Another Russula”). Their
omnipresence, anonymity, and poor culinary reputation are partly responsible, I am sure.
Also, their brittle flesh is irresistible to those who like to smash things. Furthermore, they
have a habit of forming “mushrumps” in the humus which resemble those made by Boletus
edulis. Frustrated boletivores can be uncompromisingly brutal (see p. 546), and their
habitual hunting grounds are inevitably strewn with the broken bodies of Russulas.
However, mushrooms were not created for the exclusive enjoyment of Homo sapiens, and
it is wrong to judge them accordingly. T ry to resist the sharp temptation to mash, maim, and
mutilate them. Those who follow in your footsteps will appreciate your sensitivity and
self-restraint.
Most of the mild-tasting Russulas are edible, but this should not be taken as a signal to
sample them indiscriminately. Always identify what you intend to eat! Those which red¬
den or blacken and/ or have a peppery (acrid) taste should be avoided—at least some cause
vomiting and diarrhea or worse. Thorough cooking may render them edible, but it hardly
seems worth the effort or risk. Russulas are not widely revered as esculents, and it is
true that their dry, granular flesh does not blend well with some dishes. However, R.
xerampelina and R. cyanoxantha (to name just two of the local species) are marvelous
delicacies in their own right, and R. virescens and R. crust osa of eastern North America are
also excellent.
About 200 species of Russula are reported from North America, but the vast majority of
them are “JAR’s”—i.e., depressingly difficult to demystify. Only some of the more
distinctive types are described here. You will doubtlessly encounter many, many more—
some of them unclassified. If your “JAR” does not key out convincingly, you can send
it to a specialist, or better yet, the nearest compost pile.
RUSSULA 85

Key to Russula
l. Fruiting body medium-sized to very large, hard (especially the stalk); cap white to brown, black,
or grayish (never brightly colored); gills typically adnate to decurrent and usually alternating
long and short; cap cuticle (skin) not peeling easily, margin not striate .2
1. Not with above combination of characteristics . 10
2. Fruiting body basically white (but often with brown to yellowish stains); flesh and stalk surface
not staining when bruised or cut . 3
2. Flesh and stalk surface changing color when bruised (within 10 minutes) and/ or fruiting body
not white .4
3. Cap 5-10 cm broad; taste very acrid; gills and stalk never blue-green .
. R. cascadensis(see R. brevipes, p. 87)
3. Cap 7-20 cm or more broad; taste mild to slightly acrid, or if distinctly acrid then gills and/ or
stalk apex usually tinged blue-green or green.R. brevipes & others, p. 87
4. Fruiting body turning dark gray to black Ji'recr/y whenbruised(within5-10 minutes), withoutan
intermediate reddish phase; fruiting body blackening in age .../?. albonigra & others, p. 89
4. Not as above; if staining black, then with a preliminary reddish phase . 5
5. Flesh and stalk surface distinctly staining reddish to orange when scratched or bruised (within
5 minutes), then eventually darkening to dark gray, dark brownish-gray, or black .6
5. Not as above (but may stain reddish without blackening afterward or may stain smoky-brown
directly) . 8
6. Gills thick and widely spaced .R. nigricans (see R. densifolia, p. 90)
6. Not as above . 7
7. Cap viscid when moist; taste usually acrid .R. densifolia, p. 90
7. Cap not viscid; taste typically mild .R. dissimulans (see R. densifolia, p. 90)
8. Gills usually with sordid reddish stains, often entirely reddish in old age; fruiting body not
blackening; associated with oak in California and southern U.S. R. subnigricansgroup, p. 90
8. Not as above .9
9. Flesh typically staining faintly reddish when cut (within 20 minutes), and then eventually pale
smoky-brown (or at times without reddish phase) R. adusta( see R. subnigricans group, p. 90)
9. Not as above . 10
10. Fruiting body very firm and stalk hard; cap whitish becoming ochre, rusty-brown, or reddish-
brown, margin not striate; all parts bruising or discoloring ochraceous to cinnamon-brown;
odor often unpleasant in age, never sweet; spore print white; common in eastern North
America .R. compacta (see R. subnigricans group, p. 90)
10. Not as above . 11
11. Odor heavy and sweet (like maraschino cherries or benzaldehyde), but in age often becoming
fetid; cap medium-sized to large (5-15 cm broad), yellow-brown to straw-color, dull ochre, or
yellow-orange .R. fragrantissima group, p. 92
11. Not as above . 12
12. Cap 5-12 cm broad, typically dark red with a nearly blackish center; taste usually acrid but some¬
times mild; spore print whitish; common under hardwoods in eastern North America .
. R. atropurpurea (-R. krombholzii)
12. Not as above . 13
13. Flesh and stalk surface staining gray to ashy or black (sometimes slowly and sometimes with
an intermediate reddish phase) . 14
13. Not as above . 17
14. Cap yellow to ochre . 15
14. Cap orange to red, coppery, purple-red, etc. (sometimes mixed with yellow) .. 16
15. Taste mild .R. claroflava, p. 92
15. Taste acrid .R. ochroleuca{see R. claroflava, p. 92)
16. Cap red to orange or coppery-brown; flesh bruising gray directly . R. decolorans group, p. 91
16. Cap purplish, greenish, brownish, etc. (often a mixture of colors); flesh usually turning reddish
before graying . R. occidentalis & others (see R. decolorans group, p. 91)
86 RUSSULACEAE

17. Cap bright yellow to golden-yellow; mature gills yellowish R. lutea (see R. claroflava, p. 92)
17. Not as above . 18
18. Cap white or whitish to yellowish-white or pale yellow . 19
18. Cap some other color (including yellow-brown) when fresh, but may fade to whitish in age 23
19. Cap cuticle (skin) thick and rubbery .R. crassotunicata (see R. cremoricolor, p. 97)
19. Not as above . 20
20. Spore print and gills white or whitish . 21
20. Spore print and mature gills yellowish . 22
21. Cap pale yellow to yellowish-white, viscid when moist, the margin usually striate in age; taste
acrid; usually scattered to gregarious .R. cremoricolor & others, p. 97
21. Cap dull white or tinged buff or pink at center, often dirty, viscid or dry; taste mild or acrid;
usually occurring in small numbers .R. albidula group, p. 96
22. Stalk typically staining yellow and then brown when scratched; odor fishy in age .
.R. sp. (unidentified) (see R. xerampelina, p. 102)
22. Not as above .R. maculata group, p. 100
23. Taste of flesh and/ or gills distinctly acrid (taste them both!) .24
23. Taste mild or nearly so . 32
24. Stalk white (but sometimes with stains or discolorations, especially at base) .25
24. Stalk red or flushed pink to rose .31
25. Cap yellow-brown to dull straw-color, hazel-brown, grayish-brown, or even darker at center;
margin striate and with small bumps, at least in age; odor usually rather unpleasant; very
common .R. sororia group, p. 93
25. Not as above . 26
26. Spore print and gills white or whitish . 27
26. Spore print yellow to ochre; gills yellowish at least in age .
. Various JARs (see R. alutacea group, p. 102 & R. maculata group, p. 100)
27. Stalk and flesh often slowly staining orange or pink when bruised (sometimes with an inter¬
mediate yellowish phase); cap typically multicolored (usually yellowish at center and splotched
with yellow, orange, or red toward margin) .R. bicolor
27. Not as above . 28
28. Cap red suffused with gray or brown; associated with western conifers and especially common
in the Rocky Mountains .R. montana
28. Not as above (but cap may be bright red) . 29
29. Cap bright red when fresh, but often fading to pink, orange, or sometimes even white .
. R. emetica group, p. 97
29. Not as above; cap variously colored but not bright red . 30
30. Many gills forked at quite a distance from the stalk; stalk firm, 1-3 cm thick, cap 5-15 cm broad
.R. variata(see R. cyanoxantha, p. 94)
30. Gills not forked, or forked only near stalk; stalk usually less than 1.5 cm thick, often quite fragile;
cap up to 7 cm broad .R.fragilis & others, p. 98
31. Stalk red to pink; cap dark red to bright red (but often fading in age) .R. rosacea, p. 99
31. Cap pink, olive-gray, etc., or a mixture of these colors; stalk usually tinged rose or grayish-rose
.R. gracilis group & others, p. 99
32. Cap surface dry and soon areolate (cracked into scales or plaques), greenish to blue-gray or with
buff, yellowish, or even brownish tones; common under hardwoods in eastern U.S.33
32. Not as above . 34
33. Cap usually greenish or greenish-gray .R. virescens, p. 95
33. Cap buff to yellowish or brownish, with green tones sometimes present also.
.R. crustosa & others (see R. virescens, p. 95)
34. Cap predominantly greenish or grayish-green (sometimes mixed with other colors); stalk
white or whitish (or sometimes brownish-stained, but never pink); odor not fishy in age; spore
print white to creamy (pale yellow), not ochre or dark yellow . 35
34. Not with above combination of characteristics; cap may have greenish tones, but typically the
greenish tones not predominating . 38
RUSSULA 87

35. Spore print white; fruiting body often robust (stalk 1.5-3 cm thick or more); stalk not staining
salmon in ferrous sulfate . R. cyanoxantha, p. 94
35. Spore print pale cream; stalk typically 0.7-2 (2.5) cm thick, staining salmon in ferrous sulfate 36
36. Cap bluish-green to greenish, with a matt (dull) appearance .
. R. parazurea (see R. aeruginea, p. 95)
36. Cap greenish to grayish, often with yellowish, rusty, or brownish areas, and viscid when
moist . 37
37. Cap usually greenish; found with aspen, birch, oak, and conifers .R. aeruginea, p. 95
37. Cap often with other colors; found with oak and beech . . . R. grisea (see R. aeruginea, p. 95)
38. Cap small to medium-sized (2-7 cm broad); surface dry, reddish to purple, but overlaid with
a fine whitish powder or velvety bloom .R. mariae, p. 96
38. Not as above; cap lacking whitish bloom . 39
39. Spore print and gills white or whitish .40
39. Spore print yellow to ochre; gills yellowish in age .42
40. Cap typically with purplish, pinkish-lilac, bluish, and/or yellow tones; fruiting body some¬
times large; gills often slightly greasy to the touch; stalk not staining salmon in ferrous sulfate
. R. cyanoxantha, p. 94
40. Cap usually browner, grayer, redder, or more flesh-colored than above; fruiting body usually
medium-sized; stalk staining salmon in ferrous sulfate .41
41. Predominant color of cap typically flesh-color or reddish; cuticle (skin) often wrinkled; found
mainly in eastern North America .R. vesca(see R. cyanoxantha, p. 94)
41. Predominant color of cap typically some shade of brown, olive-brown, gray, or gray-brown;
widespread . R. brunneola(see R. cyanoxantha, p. 94)
42. Stalk typically staining yellow when scratched, then slowly becoming brownish; stalk usually
rose-colored or at least with a blush of pink, but white in some forms; fruiting body medium¬
sized to very large; odor shrimpy or fishy in old age .R. xerampelina, p. 102
42. Not as above . 43
43. Cap red to pink, orange, or whitish, or a mixture of these colors; associated with hardwoods,
especially oak; stalk white .R. maculata group, p. 100
43. Not as above .44
44. Cap 2-6 cm broad, markedly fragile, variously colored but usually with at least some purple
. R. placita group & others, p. 100
44. Not as above; larger and/ or not markedly fragile .45
45. Cap large, firm, sometimes massive (up to 35 cm broad!); stalk often tinged pinkish .
.R. olivacea(see R. alutacea group, p. 102)
45. Not as above; medium-sized to fairly large, but not massive; stalk usually white .46
46. Spores and mature gills creamy; found with hardwoods . . R. grisea (see R. aeruginea, p. 95)
46. Spores and mature gills yellow to ochre; found with hardwoods or conifers . 47
47. Cap medium-sized (3-12 cm broad), burgundy to livid red, purple, or rusty-red, or with a
brownish center, or with reddish-buff tones; fruiting body quite firm R. integragroup, p. 101
47. Not as above; medium-sized to fairly large (cap 5-20 cm broad) and often fragile in old age
.R. alutacea group & assorted JARs, p. 102

Russula brevipes (Short-Stemmed Russula)

CAP 7-30 cm broad, broadly convex with a depressed center and inrolled margin, be¬
coming broadly vase-shaped in age; surface dry, unpolished, minutely woolly or felty,
white or whitish but often dirty and/or with yellowish to brown stains and discolorations;
margin not striate. Flesh thick, crisp, brittle, white, not staining; taste mild or slowly acrid.
GILLS thin, close or crowded, adnate to decurrent, usually alternating long and short,
white or creamy (but tinged blue-green in one variety), often brownish-stained in age.
STALK 2-7(10) cm long, 2-5 cm thick, hard and rigid, often quite short and stout, equal or
tapering downward, smooth, dry, dull white or brownish-stained, sometimes with blue-
green tinge at apex. SPORE PRINT white to pale buff; spores 8-11 x 6.5-9 microns,
elliptical, with amyloid warts and ridges.
 ■ ■ .

“Better kicked than picked,” Russula brevipes is one of the most common and mundane of all the
Russulas. Note the centrally depressed or vaselike white cap. The stem can be slightly longer—or
considerably shorter—than shown here and the cap is often incrusted with dirt.

HABITAT: Solitary or more often scattered or in groups or troops on ground in woods

of all kinds; widely distributed and often abundant. It hugs the ground closely and may be
visible only as a low mound or “mushrump” in the humus. In our area the major crop is
typically in November or December, but it can be found most any time.
EDIBILITY: Edible, but better kicked than picked. In my experience it is insipid and
granular, but some people are apparently fond of it. One authority recommends stewing
it slowly in soups; another says to “throw it in with the sauerkraut.” The flavor and texture
are substantially improved when it is parasitized by Hypomyces lactifluorum(see p. 884).
COMMENTS: “So large, so mediocre” and “Better kicked than picked” typify the
comments this tedious, vulgar, mundane mushroom received when overwhelmingly voted
the “Most Boring Mushroom” of the 8th Annual Santa Cruz Fungus Fair. (Second place
went to “the one behind the desk answering questions.”) Actually, it is a rather attractive
mushroom when first encountered, the pristine cleanness (clean pristineness?) of the gills
contrasting prettily with the dirty hunks of humus hoisted aloft by the concave cap. How¬
ever, as the above comments attest, one quickly tires of finding it—not only because of its
ubiquitousness (it is by far the most common of the larger Russulas), but also due to its
annoying habit of forming deceptively promising “mushrumps” in the humus (thwarted
boletivores are not known for their ability to perceive beauty).
R. brevipes is easily recognized by its dull white color, centrally depressed cap with an
inrolled margin when young, hard stem, and crisp flesh that does not blacken or redden
when bruised. It has the general aspect of a large Lactarius (e.g., L. piperatus), but lacks a
latex, though the gills when young may be beaded with water droplets. Var. acrior is the
form with an acrid taste and blue-green tints in the gills and stalk apex. Other species: R.
romagnesiana is a rare but very similar eastern species with smaller spores;i?. cascadensis
also has smaller spores, but has an intensely acrid taste and smaller cap (4-12 cm broad)
and occurs under conifers in the Pacific Northwest. R. delica is also close to R. brevipes,
but is said to have thick, well-spaced gills. It was originally described from Europe and is
apparently rare in North America.

88
Russula albonigra is a hard, robust species that is decidedly schizophrenic: white in youth, black in age
(note the young whitish cap at the bottom). It also blackens when bruised.

Russula albonigra (Blackening Russula; Integrated Russula)

CAP 7-20 (25) cm broad, broadly convex or centrally depressed becoming broadly
depressed in age; surface dry to slightly viscid, sometimes polished; smooth, at first white,
soon becoming grayish or blackish-brown and finally entirely black; margin not striate.
Flesh thick, crisp, brittle, white, bruising gray and then black; taste mild or slightly acrid.
GILLS adnate to slightly decurrent, close to rather well-spaced, thick, brittle, usually
alternating long and short; creamy-white staining gray or black, often entirely black in old
age. STALK 3-13 cm long, 2-5 cm thick, very hard, stout, solid, rigid, smooth, equal or
tapered downward, white becoming grayish or brownish-gray in age or where wounded,
then black. SPORE PRINT white; spores 7-10 x 5.5-7.5 microns, broadly elliptical to
nearly round, with amyloid warts and ridges.
HABITAT: Scattered or in groups or troops under both hardwoods and conifers; wide¬
ly distributed. It is sporadically common in our oak-madrone woodlands but appears to
be more abundant than it actually is because the fruiting bodies do not decay readily.
In the Pacific Northwest it favors conifers.
EDIBILITY: Better punted than hunted. Although said to be edible if thoroughly cooked,
it is hardly tempting and a closely related Oriental species is poisonous.
COMMENTS: This sometimes massive Russula is easily identified by its hard cap and
stem and the blackening of all parts in age or when handled. It is decidedly schizophrenic
in appearance—young specimens are reminiscent of R. brevipes and Hygrophorus
sordidus in their whiteness, while elderly individuals are completely jet-black (hence the
epithet albonigra, meaning “white-black”). Most dramatic, however, are those specimens
in transition, i.e., with a black cap, black stem, and white gills (the gills generally take longer
to blacken than the cap and stem). R. sordida is a passe synonym. R. atrata is a very similar
west coast species that also blackens, but has a much thicker cap cuticle. Several allied
species stain reddish before turning black (see R. densifolia), and R. adusta (see comments
under the R. subnigricans group) darkens less dramatically.

89
90 RUSSULACEAE

Russula densifolia (Reddening Russula)

CAP 5-15 cm broad, broadly convex to plane or depressed; surface viscid when moist, soon
dry and often polished; smooth, whitish to pale buff becoming grayish or brownish or
eventually blackish in old age; margin not striate. Flesh thick, crisp, white, slowly bruising
reddish or orange-red, then eventually grayish-brown to black; taste usually acrid. GILLS
adnate to slightly decurrent, close, brittle, whitish, developing sordid reddish to smoky
stains in age, usually alternating long and short. STALK 4-10 cm long, 1-3 cm thick, hard,
rigid, smooth, equal; whitish when fresh but staining like the flesh (reddish or orangish,
then smoky-brown to black). SPORE PRINT white; spores 7-10 x 6-8 microns, broadly
elliptical with amyloid warts and ridges.
HABITAT: Solitary, scattered, or in groups in woods; widely distributed. It is fairly
common locally, along with R. dissimulans (see comments), but is not nearly as numerous
as R. albonigra, R. brevipes, or R. subnigricans. The major crop is in the fall or early winter.
EDIBILITY: Better dribbled than nibbled. The sharp taste is said to disappear in cooking,
but the end product is insipid at best and indigestible or even poisonous at worst.
COMMENTS: The slow staining of wounded tissue to reddish and then grayish-brown,
sooty-gray, or black is the calling card of this coarse Russula and its close relatives (see
below). The staining is a tactful compromise between R. albonigra, which blackens
directly, and the R. subnigricans group, which stains reddish but not black. The full
staining sequence may take half an hour and is best seen by scratching the stalk. Other
coarse, closely related Russulas with the same staining reactions include: R. dissimulans,
common and widely distributed, with a dry (not viscid) cap, mild taste, and slightly larger
spores; and R. nigricans, fairly common (at least on the west coast), with very thick, well¬
spaced gills and smaller spores. Both of these species are “better beaten than eaten.”

Russula subnigricans group (Rank Russula)

CAP (5) 9-20 (30) cm broad, broadly convex to plane or depressed; surface slightly viscid
when moist, smooth, whitish when young, soon becoming pale smoky-brown or dingy
reddish-brown or dull ochraceous-stained, or often a sordid mixture of these shades;
margin at first inrolled, not striate. Flesh thick, crisp, white, typically bruising slowly
reddish (in 5-20 minutes), then usually browner or grayer within an hour, but sometimes
staining grayish-brown directly (especially in stalk), or not at all; odor strong and
unpleasant, at least in age or upon drying; taste mild to slightly bitter. GILLS thick, brittle,
adnate to slightly decurrent, usually alternating long and short; pallid, soon stained dark or
sordid reddish; close to fairly well-spaced. STALK (3) 7-13 cm long, (1.5) 3-7 cm thick, very
hard, rigid, solid; equal or with tapered base, smooth, white, staining sordid reddish and / or
smoky-brown in age or when handled. SPORE PRINT white; spores 6-10 microns, nearly
round with low amyloid warts and ridges.
HABITAT: Solitary, scattered, or in groups under live oak; common in our area fromlate
summer through early winter and also reported from Mississippi. In our area it is a good
chanterelle-indicator (see comments).
EDIBILITY: Better trampled than sampled. The rank odor is hardly enticing and the
Japanese version is extremely poisonous according to mycologist Tsuguo Hongo.*
COMMENTS: This vulgar giant of the genus is very close both macro- and micro¬
scopically to R. subnigricans, a species originally described from Japan. (Our version,
upon which the above description is based, seems to differ only in its slightly larger size,
closer gills, and rank odor at maturity.) The principal fieldmarks are its hard stem, brittle

* Hongo, coincidentally, is the Spanish word for “mushroom.”

Russula subnigricans group. This large rank Russula develops dingy reddish stains on the gills,
but never blackens. It seems to grow only with oak—usually in the company of chanterelles.

texture, large size, coarse appearance, and reddening of the gills in age or where wounded.
The staining reactions elsewhere on the fruiting body are rather variable, but at no point
does it blacken as in R. albonigra, R. densifolia, R. dissimulans, or R. nigricans. The cap
color is rather amorphous—generally an unbecoming blend of murky gray, smoky-brown,
and sordid reddish or ochraceous—but in one collection I made the fruiting body had
bluish-green tinges as in R. brevipes var. acrior. Another interesting feature is its affinity for
chanterelles, one or more of which will often grow right out of the base of its stem—or if not,
then in the immediate vicinty! Other species: R. adusta is a somewhat similar, widespread
species which has rather erratic staining reactions (the flesh normally bruises smoky-
brown or grayish-black, but may show a slight reddish intermediate phase); its gills,
however, do not redden in age. R. compacta of northern and eastern North America is
also somewhat similar, but stains ochre or brown and has a pallid to rusty- or reddish-
brown to ochre-brown cap. Both of these are “better stomped than chomped.”

Russula decolorans group (Graying Russula)

CAP 5-15 cm broad, nearly round becoming convex to plane or slightly depressed; sur¬
face smooth, viscid when moist, color variable: dull red to orange, red mixed with yellow,
coppery-brown, or at times with cinnamon or tawny tones; margin striate in old age. Flesh
thick, firm, white, becoming gray, ashy, or black when bruised or exposed (sometimes
slowly); odor and taste mild. GILLS typically adnate to adnexed, fairly close, creamy
becoming pale ochre or yellowish, sometimes grayish-stained in old age. STALK 4-12 cm
long, 1 -3 cm thick, equal or often thicker below, very firm when young (but often softer in
age), white, staining and aging grayish. SPORE PRINT yellowish to pale ochre; spores
9-14 x 7-10 (12) microns, elliptical, with isolated amyloid warts.
HABITAT: Solitary to widely scattered or gregarious on ground in mixed woods and
under conifers, associated principally with pine; widely distributed.
EDIBILITY: Better smashed than stashed—it is said to be edible, but there are several
look-alikes of unknown edibility.

91
92 RUSSULACEAE

COMMENTS: The graying flesh, reddish to orange cap, and mild taste are the fallible
fieldmarks of this firm, rather robust Russula, which is actually a close-knit “complex” of
species. One striking form found in our coastal pine forests is very firm and has a somewhat
shiny cap in dry weather. Other species: R. paludosa has a reddish cap and flesh that grays
somewhat, but the taste is usually slightly bitter or acrid. R. obscura has a darker (purple-
red to brownish) cap, mild taste, and flesh that stains pinkish or reddish before blackening.
Both of the above species are quite firm and robust, have ochre spores, and grow with
conifers. Another distinctive species,/?, occidentals, has a grayish to olive-brown to buff,
brownish, or vinaceous cap (usually a mixture of these colors) and mild taste. Its stalk
and flesh are white but usually stain reddish and then gray (or gray directly) when bruised.
It is fairly common under mountain conifers in the Pacific Northwest.

Russula claroflava (Yellow Russula)

CAP (3) 4-12 cm broad, nearly round becoming convex to plane or slightly depressed;
surface slightly viscid, soon dry, yellow to golden-yellow, smooth; margin slightly striate in
age. Flesh white, slowly staining grayish when rubbed; odor and taste mild. GILLS adnate
to adnexed or free, close, creamy becoming pale ochre; sometimes grayish-stained in age.
STALK 3 -8 cm long, 1-2 cm thick, more or less equal, white to pale yellow, dry, smooth,
aging or slowly bruising grayish. SPORE PRINT yellow-ochre; spores 8.5-10 x 7.5-8
microns, broadly elliptical with amyloid warts and ridges.
HABITAT: Scattered or in small groups in woods; mainly northern or montane in distri¬
bution and partial to birch, aspen, and various conifers. I have found it in New Mexico
under aspen, but not in California. It fruits mainly in the summer and fall.
EDIBILITY: Edible.
COMMENTS: The pretty yellow cap and tendency to stain or age ashy-gray rescue this
species from “JAR” status. R.flava is a synonym. R. ochroleuca is a similar species with
a yellow-ochre cap, acrid taste, and flesh that grays quickly. Another yellow-capped spe¬
cies, R. lutea, is rather fragile, has a mild taste, ochre gills, and does not stain gray. All of
these are fairly common in northern North America under conifers, birch, and aspen.

Russula fragrantissima group Color Plate 11

(Fragrant Russula; Fetid Russula)
CAP 5-15 (20) cm broad, nearly round becoming convex, then plane or somewhat
depressed; surface viscid or slimy when moist, smooth, yellowish to yellow-brown to bright
ochre, straw-colored, orange-brown, or brown, often darker (or reddish-spotted) at the
center; margin usually radially furrowed with small bumps (tuberculate-striate), at least in
old age. Flesh thick, white or tinted cap color; odor heavy and penetrating—at first sweet
(like maraschino cherries, almond extract, or benzaldehyde), but in age with a nauseating
(fetid) component; taste nauseating; taste of gills acrid. GILLS creamy-white becoming
yellowish or pale ochre, brownish-spotted in age, often beaded with water droplets when
fresh; adnate to adnexed or free, fairly close. STALK 3-20 cm long, 1 -4 cm thick, equal or
somewhat tapered below, whitish to buff becoming yellowish or brownish-stained in age,
especially near base; smooth or longitudinally lined, dry. SPORE PRINT pale orange-
yellow; spores 6-9 x 6-8 microns, broadly elliptical to nearly round with amyloid warts and
at least partially reticulate.
HABITAT: Solitary or scattered to gregarious on ground under both hardwoods and
conifers, widely distributed. In our area it fruits mainly in the fall under oak, tanoak, and
madrone; it can be found every year but is not nearly as numerous as the R. sororia group.
RUSSULA 93

EDIBILITY: U nequivocally inedible and possibly poisonous—in other words, it is better

punched than munched. The disgusting taste and strong odor should discourage most
people from sampling it.
COMMENTS: Anyone under the mistaken impression that all mushrooms smell alike
should get a whiff of this robust, odoriferous Russula. The penetrating odor is reminiscent
of maraschino cherries, but with a fetid component that becomes more and more pro¬
nounced as the mushroom matures. The cap is often merely flat at maturity, rather than
depressed as in most of the larger Russulas, and the stalk isoftenlonginrelationtothecap.
The yellow-brown to ochre cap, fairly large size, and water droplets on the gills (when
moist) are also distinctive. Other species in the “complex” include:: R. laurocerasi
(possibly the one in the color plate), a slightly yellower, cleaner, slimmer, larger-spored
version of R.fragrantissima whose odor is strongly fragrant through maturity, albeit with a
subtle fetid component; it is also widely distributed and the two appear to intergrade. Both
have passed under the namzR.foetens, a European species with a nauseating odor even in
youth and non-reticulate spores. Its North American counterpart,/?, subfoetens (-R.
foetentula?) is smaller, has a bright rusty-orange to rusty-yellow to brownish cap, and
does not smell as strong. Finally, there is R. ventricosipes, which grows in sand under
northern conifers and has a large, conspicuously swollen (ventricose) stalk.

Russula sororia group (Comb Russula)

CAP 4-12 cm broad, convex becoming plane or centrally depressed; surface viscid when
moist, smooth, dark grayish-brown to hazel-brown when young (rarely whitish), usually
paler (yellow-brown to straw-colored or pale grayish-brown) toward margin or in age;
margin usually furrowed with low bumps (tuberculate-striate) in age. Flesh brittle, firm,
white, not bruising; odor unpleasant (rancid or “spermatic”); taste mild or slightly acrid;
taste of gills slowly but distinctly acrid. GILLS white or creamy, but often brownish- or
rusty-stained in age; close, adnate to adnexed or free. STALK 3-8 cm long, 1-2.5 cm thick,
equal or tapered downward, whitish, but often rusty-stained near base; often with large
cavities (usually 3) within. SPORE PRINT creamy to pale yellow; spores 6-9 * 5-7
microns, broadly elliptical with amyloid warts.
HABITAT: Scattered to densely gregarious in woods or at their edges, on lawns under
trees, etc; common and widely distributed. In our area it is often abundant in the fall and
winter, especially under pine, oak, and Douglas-fir.

Russula sororia is abundant under both hardwoods and conifers. The yellow-brown to grayish-
brown cap has a striate margin in age.
94 RUSSULACEAE

EDIBILITY: Better stomped than chomped—it might be poisonous and doesn’t taste
good anyway.
COMMENTS: The brown to grayish-brown to dingy straw-colored cap, tuberculate-
striate margin in age, peppery-tasting gills, unpleasant odor, and moderate size typify a
group of Russulas that are at times exceedingly abundant. They differ from the R.fragran-
tissima group in their smaller size, grayer color, and different odor. They have passed under
the names R. peciinata, a European species with a yellower cap, and R. pectinatoides,
a North American species that is also yellower than R. sororia, and has a less acrid taste
and darker (yellower) spores. Other species: R. amoenolens is the same as R. sororia
(as defined here); R. cerolens is practically the same but has partially reticulate spores;
R. granulata and R. pulverulenta have scurfy-granulose caps.

Russula cyanoxantha (Variegated Russula) Color Plate 12

CAP 5-18 cm or more broad, convex becoming plane or depressed; surface viscid when
moist but soon dry, smooth, color variable, sometimes entirely but more often a mixture of:
pinkish-lilac, dull purple, green, olive, yellow, blue-green, white, and/ or brown; margin
sometimes obscurely striate. Flesh thick, firm, white, crisp; odor mild or pleasant, taste
mild. GILLS adnexed to adnate or slightly decurrent, many of them forked at least once;
white or with a few brownish stains, slightly greasy to the touch. STALK 5-13 cm long, 1.5-5
cm thick, solid or stuffed, rather hard; more or less equal, smooth, white, but the flesh in
base sometimes grayish. SPORE PRINT white; spores 7-10 * 6-8 microns, broadly
elliptical to nearly round with isolated amyloid warts. Stalk not staining salmon in ferrous
sulfate.
HABITAT: Solitary, scattered, or in groups or troops under hardwoods or sometimes
conifers; widely distributed. It is often abundant shortly after the first fall rains in our
live oak-tanoak-madrone forests, but fruits on into the winter. Like many Russulas, it
is often concealed by fallen leaves.
EDIBILITY: Edible and choice; it is delectably sweet when young and rated highly in
Europe. The hard, crisp buttons are hard to beat in omelets or with scalloped potatoes, but
alas, they are also considered “edible and choice” by maggots!

COMMENTS: This variable species is more difficult to characterize than it is to recognize

(see photo on p. 897). The telltale traits are its fairly large size, white gills and spores, firm
white stalk, mild taste, and variegated cap (typically a mixture of pinkish-lilac, dull purple,
green, yellow or even blue). In our form young specimens—especially those still covered
by leaves—are often uniformly colored a beautiful pinkish-lilac-mauve. When exposed
to light they gradually become spotted with yellow, green, etc., and in old age are often a
washed-out whitish color at the center, with slight greenish or lilac tones showing on the
margin. Another distinctive feature is its soft, somewhat greasy gills: they are not as brittle
as those of most Russulas. R. parazurea (see comments under R. aeruginea) is easily
mistaken harmlessly for greenish forms of R. cyanoxantha, but is usually slightly smaller,
not so variegated, and has reticulate or partially reticulate spores. R. variata (-R. cyano¬
xantha var. variata), common in northern and eastern North America, differs in its more
promiscuously forked gills, often acrid taste, and somewhat different cap color—tending
more toward brownish-olive, greenish, dull purplish, or vinaceous-brown. Two other
species, R. vesca and R. brunneola, also have a mild taste and whitish spores, but their
stalks stain salmon or deep salmon in ferrous sulfate. The cap cuticle in R. vesca tends
to be slightly redder or more flesh-colored (and is often wrinkled and/or recedes from
the margin in age), while that of R. brunneola tends to have a brownish to yellowish-
brown to olive-brown or grayish-olive cast. Both are widely distributed and easily
RUSSULA 95

confused with each other (as well as with R. cyanoxantha) when not “typically” colored.
Both are edible, however, so it hardly matters. For similar species with creamy or yellow-
tinged spores, see R. parazurea and R. grisea (under R. aeruginea).

Russula aeruginea (Green Russula)

CAP 3 -9 cm broad, convex becoming plane or slightly depressed; surface viscid when
moist, dull greeen to dark green, sometimes with brown, gray, or yellowish tints or blotches;
margin often striate. Flesh white, brittle; taste mild. GILLS adnate to adnexed or free,
close, brittle, white becoming pale yellowish, often with brownish stains. STALK 4-8 cm
long, 1 -2 cm thick, equal or with tapered base, white or faintly yellow, base often with pale
brown stains. SPORE PRINT creamy to pale yellow or pale orange-yellow; spores 6-9 *
5-7 microns, nearly round with amyloid warts and ridges.
HABITAT: Solitary, scattered, or in groups in woods; widely distributed. In California
I have found it only a few times, but it is sometimes abundant under aspen in the Rocky
Mountains and Southwest, and under oak in the South.
EDIBILITY: Edible. I have fried it.
COMMENTS: This moderate-sized Russula can be told by its smooth green cap, pallid
or brownish-spotted gills, and pale yellow spore print. There is no red or purple in the cap
as in some of the larger, yellow-spored Russulas (e.g., R. olivacea and R. xerampelina).
There are several similar species, including: R.parazurea, a beautiful green-capped species
with a matt appearance, firm white stem, and white spores, found under oak and tanoak
in our area; and R. grisea, fairly common in our area under oak, with a slightly duller
or grayer cap (green mixed with brown, gray, etc., or sometimes purplish or lilac) and
yellowish spores. Also see R. cyanoxantha and the species listed under it.

Russula virescens (Quilted Green Russula)

CAP 5-15 cm broad, nearly round at first becoming convex to plane or slightly depressed;
surface dry and sometimes slightly velvety, markedly areolate at maturity (i.e., cracked
into small flattened scales or patches); dull green to greenish-gray or sometimes dull
bluish-green, at times with ochre, buff, or creamy discolorations; margin not striate or
only very slightly so, often lobed. Flesh thick, white, brittle; odor and taste mild. GILLS
white or creamy, fairly close, adnate to adnexed or even free. STALK 3-9 cm long, 1-3 (4)
cm thick, more or less equal, dry, firm, brittle, white, more or less smooth. SPORE
PRINT white or with a faint yellow tinge; spores 6-9 (10) * 5-7 microns, elliptical to nearly
round, with amyloid warts and/ or ridges.

HABITAT: Solitary to widely scattered or in groups under hardwoods, especially oak

and beech; common (along with R. crustosa—see comments) during the summer months
in eastern North America. It has also been recorded from Montana under birch and is to
be expected in southern Arizona and New Mexico, where several “eastern” species occur.
EDIBILITY: Edible and choice—along with R. cyanoxantha and R. xerampelina, among
the best of the Russulas for the table.
COMMENTS: One of the most distinctive of all the Russulas, this species is easily told by
its greenish to greenish-gray, areolate (cracked or quilted) cap, white spores, and asso¬
ciation with hardwoods. It is one of the characteristic summertime mushrooms of eastern
hardwood forests, where it sometimes fruits with the equally edible Lactarius volemus
and/ or L. corrugis. Other species: R. crustosa is a closely related edible species
with an areolate cap that is slightly viscid when wet and more variable in color (usually
browner or more ochre-buff, but sometimes with a greenish tinge); R. maculosa of the
Southeast resembles R. crustosa, but has a paler cap and disagreeable odor in age.
96 RUSSULACEAE

Russula mariae (Powdered Russula)

CAP 2-7 cm broad, convex becoming plane or slightly depressed; surface dry, purple to
reddish-purple, amethyst, dark crimson, or maroon (sometimes toned with gray or olive),
finely powdered or velvety from a whitish bloom; margin sometimes striate when old.
Flesh thin, brittle, white; taste mild or slightly acrid. GILLS close, white becoming creamy
or pale yellowish, typically adnate to adnexed. STALK 2.5-7.5 cm long, 1-2.5 cm thick,
equal or tapering downward, firm, rigid, white or tinged reddish to purple. SPORE PRINT
creamy-yellow; spores 7-10 * 6-8 microns, broadly elliptical to round, with amyloid warts
and ridges.
HABITAT: Solitary or in groups on ground in woods; rather rare in California, where I
have found it in the fall under oak and Douglas-fir, but a common mid-summer mushroom
in the hardwood forests of eastern North America.
EDIBILITY: Edible, but hardly incredible.
COMMENTS: This exquisite, rather dainty Russula is one of the more easily recognized
species in the genus. The dry, velvety or finely powdered reddish to purple cap, creamy
spore print, and mild taste distinguish it. The “splitters” recognize two species, the purple-
capped forms being called R. alachuana. The name R. mariae should not be confused
with R. mairei, a member of the R. emetica group.

Russula albidula group (Boring White Russula)

CAP 3-8 cm broad, convex becoming plane or shallowly depressed; surface smooth,
slightly viscid when moist, then dry; white or tinged buff at center; margin not striate or
very obscurely so. Flesh fragile, white; taste acrid. GILLS adnate to adnexed or free, brit¬
tle, close, white or creamy. STALK 2.5-7 cm long, 1-2.5 cm thick, equal or thicker below,
dry, smooth, rigid, white; often spongy or hollow in age. SPOREPRINT white or creamy-
white; spores 7-10 microns, nearly round with amyloid ridges.
HABITAT: Solitary or in small groups in woods or at their edges, widely distributed. It is
fairly common in our area from late summer through the spring, usually with oak butnever
in large numbers (like lips and scissors, it tends to occur in pairs). It is partial to poor
soil along trails or roadcuts.
EDIBILITY: Better whacked than sacked—the acrid taste is a deterrent.
COMMENTS: This plain, unprepossessing, profoundly forgettable Russula can be
recognized by its white color, modest size, and acrid taste. It is smaller and more fragile
than R. brevipes and does not have a strongly inrolled cap margin when young. There are
several very similar, confusing, widespread, wishy-washy Russulas. They include R. al-
bella and R. albida, with a mild to slightly bitter taste, and R. anomaAland/?, subalbidula,
with an acrid taste, but distinguishing between them is a matter for the specialist. For a
comparison of the R. albidula group to R. cremoricolor, see comments under the latter.

Russula albidula group. A dingy and often dirty whitish “JAR” of moderate size. R. cremoricolor
(illustrated on next page) is usually much cleaner.
Russula cremoricolor resembles R. emetica except for its cap color, which is yellowish-white.

Russula cremoricolor (Creamy Russula)

CAP 3-10 cm broad, convex to plane to centrally or broadly depressed; surface viscid
when moist, smooth, white to pallid to very pale yellowish-white, the center often slightly
darker (yellowish or buff); margin usually striate in age (but sometimes obscurely). Flesh
brittle, white; odor mild or slightly fragrant; taste distinctly acrid. GILLS white, adnate
to adnexed or ocasionally free, fairly close. STALK 3-10 cm long, 0.7-2.5 (3) cm thick,
equal or slightly thicker below, dry, white or whitish, usually longitudinally lined or striate.
SPORE PRINT white; spores 7.5-10 * 7.5-8 microns, round to broadly elliptical, with
amyloid warts and ridges.
HABITAT: Solitary, widely scattered, or ingroupsortroopsin woods; widelydistributed.
In our area it is often abundant in the late fall and winter in the same habitats as R. emetica.
In the Pacific Northwest it favors conifers.
EDIBILITY: Better obliterated than refrigerated—the acrid taste is a deterrent.

COMMENTS: This species looks like an albino R. emetica, and save for the whitish cap,
there is little difference. The cap, although whitish, is slightly yellower than that of R.
albidula, and is usually cleaner and more markedly striate. Also, R. cremoricolor tends to
fruit in larger numbers. Other species: R. crenulata and R. raoultii are two very similar
species that are yellower (cap more or less pale yellow); the latter also has smaller spores.
Another pale capped, bitter- or acrid-tasting species, R. crassotunicata, can be distin¬
guished by its thick, rubbery or leathery cap cuticle that is often slightly scurfy and its
white stalk that frequently develops brownish spots in age. It occurs under northern
conifers. “Eating raoultii” (or any of the other species discussed here) is not recommended.

Russula emetica group (Emetic Russula; The Sickener)

CAP 3-10 cm broad, rounded-convex becoming plane or broadly depressed; surface viscid
when moist, smooth, bright red to scarlet, the center often darker; fading in age or wet
weather to pink, orange, or blotched with white; margin eventually striate (but some¬
times obscurely). Flesh white (but pink under the cuticle), brittle, odor mild, taste very
acrid. GILLS white or creamy-white, brittle, close, adnate toadnexed or free. STALK 3-10
cm long, 0.7-2.5 cm thick, equal or thicker below, dry, white or whitish, usually with
longitudinal lines. SPORE PRINT white; spores 7-11 * 6.5-9 microns, elliptical with
amyloid warts and ridges.

97
Russula emetica group. The bright red cap (that often fades), white gills, white stem, and peppery
taste make this an easy mushroom to identify.

HABITAT: Solitary, scattered, or in groups or troops in woods, widely distributed and

common. Our member of the R. emetica group fruits late in the season (winter) in chaparral
and mixed woods (oak, madrone, manzanita, and pine), often intermingled with R.
cremoricolor and Lactarius vinaceorufescens. It also grows on very rotten wood—
unorthodox behavior for a Russula.
EDIBILITY: Pleasing to the eyes of all, to the tongues of some, but to the stomachs of none.
As its name implies, it is poisonous, at least raw, and can be used to induce vomiting.
Parboiling may destroy the toxins, but it hardly seems worth the trouble or risk. It is
collected by Himalayan villagers, presumably for use in curries (consult your local travel
agent for details).
COMMENTS: This attractive mushroom is easily recognized by its red cap, white stem
and gills, and red-hot taste. As in many Russulas, the skin peels rather easily from the cap.
For years the name R. emetica has been applied to a confusing group of Russulas with the
above features. I see no compelling reason to discontinue this trend, though the present
consensus seems to be that the “real” R. emetica occurs principally with conifers and favors
sphagnum bogs. Other species in the group include: R. silvicola, an eastern species with a
red to pinkish-yellow cap and slightly smaller spores, common with both hardwoods and
conifers; and R. mairei, colored like R. emetica but associated with hardwoods and with a
firmer texture and smaller spores, 6-8.5 x 5-7 microns (it may well be our local species).

Russula fragilis (Fragile Russula)

CAP 2-5 cm broad, convex becoming plane or depressed; surface smooth, viscid when
moist, color variable: purplish to pinkish, olive-brown, greenish, or even yellow, or a
mixture of these colors, but most often blackish at the center and pinkish or pinkish-yellow
at the margin, with a grayish-olive zone in between; margin striate in age. Flesh thin, very
fragile, white; odor variable; taste acrid. GILLS adnate to adnexed or even free, fairly close,
white or creamy-white. STALK 2.5-7 cm long, 0.5-1.5 cm thick, equal or slightly thicker at
either end, dry, whitish; fragile. SPORE PRINT whitish; spores 6-9 x 5-8 microns, broadly
elliptical to nearly round, with amyloid warts and ridges.
HABITAT: Solitary, scattered, or in groups in woods of all kinds, but especially under
conifers; widespread. I have not seen it in our area, but several similar species occur.

98
RUSSULA 99

EDIBILITY: Not recommended—the acrid taste is a deterrent.

COMMENTS: This species is apt to befuddle the color-conscious beginner with its
numerous color guises, but is rather distinctive when “typical,” i.e., dark at the center,
pinkish toward the margin and grayish-olive between. The other critical features are the
fragile texture, peppery taste, white gills, and white spores. Other species: R. aquosa is a
closely related but slightly larger species; its cap is viscid with a greasy feel and shiny
appearance when wet, and is usually reddish to orange-red or pink with grayish, brownish,
or yellow tints.

Russula gracilis group

CAP 2 -8 cm broad, convex becoming plane or slightly depressed, sometimes retaining a
slight umbo; surface smooth, viscid when moist, color variable: usually some shade of
pink, but often with a dull greenish or grayish-olive to brownish-black center and pale
pinkish margin, and sometimes a rather deep pink or violet-tinged; margin usually striate
in old age. Flesh fragile, white; taste acrid. GILLS typically adnate to slightly decurrent,
whitish becoming creamy and finally tinged yellow, close. STALK 3.5-8 cm long,0.5-1.5
cm thick, equal or slightly thicker below, usually quite soft and fragile, whitish, but usually
tinged or flushed pink to grayish-rose. SPORE PRINT creamy to pale ochre; spores 7-9 *
5-7 microns, elliptical, with isolated amyloid warts.
HABITAT: Widely scattered to gregarious on ground in woods; it is said to grow only
with birch, but a very similar if not identical species is abundant in our area with Douglas-
fir in the fall and winter.
EDIBILITY: Not edible due to the acrid taste.
COMMENTS: Also known asR. gracillima, this species can be told by its fragile pinkish-
tinged stalk, yellowish spores, acrid (peppery) taste, and pinkish or pink-and-greenish cap.
R.fragilis is quite similar in cap color but has whitish spores and a white stalk, while the
R. placita group, which also occurs commonly with Douglas-fir in our area, has a mild
taste, white stem, and darker cap. Other species: R. pelargonia also has a pinkish-tinged
stalk and acrid taste and grows with conifers (especially Douglas-fir), but it has creamy
spores, a more erratically colored cap (grayish, pinkish, brownish, reddish, etc.), and a
distinct geranium-like odor (at least when the flesh is crushed).

Russula rosacea (Rosy Russula) Color Plate 13

CAP 3-12 cm broad, convex to plane or somewhat depressed; surface viscid when moist,
smooth, dark red to bright red, fading in old age to pink or pink blotched with white. Flesh
white, firm, brittle, odor mild, taste very acrid. GILLS creamy-white to pale yellow, adnate

Russula gracilis group. Our member of this species “complex” (shown here) is abundant under
Douglas-fir. Note the slightly colored (pinkish-tinged) stalk.
100 RUSSULACEAE

to decurrent, close, brittle. STALK 4-10 cm long, 1 -2.5 cm thick, equal or with a narrowed
base, smooth, dry, pink or red, or at times only with a blush of red; hollow in age. SPORE
PRINT pale yellow; spores 7-9 * 6-8 microns, nearly round with amyloid warts.
HABITAT: Scattered or in large troops under conifers, especially pine. Abundant on the
Pacific Coast from fall through early spring, fruiting in our area mainly in coastal stands
of Monterey pine, often in the company of Lactarius deliciosus.
EDIBILITY: To be avoided because of the acrid taste.
COMMENTS: This common, conspicuous, colorful inhabitant of our coastal forests is
readily recognized by its red cap, red to rosy stem, creamy gills, and acrid (peppery) taste.
It is one of our prettiest mushrooms—the clean, pale gills contrasting beautifully with the
red cap and stem. At a distance it is sometimes mistaken for Amanita muscaria, which also
grows commonly with pines but has a white stem with a ring (annulus) and volva. The pale
yellow spores and red or pink stem distinguish R. rosacea from R. emetica, while the acrid
taste separates it from R. xerampelina and the R. alutacea group. Other species: R. san¬
guined is probably a synonym. A form with reddish-purple cap and stem—possibly
distinct—is sometimes found in our area growing with R. rosacea.

Russula maculata group

CAP 4-10 (12) cm broad, broadly convex becoming plane or centrally depressed; surface
viscid when moist, smooth, color variable: red to pink, reddish-orange, peachy-pink,
yellowish-buff, or whitish, or often a mixture of these colors, or buffy-white with pinkish
tints on the margin; margin obscurely striate in age. Flesh white, firm becoming fragile in
age; taste mild or slightly peppery. GILLS creamy becoming pale ochre, close, adnexed,
or free. STALK 4-10 (13) cm long, 1-3 cm thick, equal or thicker below, white, the base
sometimes brownish-stained. SPORE PRINT yellow-ochre; spores 7-10 x 6-9 microns,
nearly round, with amyloid warts and ridges.
HABIT AT: Solitary, scattered, or in groups under hardwoods; common in our area in the
fall and winter (sometimes spring) with live oak, but more widely distributed.
EDIBILITY: Unknown.
COMMENTS: The cap color in this species “complex” is quite variable, but within a well-
defined range: red, pink, orange, yellowish-buff, and whitish. The association with oak,
yellow spores, and rather fragile texture in age are also distinctive. The “true” R. maculata
of Europe is said to have a mild to peppery taste and slightly fragrant odor. Our version has
a consistently mild taste and little or no odor. Whether it is the “true” R. maculata or a
closely related species is for the Rwssi//a-researchers to decide. There are a number of
similar reddish-capped, mild-tasting species, such as/?, velenovskyi, which do not fade
as drastically and/ or differ microscopically.

Russulaplacita group (Pleasing Russula)

CAP 2-7 (10) cm broad, convex to plane or somewhat depressed; surface viscid when
moist, smooth, dark purple to wine-colored, reddish-violet, or brownish-purple, often
darker at the center and paler toward margin, but sometimes brown to yellowish at center,
and sometimes washed out in age; margin usually striate in age, at least obscurely. Flesh
rather thin, fragile, white, odor and taste mild. GILLS close, white or creamy soon
becoming yellow and finally dull ochre-yellow; adnate to adnexed or free. STALK 2-8 cm
long, 0.5-2 cm thick, equal or slightly thicker at either end, white, dry, soft and spongy
inside; fragile in age. SPORE PRINT yellow-ochre; spores 7-10 x 6-9 microns, broadly
elliptical to nearly round, with amyloid warts and sometimes ridges.
Russula placita group. As defined here, this group includes many small, fragile Russulas with a
mild taste, purplish cap, and yellow gills. They usually grow under conifers such as Douglas-fir.

HABITAT: Solitary to scattered or gregarious in woods, associated mainly with conifers;

widely distributed. In our area this group is quite common in the fall and winter, especially
with Douglas-fir (often accompanied by R. xerampelina, R. Integra, and Lactarius
rubrilacteus)\ elsewhere it often grows with spruce, fir, hemlock, and pine.

EDIBILITY: Edible but forgettable—thin-fleshed and virtually tasteless. Similar species

are probably—but not definitely—harmless.
COMMENTS: A large number of small, fragile, mild-tasting, yellow-spored Russulas will
fit the above description. Their caps vary considerably in color but usually have some
purple or reddish-purple in them. Their fragility is such that it’s difficult to transport them
home without crushing them (the stalk, though fragile, breaks cleanly like chalk). They lack
the shrimpy odor of R. xerampelina and do not stain brown, and are not as firm as R.
Integra. Their identification is best left to the specialist armed with monographs and a
microscope. Some related species that bear mentioning are: R. lilacea, whose spores
have isolated warts; R. abietina, cap often greenish or brownish; R. puellaris, with pale
creamy-yellow spores;/?, caerulea, with frequently umbonate cap; and/?, chamaeleontina,
with a small, thin, fragile, purplish to lilac to reddish to orange or yellow cap. The latter has
proved invaluable to compulsive categorizers exasperated by the endless nuances of color
in these Russulas. Aberrant forms with yellow spores, fragile texture, and a mild taste can
simply be referred to R. chamaeleontina and conveniently forgotten about. R. fragilis
has served the same purpose in the fragile, white-spored, acrid-tasting group.

Russula integra group

CAP 3-12 cm broad, convex becoming plane or slightly depressed; surface smooth, viscid
when moist, color variable: usually burgundy to vinaceous-brown to livid red or rusty-
red with the margin paler (reddish-buff), but in older specimens often buff or brownish in
the central area and sometimes entirely reddish-buff; margin obscurely striate in old age.
Flesh crisp, firm, white, not staining, odor and taste mild. GILLS adnate to adnexed or
free, white becoming yellowish or pale ochre; close. STALK 3-9 cm long, 1-3 cm thick;
usually equal or tapering upward, dry, white, sometimes with brownish or yellow-brown
spots and stains. SPORE PRINT yellow-ochre; spores 8-11 * 6-9 microns, broadly
elliptical with amyloid warts.
HABITAT: Scattered to gregarious under conifers, widely distributed. Common at times
in our area in the fall and early winter. I have seen large fruitings under Douglas-fir.
EDIBILITY: Edible and good when young and crisp; however, there are many similar
species of unknown edibility, so exercise caution.
COMMENTS: Also known as /?. polychroma, this medium-sized species and its close
relatives can be recognized by their mild odor and taste, yellow spores, white or brownish-
stained stem, and reddish to wine-colored to brownish cap. They are larger than and not
nearly so fragile as the R. placita group, and smaller than R. xerampelina and R. alutacea.
As is commonly the case in Russula, this “complex” is in need of critical study.

101
102 RUSSULACEAE

Russula alutacea group

CAP 5-20 cm broad, convex to plane or somewhat depressed; surface viscid when moist
but soon dry, smooth, color variable: dark red to red, purplish, purple-brown, or purple-
red, or buff to straw-colored at the center or throughout, or with olive shades, or often
a mixture of these colors; margin striate in age. Flesh brittle, white, firm at first, fragile in
age; odor and taste mild. GILLS pale to dull ochre or straw-color; close, adnate to adnexed
or free; brittle. STALK 3-10 cm long, 1.5-4 cm thick, more or less equal, smooth, dry, white
or sometimes tinged pinkish. SPORE PRINT ochre-yellow; spores 8-11 * 6.5-9 microns,
broadly elliptical with amyloid warts. Phenol solution staining stalk and flesh purple-red.
HABITAT: Solitary to scattered or gregarious in mixed woods and under conifers, widely
distributed. In our area this group is especially abundant in old coastal pine forests in
the fall and winter.
EDIBILITY: Not recommended. A large number of species will key out here—and we are
not yet sure whether they’re all edible.
COMMENTS: The above description embraces a number of medium-to-large, mild-
tasting, yellow-spored Russulas. Distinguishing between them is a job for rabid Russula
buffs. The most common variety in our area sometimes grows with R. xerampelina, but is
easily distinguished by its mild odor and marked fragility in age; older caps often disin¬
tegrate in transit unless carefully collected and packed. Another local species is much
firmer, with a reddish, often burnished cap (see comments under R. decolorans). R.
olivacea, which occurs in Europe under hardwoods and in America under conifers, is a
very firm, sometimes massive (cap to 35 cm broad!) species with a more frequently rose-
tinted stem. It is closely related to the true R. alutacea by virtue of its purple-red phenol
reaction. There are also a number of medium-sized to fairly large yellow-spored Russulas
with an acrid taste. These include/?, veternosa and/?, tenuiceps of eastern North America
(and of questionable occurrence in California); and an unidentified western conifer-
lover that has a buffy-brown to pinkish-brown or flesh-colored cap, mild-tasting flesh,
and acrid-tasting gills.

Russula xerampelina (Shrimp Russula) Color Plate 14

CAP 5-30 cm broad, convex becoming plane orcentrally depressed; surface smooth, viscid
when moist, color variable: typically red to dark red, purple, or brownish-olive, but often
laced with(or sometimes entirely) green, brown, yellow-brown, purple-brown, etc.; margin
usually striate in age (at least obscurely). Flesh thick, creamy-white, bruising yellowish and
then brown or discolored brownish in old age, brittle; odor mild becoming shrimpy or
crablike in old age; taste mild. GILLS close, adnate to adnexed, creamy-white becoming
dull yellowish, staining like the flesh, drying brownish or grayish. ST ALK 3-12 cm long, 1 -
4 cm thick, equal or with slightly enlarged base, often longitudinally lined; dry, sometimes
entirely rose-pink, but more often white with just a tinge of pink, or occasionally pure
white; staining yellowish where scratched, then slowly brown; interior spongy and often
discolored in age. SPORE PRINT yellowish; spores 8-11 * 6-8.5 microns, elliptical to
nearly round, with amyloid warts. Flesh and stalk turning deep green in ferrous sulfate.
HABITAT: Solitary to gregarious under conifers; widely distributed and common,
ranging as far north as the Arctic Circle. In our area it is associated almost exclusively with
Douglas-fir and is often abundant in the fall, less so in the winter and early spring.
EDIBILITY: Edible and unforgettable—but one of the least appreciated of our edible
fungi, perhaps due to the mediocrity of its brethren. The young, nutty caps are superb
stuffed with grated cheese, chives, walnuts, and parsley and then broiled. In contrast to
R. cyanoxantha, they are rarely riddled with maggots!
RUSSULA 103

COMMENTS: This beautiful but extremely variable species can be recognized by the
following combination of characteristics: (1) cap viscid when wet, usually with adhering
debris when dry (2) stem usually—but not always—tinted pinkish and always staining
yellowish and finally brown when handled or bruised (3) yellow spore print (4) mild (not
peppery) taste (5) fishy odor at maturity, which is accentuated by cooking or drying
(6) tendency of the gills to age or dry brownish to grayish. In addition, the young buttons
are remarkably rotund and symmetrical—but this character can be appreciated only by
religiously observing a large number of Russulas. Given the extreme variation in color and
size, R. xerampelina may well be a composite species. At least two distinctive—but
intergrading—varieties occur: one with a red to purple cap and lovely rosy stem, the other
with a brownish-olive cap and white or nearly white stem (in addition to the color plate,
there is a black-and-white photo on p. 83). Another distinctive variety occurs in our area
with oak and manzanita; it differs in having a pale greenish-yellow to buff or whitish cap.

HYGROPHORACEAE (Waxy Caps) sp°res

Small to medium-sized, mostly terrestrial mushrooms. CAP typically smooth, often viscid and
brightly colored. GILLS soft, waxy, usually thick and well-spaced, usually attached. ST^LK
central; fleshy or hollow. VEIL absent or evanescent, rarely forming a slight ring (annulus).
VOLVA absent. SPORE PRINT white. Spores round to elliptical, but often shaped like corn
kernels when immature; typically smooth and not amyloid. Basidia long and narrow. Gill tissue
interwoven, parallel, or divergent.

THESE are attractive, often colorful, white-spored mushrooms with soft, clean, waxy-
looking gills. The flesh is not dry and brittle or chalky as in the Russulaceae, and the gills,
though often thick, do not have the blunt edges typical of the chanterelles (Cantharella-
ceae). However, there is little aside from the “waxy” gills to separate them from the
numerous white-spored genera in the Tricholomataceae (particularly Clitocybe, Mycena,
Omphalina, Laccaria, and Marasmius). The “waxiness” is admittedly an ambiguous
character, but with a little experience is readily apparent—the gills are characteristically
soft, clean, fleshy, and . . . they look (and often feel) waxy. (The cap sometimes also has a
waxy appearance, hence the common name “waxy caps.”) Laccaria species have somewhat
waxy gills, but they have tough, fibrous stems and spiny spores. Chroogomphus and
Gomphidius also have waxy gills, but their spores are smoky-black.
The waxy caps have traditionally been grouped together in a single large and diverse
genus, Hygrophorus, but most mycologists now split them into two or more genera based
primarily on the arrangement of the hyphae in the gill tissue (see illustrations, p. 19). As
this character can only be determined with a microscope, the family is treated as one entity
here, but three genera are recognized:
Camarophyllus (hyphae in gill tissue intricately interwoven): Fruiting body small to
medium-sized, usually white or dull colored; cap usually not viscid or only slightly so;
gills typically decurrent, thick, well-spaced, somewhat waxy; stalk dry; veil absent.
Hygrocybe (hyphae in gill tissue more or less parallel): Fruiting body small to medium¬
sized, usually brightly colored and quite waxy; cap dry to moist, viscid, or slimy; gills
free to adnexed, adnate, or sometimes decurrent, obviously waxy; stalk dry or viscid
or slimy, often slender and hollow; veil absent.
Hygrophorus in restricted sense (hyphae in gill tissue divergent from a central strand):
Fruiting body medium-sized to fairly large, usually white or dull colored; cap viscid or
slimy; gills typically adnate to decurrent, well-spaced or close, often only slightly
waxy, usually white or pale colored; stalk slimy, viscid or dry, often fleshy (rarely
hollow), apex often punctate (adorned with pointlike scales or granules); slimy or
fibrillose veil sometimes present.
104 HYGROPHORACEAE

Hygrocybe species are the mushrooms most easily identified as “waxy caps,” while
Hygrophorus and Camarophyllus species are more likely to be confused with other genera,
particularly Clitocybe, because their gills are not so obviously waxy. Some investigators
still recognize only one genus, Hygrophorus, while others recognize only two by throwing
Camarophyllus in with Hygrocybe. Several small, rare “satellite” genera also occur, but are
not treated here.
The long, narrow protruding basidia that give the gills their “waxiness” are reminiscent
of those in Cantharellus, leading some mycologists to infer that Camarophyllus (the most
primitive genus of waxy caps) evolved from Cantharellus-Mkt ancestors. (Camarophyllus
pratensis, in fact, slightly resembles the chanterelle, Cantharellus cibarius.) A relation¬
ship with Omphalina and Clitocybe of the Tricholomataceae has also been suggested.
The waxy caps occur throughout the mushroom season but are partial to cold weather.
In colder climates they may continue to fruit after the first frosts, while in our area they
normally peak in the cold winter months (December-February). Camarophyllus and
Hygrocybe species are mostly saprophytic. Like Leptonia species, they are particularly
abundant in environments where other fleshy fungi are few. In coastal California
this means under redwood and to a lesser extent cypress, but in other regions they are
common in ungrazed fields, heaths, and boggy areas. Hygrophorus species, on the other
hand, are largely mycorrhizal—hence in our area they occur with other mycorrhiza-
formers under oak, madrone, pine, and Douglas-fir, rather than with redwood or cypress.
Hygrophorus and Hygrocybe each contain about 100 species in North America, while
Camarophyllus has about 40. Most waxy caps have wide distributions, and at least half of
the North American species occur in California.
Waxy caps are without a doubt among our most colorful mushrooms, with virtually
every hue in the rainbow represented: blue or green in Hygrocybe psittacina, pink in
Hygrocybe calyptraeformis, bright yellow, orange, or red in a great many species, black in
Hygrocybe conica, white in Hygrophorus eburneus. None are thought to be dangerously
poisonous, and some species are rated highly as edibles. However, I have yet to find one
to my liking. By and large they are too bland or too watery or too bland and too watery
to be worth eating. (See comments on the edibility of Hygrophorus sordidus and H.
russula).) Also, a few species (e.g., Hygrocybe conica and H. punicea) may cause illness.
However, their beauty is reason enough for getting to know them, and they have the
added attraction of being relatively easy to identify in the field! Consequently, a fairly
extensive selection of species is offered here.
Key to the Hygrophoraceae
1. Fruiting body whitish except for yellow to golden-orange powder, flakes, or granules on the cap
and/ or stalk apex (but powder may smear in wet weather) Hygrophorus chrysodon, p. 119
1. Not as above (if fruiting body white, then lacking yellow to golden-orange granules) . 2
2. Fruiting body white to creamy, buff, or pale yellow when fresh . 3
2. Fruiting body at least partially darker or differently colored (including bright lemon-yellow) 17
3. Stalk viscid or slimy when moist, at least over lower portion .4
3. Stalk not viscid . 7
4. Cap tinged buff to yellowish or pale yellow, at least at center; especially common under pine
(but not restricted to it).Hygrophorus gliocyclus & others, p. 120
4. Not as above . 5
5. Stalk usually at least 1.5 cm thick; cap fairly large (5-15 cm broad) .
.Hygrophorusponderatus (see H. sordidus, p. 122)
5. Stalk usually (but not always) less than 1.5 cm thick; cap medium-sized to fairly small .... 6
6. Cap conical when young, usually with a prominent umbo in age; gills not decurrent; found in
eastern North America, rarely in California (mostly under cypress) .Hygrocybepura
6. Not as above; gills adnate to decurrent; widespread .Hygrophorus eburneus, p. 119
HYGROPHORACEAE 105

7. Cap not conical; stalk 1 -7 cm thick, or if thinner then cap viscid when moist and developing dis¬
tinct yellow-brown to reddish-brown, peachy, salmon, or ochre tones in age .8
7. Not as above; stalk typically less than 1 cm thick and/ or cap distinctly conical. 10
8. Stalk often with an annulus (ring) or “volva” that is easily obliterated; typically found under
mountain conifers .Hygrophorus subalpinus, p. 121
8. Not as above; annulus or “volva” absent .9
9. Fruiting body white or discoloring slightly yellowish or buff; associated mainly with oak,
occasionally with conifers .Hygrophorus sordidus & others, p. 122
9. Either fruiting body discoloring more dramatically in age or else associated with conifers and
possessing a densely scurfy stalk apex. 74
10. Growing in grass; stalk tough and pliant; cap often umbonate (seeMarasmius oreades, p. 208)
10. Not as above . 11
11. Odor sharp and cedarlike or fragrant .Camarophyllus russocoriaceus & others, p. 109
11. Odor mild or not distinctive (or merely fungal) . 12
12. Cap conical when young and/ or gills typically notched oradnexed orattimesadnate;gillhyphae
parallel .Hygrocybe subaustraliga& H. albinella (see Camarophyllus borealis, p. 109)
12. Cap not normally conical; gills typically adnate to decurrent; gill hyphae not parallel .... 13
13. Cap viscid when moist . 14
13. Cap not viscid ... 15
14. Cap thin and often translucent-striate when moist, usually less than 3 cm broad; found in
many habitats; gill tissue interwoven .Camarophyllus niveus (see C. borealis, p. 109)
14. Not as above; found mainly under conifers; gill tissue divergent .
.Hygrophoruspiceae (see H. eburneus, p. 119)
15. Growing in groups or troops on pine needles; gills very widely spaced and usually with veins in
between; cap broadly convex soon becoming plane or umbilicate (see photo at top of p. 207)
..(see Marasmius sp. (unidentified), p. 206)
15. Not as above (see top right photo on p. 109) . 16
16. Fruiting body often developing a slightly yellowish tinge in age; cap dry .
.Camarophyllus virgineus(see C. borealis, p. 109)
16. Not as above; cap sometimes slightly lubricous .... Camarophyllus borealis & others, p. 109
17. Cap small (up to 6 cm broad but usually less), dry, honey-colored or yellowish to orange or
scarlet beneath a layer of small, gray to dark brown or blackish fibrillose scales . 18
17. Not as above; either cap viscid or larger or lacking dark scales or differently colored. 19
18. Background of cap scarlet to orange when fresh Hygrocybe turunda (see H. miniata, p. 113)
18. Background of cap buff to dull yellowish or honey-colored; found in eastern North America
.Hygrocybe caespitosa
19. Fruiting body staining gray to black when handled (often slowly); cap conical, at least when
young; common.Hygrocybe conica & others, p. 116
19. Not as above; fruiting body not blackening when handled . 20
20. Cap bright red, yellow, or orange, at least when fresh (but may fade to whitish) . 21
20. Cap some other color (including pink, salmon, vinaceous, purple-red, pale dull orange, etc.) 42
21. Cap and lower portion of stalk viscid to slimy when moist; stalk typically 4 mm thick or more
and not hollow; gills adnate to decurrent; gill tissue divergent . 22
21. Not as above; if both cap and stalk'viscid, then stalk typically slender, hollow, and fragile 23
22. Cap red to orange when young, at least at center.Hygrophorus speciosus, p. 126
22. Cap at first dark brown to olive-brown at center, the brighter colors developing in age and the
center often remaining brownish .Hygrophorus hypothejus, p. 126
23. Cap dark to bright red when fresh (but may fade in age or as it dries to orange or yellow) . 24
23. Cap orange to yellow (sometimes fading to whitish), never bright red . 31
24. Cap conical, up to 5 cm broad, bright red and not usually fading, viscid when moist; stalk also
red and usually viscid; found in eastern North America .Hygrocybe ruber
24. Not as above . 25
25. Cap and stalk distinctly viscid to slimy when moist; cap usually small or minute .
.Hygrocybe reai & others (see H. miniata, p. 113)
25. Not as above . 26
106 HYGROPHORACEAE

26. Cap distinctly conical when young and usually retaining a pointed umbo in age; stalk yellow
.Hygrocybe cuspidata (see H. acutoconica, p. 115)
26. Not as above . 27
27. Gills usually distinctly decurrent; cap typically small or minute (less than 4 cm broad) ... 40
27. Not as above . 28
28. Cap small (1-4 cm), not viscid, red to scarlet when moist but fading drastically as it dries to
orange or even yellow .Hygrocybe miniata & others, p. 113
28. Not as above (if cap not viscid, then cap not fading drastically or cap larger) . 29
29. Stalk white (base may be colored) .Hygrocybe laetissima (see H. punicea, p. 114)
29. Stalk yellow to orange or red . 30
30. Cap viscid when moist; stalk typically (0.5)1-2 cm or more thick, usually yellow or reddish fading
to orange-yellow; base of stalk often whitish .Hygrocybe punicea & others, p. 114
30. Cap not viscid or only slightly viscid; stalk typically 3-8 mm thick, usually red to reddish-orange
with a yellow to orange base .Hygrocybe coccinea & others, p. 114
31. Cap and stalk slimy or very viscid when moist; cap generally 4 cm broad or less . 32
31. Not as above (cap may be slimy, but stalk dry to only slightly viscid) . 35
32. Fruiting body showing pink, flesh-colored, or greenish (rarely bluish) tones somewhere on
fruiting body; gills not normally decurrent.Hygrocybepsittacina, p. 118
32. Not as above; fruiting body yellow to orange (but may show some white or have a lilac tinge) 33
33. Growing on rotting conifers . (see Mycena lilacifolia, p. 236)
33. Not as above; usually found on the ground . 34
34. Gills typically decurrent and cap usually depressed centrally .
.Hygrocybe nitida (see H.flavescens, p. 115)
34. Not as above .Hygrocybe chlorophana & others (see H.flavescens, p. 115)
35. Cap distinctly viscid or slimy when moist, 2-7 cm broad or more when expanded; gills not
decurrent . 36
35. Cap not viscid, or if slightly viscid then less than 3 cm broad or gills decurrent . 37
36. Cap distinctly conical, at least when young .Hygrocybe acutoconica, p. 116
36. Cap broadly convex to plane, not conical ..Hygrocybe flavescens & others, p. 115
37. Gills brilliant orange to yellow-orange, retaining their color even after the cap fades; common
in eastern North America, occasional in the Pacific Northwest Hygrocybe marginata, p. 112
37. Not as above; common and widespread . 38
38. Gills usually not decurrent; cap red to orange-red when fresh and moist (but fading).
.Hygrocybe miniata & others, p. 113
38. Not as above; gills usually decurrent . 39
39. Found in humus; cap yellow to orange-yellow (never scarlet, red, or orange) .
.Hygrocybe parvula & others (see H.flavescens, p. 115)
39. Not as above . 40
40. Stalk usually quite long (4-10 cm); found mainly in eastern North America on ground, in moss,
or on rotten wood .Hygrocybe cantharellus (see H. miniata, p. 113)
40. Not as above; found on ground, or if in lichen, moss, or wood, then stalk not so long .... 41
41. Typically found in humus or soil; cap red or scarlet when fresh.
.Hygrocybe subminiata (see H. miniata, p. 113)
41. Not as above; often found on wood, lichens, moss (see Omphalina & Xeromphalina, p. 221)
42. Cap sharply conical when young, usually retaining a pointed umbo in age, coral-pink to pink,
pinkish-orange, or salmon; gills pink .Hygrocybe calyptraeformis, p. 117
42. Not as above .43
43. Entire stalk or at least the lower part distinctly viscid to slimy when moist; cap also viscid 44
43. Stalk dry to slightly lubricous but not viscid; cap dry or viscid . 54
44. Cap small; stalk slender (usually less than 6 mm thick), hollow, not white; veil absent .... 45
44. Cap usually medium-sized to fairly large; stalk typically at least 5 mm thick, sometimes white;
veil present or absent . 47
45. Cap blackish to dark brown or grayish .Hygrocybe unguinosa (see H. psittacina, p. 118)
45. Not as above; cap brighter or lighter in color . 46
HYGROPHORACEAE 107

46. Cap sometimes green or greenish-tinged; gills typically adnate . Hygrocybepsittacina, p. 118
46. Green shades absent; gills often decurrent . Hygrocybe laeta (see H. psittacina, p. 118)
47. Cap dull yellowish becoming browner or grayer in age; stalk at least 1 cm thick; gills white or
developing greenish stains, but not yellow; found mainly under hardwoods in eastern North
America .Hygrophorus paludosus
47. Not as above .48
48. Cap (or center of cap) chestnut-brown to cinnamon-brown, reddish-brown, pinkish-tan,
salmon, or yellow-brown . 49
48. Cap olive-brown to grayish-brown, gray, blackish, etc. 51
49. Cap and gills rufous to rusty-orange, apricot, or salmon-buff; growing under oaks in eastern
North America .Hygrophorus subsalmonius
49. Not as above . 50
50. Odor fragrant, like almond extract . Hygrophorus variicolor (see H. bakerensis, p. 126)
50. Odor more or less mild . Hygrophorus laurae & others (see H. roseibrunnehs, p. 125)
51. Lower portion of stalk sheathed with gray to brown, olive-brown, or blackish fibrils, scales,
or granules; cap, gills, or stalk not yellow.Hygrophorus olivaceoalbus & others, p. 127
51. Not as above . 52
52. Gills creamy to yellow or orange; cap and stalk often exhibitng some yellow or orange tones
.Hygrophorus hypothejus, p. 126
52. Yellow and orange tones absent . 53
53. Cap evenly blackish to deep olive-brown when young; found mainly in eastern North America
.Hygrophorusfuligineus (see H. hypothejus, p. 126)
53. Cap paler when young or with a paler margin; widespread .
. Hygrophorus fuscoalbus & others (see H. hypothejus, p. 126)
54. Gills persistently bright orange to yellow-orange, even after cap fades; cap 1-5 cm broad, stalk
less than 6 mm thick . Hygrocybe marginata, p. 112
54. Not as above . 55
55. Cap green (citrine-green), at least when mature . ..Hygrocybe virescens, p. 118
55. Not as above . 56
56. Cap gray to grayish-brown, dark grayish-brown, olive, deep olive-brown, blackish, or umber;
gills and cap lacking violet or blue tones. 57
56. Cap differently colored (tan, reddish-brown, vinaceous, etc.), or if colored as above then cap
or gills with a violet, lilac, or bluish tinge when fresh . 63
57. Cap conical or with a pointed umbo, dull or dark gray.
.Hygrocybe acuta (see Camarophyllus recurvatus, p. 112)
57. Not as above . 58
58. Odor fragrant (like almond extract), but sometimes faint .
.Hygrophorus agathosmus & others, p. 128
58. Odor mild or not as above . 59
59. Base of stalk usually pale pinkish to pinkish-orange (especially interior) .
.(see Tricholoma saponaceum, p. 184)
59. Not as above . 60
60. Stalk sheathed by grayish to dark grayish-brown fibrils, scales, or granules .
. Hygrophorus inocybiformis 8c others (see H. olivaceoalbus, p. 127)
60. Not as above . 61
61. Fruiting body rather small (cap typically less than 5 cm broad, stalk typically less than 1 cm
thick and sometimes hollow) .Camarophyllus recurvatus & others, p. 112
61. Not as above; fruiting body medium-sized; stalk usually 1 cm thick or more . 62
62. Cap viscid to slimy when moist; gills white or pink . Hygrophorus calophyllus, p. 129
62. Cap viscid or dry, often appearing streaked; gills white or gray .
. Hygrophorus camarophyllus & others (see H. calophyllus, p. 129)
63. Stalk rooting deeply or with a “tap root” encased by grayish-brown volva-like material; cap
white becoming pinkish to vinaceous-red; gills white; known only from southern California
under oak; solitary or clustered .Hygrophorus marianae
63. Not as above; widespread and common . 64
108 HYGROPHORACEAE

64. Gills (or entire fruiting body) soon streaked or stained coral-red to vinaceous, pinkish-red, or
purple-red (especially in age), sometimes also with yellow stains.65
64. Not as above (but gills may be naturally pinkish) . 68
65. Associated with hardwoods (mainly oak and tanoak) .Hygrophorus russula, p. 123
65. Associated with conifers (mainly pine and spruce) . 66
66. Cap reddish at first but staining yellow and becoming pale to bright yellow in age; taste bitter;
especially common in the Rockies.Hygrophorus amarus (see H. purpurascens, p. 124)
66. Not as above (but fruiting body may yellow somewhat); widespread (including Rockies) . 67
67. Fibrillose veil present, at least when young .Hygrophorus purpurascens, p. 124
67. Veil absent .Hygrophorus erubescens & H. capreolarius (see H. purpurascens, p. 124)
68. Gills lilac or violet or pinkish to dingy flesh-colored, attached and sometimes slightly decurrent
but not deeply decurrent; stalk tough and fibrous, pliant, often fibrillose; cap often minutely
scaly or scurfy, not viscid; spores usually spiny . (steLaccaria, p. 171)
68. Not as above; spores smooth . 69
69. Gills and/ or cap with a violet, purple, or bluish tinge when fresh .
.Camarophyllus subviolaceus & others, p. 111
69. Not as above . 70
70. Gills brown to vinaceous-brown, adnate to decurrent; cap some shade of brown when moist,
fading as it dries; found with hardwoods in eastern North America Hygrophorus kauffmanii
70. Not as above; gills paler ... 71
71. Growing in grass; cap buff to tan, often umbonate; gills whitish; stalk thin, tough, pliant ....
.(seeMarasmius oreades, p. 208)
71. Not as above . 72
72. Cap 4.5 cm broad or less, translucent-striate when moist . 73
72. Not as above; cap typically opaque . 74
73. Stalk 4-7 mm thick; cap vinaceous-brown to pinkish-gray or buff, slightly viscid when moist;
gills pinkish-gray or paler, distinctly decurrent; terrestrial .
. Camarophyllus colemannianus (see C. subviolaceus, p. Ill)
73. Not as above; growing on ground or wood . (see Tricholomataceae, p. 129)
74. Stalk 1 -3 cm thick, the apex distinctly scurfy (punctate); cap 5-20 cm broad, pale tan to pinkish,
flesh-colored, pinkish-orange, or occasionally whitish; gills usually tinged pink(but sometimes
whitish); cap viscid when moist, the margin inrolled until maturity; odor not like almond
extract; associated with conifers (especially spruce) .Hygrophoruspudorinus, p. 124
74. Not with above combination of characteristics . 75
75. Gills distinctly paler than center of cap (white to creamy or pale yellow) . 76
75. Gills colored like the cap or slightly paler or darker (pinkish to salmon, etc.) . 79
76. Odor faintly fragrant (like almond extract) to strongly aromatic . 77
76. Not as above; odor usually mild or somewhat potato-like . 78
77. Odor like almond extract . Hygrophorus bakerensis & others, p. 126
77. Odor strongly aromatic (but not as above) Hygrophoruspacificus (see H. bakerensis, p. 126)
78. Cap pink or rose (but may fade!), stalk lacking sordid yellowish patches below; found under
mountain conifers, often near melting snow Hygrophorus goetzii(see H. pudorinus, p. 124)
78. Not with above features .Hygrophorus roseibrunneus & many others, p. 125
79. Odor sickeningly sweet; fruiting body pinkish-buff to pinkish-cinnamon; cap not viscid .
. Camarophyllus graveolens
79. Not as above . 80
80. Cap whitish to buff, sometimes with darker spots or zones; odor often faintly fruity (if kept in
a closed space); gills pinkish-cinnamon to salmon or ochre-salmon; associated with conifers
(but not redwood); gill hyphae divergent . Hygrophorus saxatilis (see H. pudorinus, p. 124)
80. Not with above features . 81
81. Cap whitish when young Hygrophorus albicastaneus & others (see H. roseibrunneus, p. 125)
81. Cap not white when young (but may be quite pale in age or after fading). 82
82. Gills adnexed to adnate or slightly decurrent; stalk 1 -4 cm thick, cap 5-13 cm broad; odor mild or
radish- to cucumberlike; gill hyphae divergent; usually associated with oak .
.Hygrophorus nemoreus(see Camarophylluspratensis, p. 110)
82. Not as above; gills usually decurrent .Camarophyllus pratensis, p. 110
Left: Hygrophorus subalpinus is a robust whitish conifer-loving waxy cap (see description on p.
121). Note the slight annulus (ring) on specimen at left. Right: Camarophyllus borealis and its close
relatives are white to yellowish with a slender build, decurrent gills, and little or no odor.

Camarophyllus borealis (Snowy Waxy Cap)

cap l -5 cm broad, convex or obtusely umbonate, often expanding in age to plane or
depressed; surface smooth, moist or lubricous but not viscid, watery white to dull white.
Flesh thin, soft, white, odor mild. GILLS usually decurrent, well-spaced, thick, soft,
somewhat waxy; white. STALK 2-9 cm long, 2-5 (8) mm thick, equal or tapering down¬
ward, smooth, firm, dry, often curved or sinuous, white. VEIL absent. SPORE PRINT
white; spores 7-9 (12) x 4.5-6.5 microns, elliptical, smooth. Gill tissue interwoven.
HABITAT: Scattered or ingroups onground and humus in woods or at their edges, widely
distributed. In our area this species and its relatives C. virgineus and C. niveus (see com¬
ments) are fairly common throughout the mushroom season, but seldom occur in the
large numbers typical of Hygrophorus eburneus. The best fruitings usually occur in the
winter, often in relatively dry weather.
EDIBILITY: Edible but fleshless, flavorless, and savorless.
COMMENTS: Also known as Hygrophorus borealis, this is one of several rather small,
whitish waxy caps with a dry to slightly viscid (not slimy) cap, slender non-viscid stalk,
and interwoven gill tissue. Others include: C. niveus, common, cap slightly viscid and
slightly striate when moist; C. virgineus, especially common in California, cap dry and
usually tinged yellow in age or dry weather, and spores 8-12 microns long; C. angustifolius,
dull white with spores 5-8 microns long; C. cremicolor, with pale yellow gills when young;
and three species with parallel gill hyphae and non-decurrent gills: Hygrocybesubaustrali-
ga, small, cap scarcely viscid, gills notched or adnexed, fairly common;//, albinella, cap
white and conical at first, stalk dry, rare; and H.fornicata, with a grayish-tinged viscid cap.
All of these have been placed in Hygrophorus. None have the slimy cap and stalk of Hygro¬
phorus eburneus. Alboleptonia sericella and Inocybe geophylla can be similar but have
pinkish and brown spores respectively. See also the undescribed Marasmius on p. 206.

Camarophyllus russocoriaceus (Cedar Waxy Cap)

CAP 1-3 (5) cm broad, convex to plane or slightly umbonate to slightly depressed; surface
smooth, not viscid, white or often tinged pale tan to yellowish, especially at the center. Flesh
whitish, thin; odor fragrant when fresh (like arborvitae or cedar). GILLS well-spaced,
usually decurrent, thick, slightly waxy, white or whitish. STALK 3-10 cm long, 2-5 (8) mm
thick, equal or tapered downward, often long in relation to cap; dry, smooth, colored more
or less like cap, often curved or sinuous. VEIL absent. SPORE PRINT white; spores 6.5-9 x
4-6 microns, elliptical, smooth. Gill tissue interwoven.
Camarophyllus russocoriaceus is a slim waxy cap that smells like cedar or Russian leather. Note
off-white to buff-colored cap and long, slender stem.

HABITAT: Widely scattered or in small groups in woods or at their edges, known only
from the west coast (in addition to Europe). Common in our area in the fall and winter,
especially at higher elevations in the coastal mountains, but rarely in large numbers.
EDIBILITY: Not recommended—it has a slightly medicinal flavor, at least raw.
COMMENTS: The piquant cedarlike odor distinguishes this plain-looking waxy cap
from C. borealis Sind other look-alikes. Other species: Hygrophoruspusillus is a somewhat
similar species with a viscid cap that is usually tinged cream or pale brownish-flesh-color.
It has a slight fragrant odor, divergent to nearly parallel gill tissue, and occurs in groups or
troops under conifers in northern California and the Pacific Northwest.

Camarophylluspratensis (Meadow Waxy Cap)

CAP 2-9 (10) cm broad, obtuse to broadly convex becoming broadly umbonate or plane to
depressed in age; surface smooth or sometimes cracking in age (especially at center), not
viscid; rufous to dull orange or pale dull orange, fading to salmon-buff, pinkish-tan, buff,
tawny, or even whitish; margin often wavy. Flesh white or tinged cap color; odor mild.
GILLS typically decurrent, same color as cap or paler (usually pale dull orange or salmon-
buff to nearly white); well-spaced, thick, broad, somewhat waxy. STALK 3-10 cm long,
0.5-2 cm thick, whitish or tinged cap color, equal or tapering downward, dry, smooth or
fibrillose. VEIL absent. SPORE PRINT white; spores 5.5-8 x 3.5-5 microns, elliptical to
nearly round. Gill tissue interwoven.
HABITAT: Solitary to scattered or gregarious in damp places—common and very widely
distributed—from Europe to Iceland to South America. In our area it fruits mainly in
the winter under redwood, but as its name implies (pratensis-iields), it can also occur in
open or grassy situations.
EDIBILITY: Edible and rated highly in Europe (one author even puts in on a par with
morels!). However, my own experience with it gave me no great cause for enthusiasm.
Unlike the similarly-colored chanterelle, it is readily attacked by maggots.
COMMENTS: Better known as Hygrophorus pratensis, this common waxy cap is ex¬
tremely variable in size and shape, but can generally be recognized by its pale, dull orange

110
Camarophylluspratensis is variable in both shape and color, but the gills are nearly always decurrent
and widely spaced, and the cap is never slimy.

color and decurrent gills. The cap and stem are not viscid, but the gills are thick and rather
waxy. Robust individuals are slightly reminiscent of chanterelles (Cantharellus cibarius),
but differ in their duller color and broad, well-developed gills with acute rather than blunt
edges. A similarly colored species, Hygrophorus nemoreus, is fairly common in our area
with oak in the fall and winter. It is larger and fleshier than C.pratensis{ cap5-l 3 cm broad,
stalk 1.5^1 cm thick) and has a slightly viscid to dry, orangish or cinnamon-orange cap,
adnexed to adnate or slightly decurrent gills, a mild to radish- or cucumberlike odor, and
divergent gill tissue. It is large enough and common enough to be worth sampling, but
I can find no information on its edibility.

Camarophyllus subviolaceus (Violet-Gray Waxy Cap)

CAP 2-6 cm broad, broadly convex to plane or slightly depressed with an uplifted margin;
surface slightly viscid when moist, hygrophanous: violet-gray to violet-brown at least at the
margin when moist (center often paler), fading as it dries to gray or pallid; margin trans¬
lucent striate when fresh. Flesh rather thin, colored like cap or paler; taste mild to bitter or
slightly acrid. GILLS decurrent, whitish soon becoming smoky-violet or gray with a violet
tinge; well-spaced, thick, soft, rather waxy. STALK 3-7 cm long, 0.4-1 cm thick, equal or
tapered downward, dry, smooth, white or tinged cap color, often curved. VEIL absent.
SPORE PRINT white; spores 6-8 * 4-6 microns, elliptical, smooth. Gill tissue interwoven.

Camarophyllus subviolaceus has widely spaced decurrent gills that are violet-gray when fresh.
112 HYGROPHORACEAE

HABITAT: Scattered or in small groups in woods and swamps throughout northern North
America, late summer through early winter. I have found it several times in the
Pacific Northwest but it is not particularly common.
EDIBILITY: Unknown.
COMMENTS: Better known as Hygrophorus subviolaceus, this beautiful species has
been included because its violet-tinged gills are unusual for a waxy cap. There are several
similarly-colored species, including: C. pallidas, with smaller, round or nearly round
spores and a viscid, violet-gray cap; C. angelesianus, with amyloid spores and a viscid
cap that does not fade appreciably, usually found at high elevations in the spring and
summer; C. rainierensis, with a strong green corn odor and slightly viscid cap; and C.
cinereus, with a non-viscid cap, mild taste, and larger spores. Other species include:
C. colemannianus, closely related, but with a viscid, brown to pinkish-gray to buff cap
and pinkish-gray to pale buff colors; Hygrocybe caerulescens, a bluish-tinged species
with parallel gill tissue; and Hygrocybe purpureofolia, with lavender to dull purplish
gills and a brown cap that fades to yellow-orange, found in eastern North America.

Camarophyllus recurvatus (Little Brown Waxy Cap)

CAP l -3 cm broad, convex becoming plane or slightly depressed, sometimes with a small
pointed umbo; surface smooth or cracking into small scales, dry to slightly viscid, dark to
pale olive-brown or at times pallid; margin often wavy or pleated. Flesh thin, olive-brown,
odor mild. GILLS decurrent, well-spaced, broad, thick, soft and rather waxy, white or
grayish-white. STALK 2-5 cm long, 3-6 mm thick, equal or tapering downward, smooth,
not viscid, whitish or colored like cap. VEIL absent. SPORE PRINT white; spores 7-10 * 4-
6 microns, elliptical, smooth. Gill tissue interwoven.
HABITAT: Scattered to gregarious under conifers and in open, grassy areas; widely
distributed. I’ve seen large fruitings in northern California but have yet to find it in our area.
EDIBILITY: Unknown.
COMMENTS: Also known as Hygrophorus recurvatus, this is one of a number of
undistinguished, brown to grayish waxy caps that qualify as “LBM’s.” As they are unlikely
to interest the average collector, only a few others are worth mentioning: Hygrocybe ovina,
with deeply notched to adnate gills and a convex to plane, grayish-brown cap that often has
a paler or yellower margin; Hygrocybe nitrata, with a nitrous odor; Camarophyllus
paupertinus, with a small (1-2 cm) cap and exceedingly strong, disagreeable odor (known
only from northern California under redwoods); Hygrocybe acuta, with a sharply conical,
dull gray to gray-brown cap, growing under conifers on the west coast; and Hygrocybe
atro-olivacea, with a minutely scaly cap that is dark at the center or throughout.

Hygrocybe marginata (Orange-Gilled Waxy Cap)

CAP 1 -5 cm broad, conical or convex becoming umbonate, plane, or with uplifted margin;
surface smooth, hygrophanous but not viscid: deep yellow to yellow-orange or orange
when moist, sometimes with an olive tinge, fading to pale yellow or whitish as it dries. Flesh
thin, fragile, waxy. GILLS slightly decurrent to adnate, adnexed, or even free; well-spaced,
broad, thick, soft, waxy; brilliant orange or at times orange-yellow, not fading. STALK
4-10 cm long, 3-6 mm thick, more or less equal, smooth, fragile, not viscid; pale orange to
buff or pale yellow-orange, often curved; hollow. VEIL absent. SPORE PRINT white;
spores 7-10 * 4-6 microns, elliptical, smooth. Gill tissue parallel to somewhat interwoven.
HABITAT: Solitary to scattered or in small groups in humus and on very rotten wood,
under both hardwoods and conifers; common in eastern North America in summer and
fall, known also from the Pacific Northwest and to be expected in northern California.
I have not seen it in our area.
HYGROCYBE 113

EDIBILITY: Edible but of negligible substance and flavor.

COMMENTS: The gills’ ability to retain their bright orange color long after the cap has
faded is the outstanding attribute of this modest waxy cap, also known as Hygrophorus
marginatus. The cap is at times slightly greasy or tacky to the touch, but never truly viscid.
Several color forms occur, including one with an olive cap (fairly common in Washington)
and another with yellow gills (in eastern North America).

Hygrocybe miniata (Miniature Waxy Cap)

CAP l -4 cm broad, convex to plane or slightly depressed; surface smooth or minutely scaly,
hygrophanous but not viscid: bright red to scarlet when moist, quickly fading as it dries to
orange and finally yellow. Flesh very thin, waxy, colored more or less like cap. GILLS
adnexed to adnate or very slightly decurrent, soft, waxy, thick, broad, red to orange,
yellow, or peachy (fading like cap). STALK 2-5 (8) cm long, 2-4 mm thick, equal, smooth,
dry, red to orange or yellow (fading like the cap but more slowly). VEIL absent. SPORE
PRINT white; spores 6-10 * 4-6 microns, elliptical, smooth. Gill tissue parallel.
HABITAT: Solitary, scattered, or in groups or small tufts on ground, rotting logs, or in
moss; widely distributed and common. In our area it fruits in the late fall and winter in a
variety of habitats, but usually in deep shade. The largest fruitings I’ve seen were in a
dense stand of pole-size redwood saplings and in a plot of cypresses mixed with oaks.
EDIBILITY: Edible but of negligible substance and according to most sources, bland.
However, Captain Charles Mcllvaine says: “The gunner for partridges will not shoot
rabbits; the knowing toadstool-seeker will pass all others where H. miniatus abounds.”
COMMENTS: The small size, waxy gills, and convex to plane, non-viscid cap that fades
markedly from red to yellow or orange are the far-from-infallible fieldmarks of this dainty
fungus. It is better known as Hygrophorus miniatus. The cap is neither viscid nor conical,
and is often minutely scaly (squamulose). Its frequent occurrence on rotten wood
is unorthodox behavior for a waxy cap. Other species: H. squamulosa has thicker, firmer
flesh but is otherwise similar. H. cantharellus is very similar but has a longer stem and de¬
current gills; H. moseri has a less scaly cap and decurrent gills. H. subminiata has a
minute (up to 1.5 cm broad), scarlet to orange, slightly viscid cap plus decurrent, whitish to
pale yellowish gills and a very thin (about 2 mm) stalk; it is fairly common in our area,
especially under redwood. H. turunda is a widespread miniature northern species with

Hygrocybe miniata has a bright red cap that fades to orange or yellow as it loses moisture. Note
small size (the largest is 2 cm broad) and convex (not conical) cap.
114 HYGROPHORACEAE

minute brown to grayish squamules (scales) on a red to yellow background. There are also
several minute species with a slimy-viscid cap and stalk when fresh, including: H. reai, with
a bitter-tasting cap; H. minutula, with a stalk that fades to yellowish in age; and H. sub-
minutula, whose stalk scarcely fades from red. All of the above species have a red to scarlet
cap when moist, and all were originally placed in Hygrophorus. None are worth eating.

Hygrocybe coccinea (Righteous Red Waxy Cap) Color Plate 20

CAP 1.5-4 (6) cm broad, obtusely conical or convex becoming plane or slightly umbonate;
surface smooth, dry or tacky, deep red, blood-red, or bright red when fresh, fading some¬
what in age or developing paler streaks or splotches. Flesh thin, reddish to orange, waxy.
GILLS adnate to adnexed or free, reddish to orange or peachy, or red with yellow edges;
thick, broad, soft, waxy. STALK 3-7 (10) cm long, 3-8 mm thick, equal, smooth, not viscid,
hollow; usually red to reddish-orange with a yellow base (but base sometimes appearing
whitish from mycelium); typically not fibrillose-striate. VEIL absent. SPORE PRINT
white; spores 7-10.5 x 4-5 microns, elliptical, smooth. Gill tissue parallel.
HABITAT: Solitary, scattered, or in small groups in woods and other wet places; widely
distributed, but not as common in North America as H. punicea. Like H. punicea, it is
frequent in our area in the winter under redwoods or in mixed woods. Curiously, in some
regions (such as England) it grows mainly in open fields!
EDIBILITY: Said to be edible, but easily confused with H. punicea. I haven’t tried it.
COMMENTS: Also known as Hygrophorus coccineus, this exquisite bright red waxy cap
ranks among our most beautiful mushrooms and is definitely worth seeking out. It is
redder than H. miniata and does not fade nearly as drastically. It is often confused with
H. punicea, but that species is usually more robust, has a distinctly viscid cap (when moist),
and most often has a yellow to orange, fibrillose-striate stem with a whitish base rather
than a red stem with a yellow base (see color plates 20 and 21). Other species: H. marchii
is a similar species with a viscid red cap that soon dries out and fades to orange or yellow-
orange. Like H. coccinea, it is a rather small, slender species (stalk 3-6 mm thick) that calls
to mind H.flavescens but for its color. It is widely distributed but I have not seen it locally.

Hygrocybe punicea (Scarlet Waxy Cap) Color Plate 21

CAP 2.5-12 (14) cm broad, obtusely conical or convex when young, then plane to broadly
umbonate or with uplifted margin in age; surface smooth, viscid or lubricous when moist,
deep red to bright red, fading (often in streaks or splotches) to reddish-orange and finally
orange. Flesh rather thin, waxy, watery reddish-orange to yellow-orange. GILLS adnate to
adnexed or free, reddish-orange to yellow or peachy, well-spaced, thick, broad, soft, waxy.
STALK 3-12 cm long, 0.5-2 cm thick, equal or narrowed at base, smooth, not viscid, often
fibrillose-striate; yellow or sometimes red soon fading to orange or yellow; base usually
whitish but sometimes yellow. VEIL absent. SPORE PRINT white; spores 7-12 x 4-6
microns, elliptical to oblong, smooth. Gill tissue parallel.
HABITAT: Scattered to gregarious in cool, damp places (usually in woods); widely
distributed and fairly common in our area in the winter and early spring, especially under
redwood. It is usually the last waxy cap to appear (mid-January or later).
EDIBILITY: Listed as edible in most books, but poisonous at least to some people! I know
two toadstool-testers who had very unpleasant experiences with it. It is an efficient con¬
centrator of the malleable metallic element cadmium, which is decidedly deleterious
when consumed on a regular basis.
COMMENTS: Better known as Hygrophoruspuniceus, this beautiful mushroom is as
easy to recognize as it is difficult to overlook. The bright red sticky cap and indisputably
HYGROCYBE 115

waxy gills make it stand out vividly in the dim, damp milieu where it thrives. It is just as
common in our wintertime woods as H. coccinea, but much more conspicuous because of
its larger size. Fora comparison of the two, see comments under H. coccinea. Other species
in the H. punicea “complex” include: H. laetissima, with an even brighter red cap plus a
white stalk when young; H. splendidissima, stalk white becoming reddish-striate in age;
and H. aurantiosplendens, whose cap fades more markedly. All of these favor redwoods
in California and could just as well be treated as variations of H. punicea.

Hygrocybe flavescens (Golden Waxy Cap) Color Plate 22

CAP 2-7 cm broad, broadly convex becoming plane or with margin slightly uplifted;
surface smooth, viscid when moist, bright lemon-yellow to golden-yellow (or sometimes
orange toward the center). Flesh thin, yellow, waxy. GILLS typically adnexed or free,
soft, thick, waxy, yellow or pale yellow. STALK 4-9 cm long, 3-10 mm thick, equal,
smooth, sometimes tacky or slightly viscid but not slimy; easily splitting, often grooved,
yellow to yellow-orange with a whitish base. VEIL absent. SPORE PRINT white; spores 7-
9 x 4-5 microns, elliptical, smooth. Gill tissue parallel.
HABITAT: Solitary to widely scattered or in small groups on ground, usually in woods,
widely distributed. Common in our area in the winter, especially under redwood but also
with oak, madrone, etc. It usually peaks well after H. acutoconica has finished fruiting,
sometimes when the bright red waxy caps (H. punicea and H. coccinea) appear.
EDIBILITY: Edible, but far from incredible: it is watery and has little substance or taste.
One author says it “would make a colorful and novel addition to a salad.” So would a
banana slug.
COMMENTS: The bright yellow to lemon-yellow color, convex to plane cap, and waxy
gills identify this beautiful mushroom, better knownasHygrophorusflavescens. The cap is
never red as in H. punicea, nor is it conical as in H. acutoconica. In rainy weather the
surface of the cap can be quite viscid or even slippery—giving it the appearance of a just-
licked lollipop—but in dry weather it is often somewhat silky or shiny. Other widely dis¬
tributed yellow to orange species with non-conical caps include: H. chlorophana, often
confused with H. flavescens, but slightly smaller with a distinctly slimy stalk when moist;
H. ceracea, with a slightly viscid cap and stalk and adnate to decurrent gills; H. citrino-
pallida, with a slimy-viscid cap and stalk that fade from yellow to whitish and gills that
retain their yellow color; H.flavifolia, with a slimy-viscid cap and white, slimy-viscid stalk;
H. parvula, with a non-viscid stalk and decurrent gills; and H. nitida, always yellow when
fresh, with a centrally depressed cap (even when young), gills that are soon deeply de¬
current, and a frequently viscid stalk. The latter species is widely distributed but espe¬
cially common in mossy or boggy areas in eastern North America. In both H. nitida and
H.flavifolia the cap often fades to creamy or whitish as it loses moisture or ages.

Hygrocybe acutoconica (A Cute Conic Waxy Cap)

CAP 2-7 (10) cm broad, bluntly or acutely (sharply) conical when young, sometimes
expanding in age but usually retaining pointed umbo; surface smooth, viscid or slimy when
moist, bright yellow to yellow-orange or orange (the orange usually toward center). Flesh
soft, thin, waxy, yellow. GILLS adnexed to free, thick, soft, broad, waxy, yellow, never
blackening. STALK 5-8 (12) cm long, 3-6(10) mm thick, equal or thicker below, usually
longitudinally striate and/or twisted, easily splitting; smooth, moist or slightly viscid,
yellow to yellow-orange, but usually white at base; not blackening when handled, but
base may bruise or age grayish to nearly black. VEIL absent. SPORE PRINT white; spores
9-15 x 5-9 microns, elliptical, smooth. Gill tissue parallel.
Hygrocybe acutoconica. This common yellow to orange waxy cap does not blacken when handled.
The cap is sharply conical when young, but eventually expands as shown at left.

HABITAT: Scattered to gregarious on ground in woods or undertrees, widely distributed.

In our area it fruits in the fall and early winter, mostly under redwood and oak. It is nearly
as common as its blackening counterpart, H. conica, but has a shorter season.
EDIBILITY: Harmless, fleshless, flavorless (see comments on edibility of H.flavescens).
COMMENTS: Better known as Hygrophorus acutoconicus, this cute, conic waxy
cap differs from H. conica in its yellowish gills and “failure” to blacken when handled.
Scientists may scoff at our tendency to anthropomorphize (attribute human qualities to
inhuman beings or inanimate objects), yet our language leaves us little choice. For instance,
the word “failure” (with its attendant implications of inadequacy, its connotation of
“attempting to, but not succeeding”) is often utilized by taxonomists to denote “absence.”
Before Fm accused of lending credence to this trend, let me earnestly put this question to
you philosophical few among the myopic many (or you fungophilic few among the
mycophobic many): Is the “failure” in this case an innate inability to succeed? An admirable
example of genetic (or aesthetic) restraint? A coincidental and pointless byproduct of
circumstance? Or none of the above? ... Other species: H. langei and H. aurantiolutescens
are very similar if not the same; H. cuspidata (called H. persistens by some) is also similar
but has a conical red cap when fresh that fades with age, plus an orange to yellow stem. It
is widely distributed, but I have yet to find it south of San Francisco.

Hygrocybe conica (Witch’s Hat; Conical Waxy Cap) Color Plate 19

CAP 1-5(12)) cm broad, bluntly to sharply conical, sometimes expanding in age but usually
retaining a pointed umbo; surface smooth, dry to moist or slightly viscid, sometimes red
but more often orange, yellow, or olive-yellow; blackening in age or upon handling; margin
sometimes uplifted in old age. Flesh thin, waxy, blackening in age. GILLS adnexed or
free, thick, broad, soft, waxy, usually whitish but sometimes tinged yellow, soon becoming
grayish and finally black. STALK (2) 4-20 cm long, 0.3-1 (1.5) cm thick, equal, not viscid
or only slightly so, usually striate and/ or twisted, hollow in age and easily splitting; pallid to
yellow, olive-yellow, orange, or red, with a whitish to gray base, but turning gray or black
when handled or in age. VEIL absent. SPORE PRINT white; spores 8-14 * 5-7 microns,
elliptical, smooth. Gill tissue parallel.
HABITAT: Solitary to scattered or gregarious on ground in damp places (usually in
woods); widely distributed and very common, but rarely fruiting in large numbers. It is
known from a wide variety of habitats in North America, but in our area shows a pro¬
nounced preference for redwood and cypress. It fruits in the falland winter and, along with

116
Hygrocybe conica (better known as Hygrophorus conicus) is one of our most common waxy caps.
Note conical cap (which may expand somewhat in age) and tendency to blacken.

H. acutoconica, is usually the first of the brightly colored waxy caps to appear. Bob
Winter says it often grows on lawns in Fresno, California.
EDIBILITY: Not recommended. It was once considered poisonous, perhaps due to its
blackening qualities, but also because four deaths in China were (mistakenly?) attributed
to it. Now it’s generally regarded as harmless, but Larry Stickney, chef extraordinaire,
meandering mainstay of the Mycological Society of San Francisco, and an inimitable
mushroom-loving marvel of a man, says it “elicited an odd sensation of lightheadedness
and numbness” when he tried it. At any rate, it hardly seems worth experimenting with
such a thin-fleshed, watery, tasteless morsel.
COMMENTS: The conical cap and tendency of all parts to blacken when handled im¬
mediately identify this cosmopolitan mushroom. It is likely to be among the first waxy caps
encountered by beginners and is quite beautiful and waxy-looking when growing in the
woods. By time it is brought home, however, it is often barely recognizable because of the
gray and black stains that develop. Old, withered, completely black speimens are
sometimes found growing alongside vividly colored fresh ones. When growing in deep
humus (e.g., redwood duff) the stalk is often quite long, but when growing in bare soil or
grass or shallow humus it is apt to be shorter and/ or siouizr,Hygrophorus conicus is a syn¬
onym. The “splitters” recognize several closely related blackening species, including: Hy¬
grocybe nigrescens, with a bluntly conical, red to scarlet cap, often found under oak;//.
singeri, cap and stalk distinctly viscid; and H. olivaceoniger, small, thin, and olive-green.

Hygrocybe catyptraeformis (Salmon Waxy Cap) Color Plate 23

CAP 2.5-7 cm broad, acutely (sharply) conical at first, then expanding somewhat but
retaining a pointed umbo; surface dry or slightly viscid, smooth, coral-pink to pink to
salmon when young (rarely tinged lavender), paler (or pinker or oranger, sometimes
with whitish areas) in age; margin often splitting at maturity and sometimes uplifted. Flesh
thin, watery pinkish, waxy. GILLS adnate to adnexed or even free, thick, soft, waxy, pink
to pale pink. STALK 3-16 cm long, 3-8 (10) mm thick, usually rather long and slender,
equal, smooth, often longitudinally striate and/or twisted; fragile, hollow, and easily
splitting, white or tinged pinkish (or rarely lavender), not truly viscid. VEIL absent.
SPORE PRINT white; spores 6-9 * 4.5-6 microns, elliptical, smooth. Gill tissue parallel.

117
118 HYGROPHORACEAE

HABITAT: Solitary to widely scattered or in small groups on ground in woods or at their

edges (often rooted deeply in the humus); widely distributed but not common. I have found
it only twice in our area, under redwoods, in the winter.
EDIBILITY: Edible.
COMMENTS: In my fickle fungal opinion, this striking species is one of the most beautiful
and elegantof all the waxy caps, its rarity serving to accentuate its beauty. In a genus noted
for its bright colors, this species still manages to stand out—its pointed pink to
salmon cap and pink gills render it distinct. H. psittacina and H. laeta can be pinkish, but
are smaller, usually slimy, and are never as sharply conical. Nolanea salmonea of eastern
North America is also similar, but gives a pinkish spore print and does not have waxy gills.
Hygrophorus calyptraeformis is a synonym.

Hygrocybe virescens (Lime-Green Waxy Cap)

CAP 2-5 cm broad, obtusely conical or convex becoming plane, umbonate, or with uplifted
margin in age; surface smooth, moist or tacky but not truly viscid; color variable when
young (honey-yellow or green mixed with dull orange, etc.), but soon becoming citrine-
green to yellow-green (“lime-green”) overall. Flesh thin, fragile, greenish, waxy. GILLS
adnexed to free, whitish or with citrine-green tints, thick, soft, waxy. STALK 3-7 cm long,
3-8 mm thick, more or less equal, smooth, hollow in age, not viscid; lime-green with a
whitish base. VEIL absent. SPORE PRINT white; spores 7-10 x 5-6.5 microns, elliptical,
smooth. Gill tissue parallel.
HABITAT: In scattered groups or tufts under redwood, northern California, apparently
quite rare. I have seen only one substantial fruiting—in December of 1971 at Van Damm
State Park in Mendocino County. Subsequent expeditions to the same area failed to turn
it up.
EDIBILITY: Unknown.
COMMENTS: The striking “lime-green” color of the cap and stalk make this one of the
easiest of all waxy caps to identify. Unfortunately, it is one of the most difficult of all waxy
caps to find! The above description is adapted from that of Alexander Smith, who
originally found it near Trinidad, California. Whereas many waxy caps are cosmopolitan
(particularly Hygrocybes), this one appears to have a sharply limited habitat and
geographical distribution. The only other green waxy cap, H. psittacina, is smaller,
slimmer, and slimier, and is green when young rather than in age.

Hygrocybe psittacina (Parrot Waxy Cap) Color Plate 18

CAP 1-3 cm broad, bell-shaped or convex to broadly umbonate or plane in age; surface
smooth, slimy or viscid when moist, usually shiny when dry; color extremely variable: at
first dark green to bright green or olive-green, but soon fading to some shade of yellow,
pink, orange, rufous, vinaceous, ochre-buff, tawny, etc.; margin translucent-striate when
moist. Flesh thin, soft, waxy, odor mild. GILLS adnate to very slightly decurrent, but
sometimes seceding; well-spaced, soft, thick, waxy; at first greenish, then fading like the
cap (but often yellower or redder), and often retaining slight greenish tints. STALK 2-6 (8)
cm long, 2-5 mm thick, equal or tapering upward, hollow, smooth, very slimy or viscid
when moist, greenish when young but soon fading to yellow or cap color (pink, orange,
etc.). VEIL absent. SPORE PRINT white; spores 6.5-10 x 4-6 microns, elliptical, smooth.
Gill tissue parallel.
HABITAT: Solitary to scattered or in small groups in damp soil, moss, humus, etc.; widely
distributed. Fairly common in our area in late fall and winter but easily overlooked. Like
HYGROCYBE 119

most of our Hygrocybes it favors redwood, but also occurs under other trees as well as on
mossy roadbanks or in grass. The largest fruiting I’ve seen was in a swampy hardwood
forest in Pennsylvania.
EDIBILITY: Edible, but slimy and insubstantial. Raw specimens make a colorful but
slippery supplement to salads, sliding down your throat before you can savor them.
COMMENTS: Also known as Hygrophorus psittacinus, this slippery little fungus is a
cinch to recognize when young and fresh—there is no other small green agaric with a
glutinous cap and stem! As it dries out, however, it changes color drastically, and faded
specimens have been known to fool the most seasoned Hygrocybe-hound. The rate,
extent, and nature of the color changes seem to vary according to environmental condi¬
tions, but close inspection will often reveal a faint olive tinge somewhere on the fruiting
body. Even when there is no green present, the small size and slimy-viscid stalk will dis¬
tinguish it from all but H. laeta (see below). The stalk can be so slimy that it is difficult to
pluck. Other species: H. psittacina var. californica is a rare fungus with a beautiful blue
rather than gorgeous green cap in youth. H. laeta is a widely distributed species of variable
color (violet-gray to pinkish, orange, vinaceous, etc.), but is never green when young and
usually has decurrent gills. It, too, has a slimy-viscid cap and stalk. H. unguinosa has a
slimy-viscid, gray to dark brown to nearly black cap and stem, and is also widely dis¬
tributed. For viscid-stalked Hygrocybes with bright yellow to orange or red caps, see
comments under H. flavescens and H. miniata.

Hygrophorus chrysodon (Flaky Waxy Cap) Color Plate 17

CAP 2.5-8 (10) cm broad, convex to plane or broadly umbonate; surface smooth, viscid
when moist, white except for delicate yellow to golden-orange flakes or granules on the
margin (or sometimes tinted yellow throughout); margin at first inrolled. Flesh thick, soft,
white. GILLS typically decurrent, well-spaced, white, soft, rather waxy. STALK 3-10 cm
long, 0.5-2 cm thick, equal, viscid when moist, white except for a ring of yellow to golden-
orange granules at apex (or in rainy weather, the granules dispersed throughout). VEIL
evanescent, leaving slime on stalk. SPORE PRINT white; spores 7-10 * 3.5-5 microns,
elliptical, smooth. Gill tissue divergent.
HABITAT: Solitary to gregarious in woods; widely distributed, but not very common
in our area. It is said to favor conifers, but in coastal California it shows a definite pre¬
ference for madrone and tanoak, fruiting sporadically during the late fall and winter.
EDIBILITY: Edible, but slimy and bland (for more details see H. sordidus).
COMMENTS: The exquisite flakes of gold on the cap margin and/ or stalk apex (see color
plate!) are unique to this beautiful species. However, rain may obliterate the granules or
disperse their yellow pigment over the entire cap and stem surfaces, leading to confusion
with H. gliocyclus, H. eburneus, and other white or yellowish waxy caps.

Hygrophorus eburneus (Ivory Waxy Cap)

CAP 2-7 (10) cm broad, obtuse to convex becoming broadly umbonate to plane or with
uplifted margin; surface smooth, extremely slimy or viscid when wet, pure white or some¬
times slightly yellowish in old age or occasionally with small pinkish spots; margin at first
inrolled. Flesh soft, white, odor mild. GILLS adnate to decurrent, well-spaced, thick, soft,
waxy, pure white to very slightly yellowish in old age. STALK 4-15(18) cm long, 0.3-1 (2)
cm thick, equal or tapered toward base, usually rather slender; smooth or with punctate
apex, slimy-viscid when moist, pure white, but sometimes discoloring slightly yellowish or
pinkish in age. VEIL evanescent, depositing slime on stalk. SPORE PRINT white; spores
6-9 * 3.5-5 microns, elliptical, smooth. Gill tissue divergent.
Hygrophorus eburneus. Sometimes called “Cowboy’s Handkerchief,” this common pure white waxy
cap is easily told by its slimy cap and stem. In wet weather the cap is coated with such a thick layer
of slime that it looks as if someone blew their nose on it.

HABITAT: Scattered to gregarious or tufted on ground in woods, very widely distributed

but most common on the west coast. In our area it is mycorrhizal with hardwoods and is the
most common fall and winter Hygrophorus of our oak-madrone woodlands. In other
regions it may grow with conifers, e.g., in Oregon it is often abundant in oak-pine forests
and in New Mexico it fruits in the late summer and fall under pinyon pine.
EDIBILITY: Edible and collected by some people in spite of its sliminess (see comments
on edibility of H. sordidus).
COMMENTS: The pure white slimy cap and slimy stem plus the soft, waxy white gills
typify this cosmopolitan Hygrophorus, which is the type species of the genus. Sometimes
the layer of slime is so thick that it’s difficult to pick up the mushroom. At other times the
slime dries out, in which case there is likely to be debris glued irrevocably to the cap—a sure
sign that H. eburneus ranks with Limacella illinita and H. gliocyclus (among others) as
the “slipperiest and slimiest gilled fungus among us.” (L. illinita is also white, but has free
gills, while H. gliocyclus is squatter, slightly yellower, and is monogamous with pine).
H. piceae is a very similar species with a pure white viscid cap and non-viscid stalk. It is
common under conifers, especially spruce, in the Pacific Northwest and northern Califor¬
nia. For similar species with a buff- or ochre-tinged cap, see comments under H. gliocyclus.

Hygrophorus gliocyclus (Glutinous Waxy Cap)

CAP (2) 4-10 (15) cm broad, convex or obtuse becoming broadly umbonate, plane, or even
shallowly depressed; surface smooth, very slimy or viscid when moist, white to pale cream,
usually more yellowish or buff toward the center; margin at first inrolled. Flesh white,
fairly firm. GILLS adnate to decurrent, fairly well-spaced, thick, soft, somewhat waxy;
whitish to pale or dingy yellow, or sometimes tinged faintly pinkish. STALK 2-6 cm long,
(0.6) 1-2.5 cm thick, typically rather squat, equal to ventricose (swollen in the middle) or
tapered at base, smooth, very slimy when moist except at apex; dingy white to creamy,
solid. VEIL evanescent, leaving slime on stalk and sometimes an obscure ring (annulus).
SPORE PRINT white; spores 8-11 x 4.5-6 microns, elliptical, smooth. Gill tissue divergent.
HABITAT: Scattered to gregarious in needle duff under pines, widely distributed. Fairly
common in our area in fall, winter, and early spring, sometimes growing with H. hypo¬
thecs. I have found it along the coast with Monterey and bishop pines, and in great
numbers inland with Coulter, digger, and ponderosa pines.

120
Hygrophorus gliocyclus, mature specimens. This species is just as slimy as H. eburneus, but is
stockier and slightly yellower. It is common under pine.

EDIBILITY: Edible and “choice,” according to some, but like H. eburneus, disagreeable
to collect because of the copious slime (see comments on edibility of H. sordidus). One
book describes how to remove the slime from the mushrooms, but says nothing about
removing the slime from your hands.
COMMENTS: The creamy-white to yellowish cap, stocky stature, thick layer of slime on
the cap and stem (when moist), somewhat waxy gills, and association with pine are the
hallmarks of this humdrum Hygrophorus. It is thicker, squatter, and yellower than H.
eburneus, and the stalk is indisputably viscid, in contrast to H. sordidus. A closely-related,
equally slimy pine-lover, H. flavodiscus, is also widespread but has pinkish gills in youth;
H. glutinosus of eastern North America and the Pacific Northwest is the same color as
H. gliocyclus but favors hardwoods and shows reddish-brown spots on the stalk apex as it
dries; H. whileii of northern California has the color of H. gliocyclus but the stature of
H. eburneus; H. cossus is a slimy white species that develops ochraceous tones on the cap
as it ages and has a distinctive odor (like “goat moth larvae”), but is more common in
Europe than North America; H. chrysaspis also discolors in age but has no odor and gills
that dry dark brown. H. ponderatus (see H. sordidus) is large and white.

Hygrophorus subalpinus (Subalpine Waxy Cap)

CAP 4-15 (25) cm broad, broadly convex becoming plane or slightly depressed; surface
viscid when moist but soon dry, smooth, pure white or developing a slight yellowish tinge
in age; margin sometimes with veil remnants. Flesh thick, firmat first but soft in age, white;
odor and taste mild. GILLS white when young, often creamy or tinged dingy yellowish in
age; typically adnate to decurrent, narrow, close, soft and/ or waxy. STALK 3-10 cmlong,
1-5 (7) cm thick at apex, usually thick and stout, often with a rounded basal bulb when
young (but often more or less equal in age); firm, dry, solid, white. VEIL somewhat mem¬
branous, disappearing or forming a narrow, flaring or flange-like, median to inferior
ring on the stalk (just above the bulb). SPORE PRINT white; spores 8-10 * 4.5-6 microns,
elliptical, smooth. Gill tissue divergent.
HABITAT: Solitary to gregarious on ground under conifers; known only from the
mountains of western North America. It is a common “snowbank” species in the Sierra
Nevada, Cascades, and Rocky Mountains, but also appears in the summer and late fall.
I have yet to find it on the coast.
EDIBILITY: Deer consider it a delicacy; I don’t. Although tempting because of its
robustness and penchant for growing in the spring when there are few edible agarics out

121
122 HYGROPHORACEAE

and about, it decays quickly and does not have the greatest texture and flavor (see com¬
ments on edibility of H. sordidus).
COMMENTS: This hefty white Hygrophorus is easily told from other waxy caps (H.
sordidus, et al) by its thick dry stalk and whitish color, the presence of a veil that often
forms an annulus (ring), and association with mountain conifers (see photograph at top of
p. 109). The annulus, when present, can mimic the volva of an Amanita because it tends
to sit so low on the stalk. Amanitas, however, do not have decurrent gills and are rarely as
robust. The presence of an annulus can also lead to confusion with Armillariaponderosa
and A. olida (another springtime conifer-lover), but both of those mushrooms have strong
odors. Russula brevipes is also somewhat similar, but has brittle flesh and lacks a veil.

Hygrophorus sordidus (Sordid Waxy Cap)

CAP 5-20 cm broad or more, convex becoming plane or with uplifted margin; surface
smooth, viscid when moist but not often slimy and soon dry; white or sometimes tinged
yellowish-buff at center; margin at first inrolled. Flesh thick, firm, white. GILLS adnate
to decurrent, soft but only slightly waxy; white, sometimes dingy yellowish in age. STALK
6-10 cm long, 1.5-4 cm thick, equal or narrowed at base, solid, smooth, firm, white, not
viscid. VEIL absent. SPORE PRINT white; spores 6-8 x 4-5.5 microns, elliptical, smooth.
Gill tissue divergent.
HABITAT: Solitary to widely scattered or gregarious on ground in woods, associated
principally (if not exclusively) with oaks, widespread but primarily southern. In our area
it fruits with live oak in the fall and winter. It is quite abundant some years, but almost
completely absent others.
EDIBILITY: Edible but not choice. I ruined an otherwise superb curry in my sole attempt
to make it palatable. The sliced caps neither absorbed the surrounding spices nor
contributed any special flavor of their own. The turmeric did turn them yellow, however,
making them look for all the world like undercooked and overfed banana slugs—gummy,
amorphous masses of slime that coagulated around or completely engulfed the peas
and savory chunks of potato, rendering the entire dish inedible (though unforgettable).
Since its attractive appearance belies its “slugulose” qualities, I can only conclude that the
scientific soul who named it had a similar and equally sordid experience with it.
COMMENTS: Our heftiest Hygrophorus, its size alone distinguishes it from most
other white members of the genus. The stalk is not viscid as in H. gliocyclus and H. ebur-
neus, nor is there a veil as in H. subalpinus. It is likely to be mistaken for a Clitocybe or

Hygrophorus sordidus. A robust white waxy cap with a viscid cap, dry stem, and white gills that are
only slightly waxy. H. ponderatus (not illustrated) is an equally robust, viscid-stalked white species.
HYGROPHORUS 123

Russula because of its fleshiness, but the gills are soft, and at least to the experienced
Hygrophorus-hunter, waxy. H. penarius has a slightly darker cap but is otherwise similar.
H. perfumus of the Sierra Nevada has a fragrant odor. H. ponderatus is a similar but
slightly more flavorful, widespread species whose stalk is viscid or lubricous (at least over
the lower portion); in California it usually grows with conifers.

Hygrophorus russula (Russula-Like Waxy Cap)

CAP 5-13 cm broad, convex to plane or with uplifted margin in age; surface viscid when wet
but soon dry; coral-pink to vinaceous-red, usually streaked with purple-red or vinaceous
fibrils; smooth or minutely scaly, occasionally staining yellowish when rubbed or in age;
margin often paler or whitish and incurved when young. Flesh thick, white or tinged pink;
odor and taste typically mild. GILLS usually adnate but sometimes adnexed or slightly
decurrent, close to crowded (120-150 reach the stalk), soft, slightly waxy; white at first but
soon flushed pink and developing purple-red to vinaceous stains in age. STALK 3-10 cm
long, 1.5-3.5 cm thick, usually rather stout; solid, dry, smooth, equal or tapered below;
white, soon stained or streaked pink to reddish or vinaceous. VEIL absent. SPORE
PRINT white; spores 6-8 * 3-5 microns, elliptical, smooth. Gill tissue slightly divergent.
HABITAT: Scattered to gregarious or in rings in mixed woods and under hardwoods,
associated mainly with oaks; widely distributed and common in eastern North America,
but infrequent on the west coast. In our area I find it occasionally in the late fall and early
winter in tanoak-madrone woods at higher elevations in the coastal mountains.
EDIBILITY: Edible and choice, according to some, but my lingering distaste for fleshy
waxy caps (see comments on edibility of H. sordidus) has discouraged me from sampling it.
One authority calls it “the best of the family for the table”—a classic example of damning
with faint praise, if you ask me. On the other hand, local Hygrophorus-hound Luen Miller,
who suffers from an acute case of “Mcllvaine-mania,”* says of H. russula: “Not as good as
H. sordidus, but as an edible species it is not to be despised. It has a noble waxy texture and
makes a toothsome meal. Although it lacks the supreme succulence of H. sordidus, it
largely makes up for it in possessing a copious supply of wax, which coats the mouth and
throat for hours after eating it, the way good ice cream does. All authorities pronounce it
excellent. It is delightful scalloped or stewed, or better yet, made into catchup and poured
over food. The Tartars, I am told, know it as ‘poor man’s candle’ since the dried ’carps
offer a weak flickering flame when lit.”
*See footnote on p. 419!

Hygrophorus russula, mature specimens. The entire fruiting body develops dark red streaks and
stains; the gills are close together and only slightly waxy.
124 HYGROPHORACEAE

COMMENTS: A beautiful, robust Hygrophorus, best recognized by its coral-pink to

vinaceous (wine-colored) or vinaceous-red color, absence of a veil, and growth with
hardwoods. As the name implies, it has somewhat the stature of a Russula, but the stalk
does not snap open cleanly like chalk. It was originally placed in Tricholoma, but the gills
are usually adnate rather than notched, and quite soft and waxy. For similar waxy caps
associated with conifers, see H. purpurascens.

Hygrophorus purpurascens (Purple-Red Waxy Cap)

CAP (3) 6-12 (20) cm broad, convex becoming broadly convex or plane; surface slightly
viscid when wet, otherwise dry; whitish to coral-pink with darker (vinaceous-red to purple-
brown) splashes, streaks, and/or fibrils; margin incurved at first and usually paler. Flesh
thick, firm, white; odor mild, taste mild or bitter. GILLS adnate to decurrent, white at first
but soon flushed pink, then spotted or stained purple-red to vinaceous; fairly close, soft,
slightly waxy. ST ALK (3)5-12(15) cm long, 1 -2.5 cm thick, equal or tapered below, solid,
firm, not viscid; colored or stained more or less like the cap. VEIL fibrillose, white, forming
a slight superior or apical ring or hairy zone on stalk, or disappearing entirely. SPORE
PRINT white; spores 5.5-8 x 3-4.5 microns, elliptical, smooth. Gill tissue divergent.
HABITAT: Solitary to scattered, gregarious, orin troops underconifers, especially spruce
and pine; widely distributed in northern and western North America. It does not occur in
our area, but is fairly common in the Sierra Nevada and Cascades from spring through
fall, and fruits prolifically in late summer in the spruce forests of the Rocky Mountains.
EDIBILITY: Edible, but some variants are unpalatable (bitter) even when cooked—as
I can personally attest.
COMMENTS: This waxy cap and its close relatives (see below) are very common at times
in our western mountains. They resemble H. russula in their reddish-pink to vinaceous-
red color, but favor conifers rather than hardwoods. H. purpurascens is distinct by virtue
of its fibrillose veil, which is often evident only in young, unexpanded specimens. Its
close relatives include: H. erubescens, quite similar but lacking a veil and with a frequent
tendency to slowly stain yellow when bruised or left overnight in the refrigerator. It is
normally a fairly robust mushroom, but a slender form occurs rarely in our area with pine.
H. capreolarius is a similar but smaller, more slender (stalk about 1 cm thick), spruce-
loving species with well-spaced gills and fruiting body evenly colored dark vinaceous-red
in age. I have collected it several times under Sitka spruce in northern coastal California,
but its distribution parallels that of H. purpurascens. H. amarus is also similar, but
is very bitter-tasting and tends to develop pronounced pale yellow to bright yellow tones on
the cap (or whole fruiting body) in age. It may or may not have a very slight fibrillose veil
(check young specimens!); it’s fairly common under spruce and fir in the Rocky Mountains.

Hygrophoruspudorinus (Spruce Waxy Cap) Color Plate 16

CAP 5-15 (20) cm broad, obtuse or convex with an inrolled margin, becoming broadly
convex or plane in age; surface smooth, viscid when moist, pale tan to pinkish or flesh-
color, or pinkish-orange toward the center and pinker or paler at the margin. Flesh thick,
firm, white or tinged cap color; odor mild to faintly fragrant. GILLS adnate to decurrent,
fairly close, soft, waxy, sometimes whitish but more often pinkish to pale flesh-color; not
developing reddish stains. STALK 4-20 cm long, 1-3 cm thick, equal or narrowed below,
solid, dry to tacky but not truly viscid, whitish to buff or pinkish or colored like cap; lower
portion fibrillose, upper portion conspicuously punctate (with tiny whitish scurfy scales or
tufts that darken to reddish-brown when dried or in age, and turn yellow-orange in KOH).
VEIL absent. SPORE PRINT white; spores 6.5-9.5 x 4-5.5 microns, elliptical, smooth.
Gill tissue divergent.
HYGROPHORUS 125

HABITAT: Scattered or in groups on ground under conifers, particularly spruce; widely

distributed but erratic in its fruiting habits. It does not occur in our area (probably due to
the absence of spruce), but I have seen large fruitings in December with Sitka spruce in
northern California, and in August with Engelmann spruce in the Southwest.
EDIBILITY: Edible but mediocre. Some variants are said to have a “turpentine-like taste,”
but I cannot vouch for the accuracy of this comparison, since I have never tasted tur¬
pentine. (Have you?)
COMMENTS: This attractive, rather robust waxy cap is difficult to characterize but
relatively easy to recognize (see color plate). The viscid pinkish to pinkish-salmon or pale
tan cap, punctate stalk apex, and waxy gills are good fieldmarks. The viscid cap distin¬
guishes it from the similarly colored Camarophylluspratensis. Several varieties and color
forms have been described, including var. fragrans, large and tall, with a yellow-orange
stalk base and tendency to stain yellow or orange when bruised. Another beautiful waxy
cap, H. saxatilis, occurs with conifers in the Pacific Northwest, especially on rocky hill¬
sides. It has a viscid, whitish to pale buff cap (sometimes with darker watery spots) and
lovely pale apricot to pinkish, decurrent gills. Still another distinctive species, H. goetzii,
is rather small and slender, with a rosy or pinkish cap and creamy gills often tinged with the
cap color. It is sometimes common in the late spring and summer in the Sierra Nevada and
Cascades, often near or even in melting snow.

Hygrophorus roseibrunneus (Rosy-Brown Waxy Cap)

CAP 2-10 cm broad, convex becoming broadly umbonate or plane, or with an uplifted
margin in age; surface smooth, viscid when moist, reddish-brown to pinkish-cinnamon to
rosy-brown or brown at least at the center and often overall; margin often paler or whitish.
Flesh white, soft, odor mild. GILLS usually adnate, but sometimes decurrent or adnexed;
close, soft, somewhat waxy, white. STALK 3-12 cm long, 0.5-1 (1.5) cm thick, equal or
tapered below, often curved near base; smooth, white, not viscid; apex often prominently
punctate. VEIL absent. SPORE PRINT white; spores 6-9 * 3.5-5 microns, elliptical,
smooth. Gill tissue divergent.
HABITAT: Solitary to widely scattered or in groups, associated with oak in our area
and sometimes common in the late fall and winter (along with H. brunneus). It was first
collected in Palo Alto, California, but also occurs in eastern North America.
EDIBILITY: Edible. I have tried it.
COMMENTS: The reddish- to rosy-brown color of the cap (or center of the cap), non-
viscid stalk, waxy white gills, and mild odor are the fallible fieldmarks of this species. The
punctate stem apex is characteristic of many waxy caps, but in this one it is especially at¬
tractive. Leucopaxillus amarus is similarly colored but has a dry cap, non-waxy gills, bitter
taste, and white mycelium at the base of the stem. Just as common in our area isH. brun¬
neus, very similar but for its yellow-brown to tawny-brown cap (or cap center). A species
endemic to California, H. albicastaneus, is also similar, but ranges from pure white (when
young) to yellow-brown to peach- or ochre-tinged (in age). It has a non-viscid stalk and
usually grows with oak, but stains rusty or fulvous in potassium hydroxide.H. subpungens
is a small, slender-stemmed species with a pinkish-brown to yellow-brown to salmon-
centered cap and very faint fruity odor; it occurs locally under alder, oak, and Douglas-fir.
H. tennesseensis, widespread, is also similar but favors conifers and has a bitter taste and
potato-like odor. There are also several similar species with a viscid stalk, including: H.
laurae, with a whitish cap margin, favoring hardwoods; H. discoideus, slightly darker,
favoring conifers; and H. vernalis, a western springtime “snowbank” species with a pink¬
ish-buff to pale vinaceous cap and sordid yellowish patches on the lower half of the stalk.
126 HYGROPHORACEAE

Hygrophorus bakerensis (Brown Almond Waxy Cap)

CAP 4-15 cm broad, obtuse to convex becoming broadly convex or plane; surface smooth,
slimy or viscid when moist, cinnamon-brown to yellow-brown or tawny, the margin
usually paler or whitish. Flesh thick, white; odor sweet but sometimes faint, like almond
extract or crushed peach pits. GILLS white to creamy or pinkish-buff, usually decurrent
but varying to adnate; soft, somewhat waxy. STALK 4-15 cm long, 0.8-2.5 cm thick, equal
or tapered downward, smooth, solid, dry; white to pinkish-buff. VEIL absent. SPORE
PRINT white; spores 7.5-10 * 4.5-6 microns, elliptical, smooth. Gill tissue divergent.
HABITAT: Widely scattered to gregarious under conifers, known only from the Pacific
Northwest and northern California; very common at times in the fall and early winter.
EDIBILITY: Edible but bland (it doesn’t taste like it smells).
COMMENTS: This species is one of the commonest and most characteristic waxy caps of
the Pacific Northwest. The reddish-brown to yellow-brown cap, waxy decurrent gills, non-
viscid stalk, and almondy odor set it apart. The gills and stalk are sometimes beaded with
droplets in moist weather, but there is no latex as in Lactarius. The gills are usually decur¬
rent, in contrast to Collybia oregonensis, a similarly colored, strongly fragrant mushroom
with adnexed or notched, non-waxy gills. H. agathosmus has the same odor, but its cap is
gray, not brown. Other fragrant waxy caps from the Pacific Northwest and California
include: H. monticola and H. vinicolor, similar but with larger spores (the latter with an
unpleasant taste); H. variicolor, also similar but with lower half of stalk distinctly viscid
or glutinous; and H. pacificus, a strongly aromatic (but not almondy) species with a russet
to tawny to pinkish-buff, often lobed or wavy cap, pale yellowish gills, and larger spores.

Hygrophorus speciosus (Larch Waxy Cap) Color Plate 15

CAP 2-5 (8) cm broad, convex to broadly umbonate or expanding to plane or even centrally
depressed; surface slimy or viscid when moist, smooth, bright orange-red to orange, often
fading to orange-yellow; margin often paler. Flesh white or tinted yellow, soft. GILLS
adnate to decurrent, white to pale yellow, the edges usually yellow; well-spaced, thick, soft,
waxy. STALK 3-10 cm long, 0.4-1 (1.5) cm thick, equal or thicker below, white or with
yellow to orange stains over lower portion; viscid to slimy when moist, at least below. VEIL
single (var. speciosus) or double-layered (var. kauffmanii), outer layer evanescent, leaving
slime on stalk; inner layer when present fibrillose, sometimes forming a slight ring on stalk.
SPORE PRINT white; spores 8-10 x 4.5-6 microns, elliptical, smooth. Gill tissue divergent.
HABIT AT: Scattered to densely gregarious in woods and bogs under conifers, especially
larch but also pine; common in the late summer and fall throughout the range of larch, but
also frequent with ponderosa pine in the Southwest.
EDIBILITY: Edible; I haven’t tried it.
COMMENTS: This is one of the few brightly colored waxy caps belonging to the genus
Hygrophorus in its strictest sense (i.e., waxy caps with divergent gill tissue). It is easily told
from the numerous red to orange Hygrocybes by its habitat, viscid stalk, and white to pale
yellow decurrent gills. The cap is never olive-brown as in H. hypothejus. H. pyrophilus
is similar; it has a red cap, less viscid stalk, and was found in burnt ground near Mt. Shasta.

Hygrophorus hypothejus (Olive-Brown Waxy Cap)

CAP 2-8 cm broad, convex to broadly umbonate, plane, or depressed; surface smooth,
viscid or slimy when wet, color variable: typically dark brown to olive-brown at the center
and greenish-yellow to yellow at the margin when young (but sometimes entirely olive-
brown), often developing yellow-orange to reddish-orange tones in age, especially near
the margin. Flesh thin, yellowish to whitish; odor mild. GILLS decurrent or occasionally
Hygrophorus hypothejus commonly grows with pine, often in the company of H. gliocyclus. Gills
are pale yellow to orange, decurrent, and fairly waxy, and the cap is slimy when wet.

adnate, well-spaced, thick, soft, waxy, at first pallid but soon becoming pale yellow, and in
age sometimes brightly colored (like margin of cap). STALK (3) 5-15 cm long, 0.5-1.5 (2)
cm thick, equal or tapered downward; yellow at apex, otherwise pallid or variously colored
(like cap) and viscid or slimy when moist. VEIL evanescent, leaving slime on stalk and
sometimes an obscure fibrillose ring. SPORE PRINT white; spores 7-9 * 4-5 microns,
elliptical, smooth. Gill tissue divergent.
HABITAT: Scattered to gregarious or in troops under conifers, particularly pine—very
widely distributed and often abundant in cool weather. It is common from late fall through
early spring in our coastal pine forests.
EDIBILITY: Bountiful, but bland. See H. sordidus for details.
COMMENTS: This waxy cap is best recognized by its olive-brown to olive-yellow cap
when young which is sticky or slimy and convex to depressed in age, the decurrent gills, and
association with pine. The colors are extremely variable, especially in age, and young
specimens bear little resemblance to old ones until you find stages in between. Both slim and
relatively robust specimens can be found. Other species with an olive-brown to grayish cap
and viscid stalk include: H. fuligineus, very similar but with a darker (olive to blackish)
cap, common in cool weather under conifers (mainly in eastern North America); H.fusco-
albus, with larger spores (9-13 microns long) and H. limacinus, with even larger (10-17
microns) spores; and two species which lack an inner (fibrillose) veil: H. occidentalis,
small-spored, cap brown to grayish with a pallid margin, often under oak; and H. mega-
sporus, with spores 12-18 microns long. All of these species favor conifers and have white
to grayish gills; none develop the bright colors typical of mature H. hypothejus.

Hygrophorus olivaceoalbus (Sheathed Waxy Cap)

CAP 3-12 cm broad, convex or broadly umbonate to more or less plane; surface slimy or
viscid when moist, dark brown to nearly black at the center, usually paler (grayish to
olive-brown) toward the margin, and usually with a streaked appearance from darker
fibrils. Flesh thick, white, soft, odor mild. GILLS adnate to decurrent, thick, soft, waxy,
pure white or pallid, sometimes becoming grayish. STALK (3) 5-12(15) cm long, 1-3 cm
thick, equal or thicker below; smooth and white above the ring, viscid to slimy (when moist)
below and sheathed with blackish to grayish-brown, olive-brown, or gray fibrils which
break up into patches or scaly rings. VEIL double-layered, the outer layer evanescent and
depositing slime on stalk, the inner layer fibrillose and sometimes forming an obscure ring
at top of fibrillose sheath. SPORE PRINT white; spores 9-12 * 5-6 microns, elliptical,
smooth. Gill tissue divergent.
128 HYGROPHORACEAE

HABITAT: Solitary, scattered, or in groups on ground under conifers (especially spruce)

in western and northern North America, fruiting from late summer through early winter.
I have seen it in northern California under Sitka spruce and in the southern Rocky
Mountains under Engelmann spruce and blue spruce.
EDIBILITY: Said to be edible, but bland and slimy.
COMMENTS: This handsome Hygrophorus can be distinguished from similarly colored
waxy caps by the gray to dark brown fibrillose sheath on the stem. The contrasting colors
are very striking. The coastal variety (also called H. persoonii) is usually quite dark and
slimy; the Rocky Mountain version is often paler (grayer) and only slightly viscid if at all.
Other species: H. inocybiformis has a grayish-brown sheath and veil, but its stalk is not
viscid, and its dark gray cap is only 3-7 cm broad; it occurs in the Pacific Northwest under
conifers. H. tephroleucus, H. pustulatus, and H. morrisiiare even smaller and have small,
pointlike grayish scales on the stalk. See also the species listed under H. hypothejus.

Hygrophorus agathosmus (Gray Almond Waxy Cap)

CAP 3-10 cm broad, convex to plane or with margin uplifted; surface smooth, viscid when
moist, dull gray to ashy-gray, brownish-gray, or at times grayish-olive; margin incurved at
first. Flesh soft, whitish; odor sweet, like almond extract (but sometimes faint). GILLS
adnate to slightly decurrent, close or well-spaced, soft, waxy, white or sometimes grayish in
age. STALK 4-10 (16) cm long, 0.5-1.5 (2) cm thick, equal or narrowed below, smooth, not
viscid; white or tinged gray. VEIL absent. SPORE PRINT white; spores 7-10.5 * 4.5-5.5
microns, elliptical, smooth. Gill tissue divergent.
HABIT AT: Scattered to gregarious under conifers, widely distributed and fairly common
in cool weather, but rather infrequent in our area. It is partial to spruce, but is associated
locally with Douglas-fir (probably because there isn’t any spruce).
EDIBILITY: Edible, but bland. It is unfortunate that it doesn’t taste like it smells.
COMMENTS: The grayish cap, waxy gills, dry stalk, almondy fragrance, and association
with conifers are the telltale traits of this fine fungus. The odor is sometimes faint, but if
several are placed together in a closed container it usually becomes evident. H. calophyllus
and H. camarophyllus are somewhat similar but do not have an almondy odor. Other
species: H. odoratus is a small, slender version of H. agathosmus with an almondy odor
and larger spores(l 1-14 microns long); it occurs in the Pacific Northwest under conifers.
H. occidentalis has a viscid stalk, weaker odor, and grows under oak or conifers.

Hygrophorus agathosmus, mature specimens. Note the Douglas-fir needles stuck to the cap, indi¬
cating that it was viscid. Cap is grayish, gills white and waxy, odor almondy. It also grows with spruce.
HYGROPHORUS 129

Hygrophorus calophyllus (Gray-Brown Waxy Cap)

CAP 4-11 cm broad, convex to broadly umbonate or plane; surface smooth, viscid or
slimy when moist, evenly colored deep olive-brown to dark gray-brown to umber, the
margin sometimes slightly paler. Flesh soft, white, thick; odor mild or faintly fragrant.
GILLS white or flushed a delicate pink, well-spaced, thick, soft, waxy, usually decurrent.
STALK 5-12 cm long, 1-1.5 cm thick, equal or narrowed below, smooth, not viscid; apex
white, otherwise colored like the cap but usually slightly paler. VEIL absent. SPORE
PRINT white; spores 5.5-8 x 4-5 microns, elliptical, smooth. Gill tissue divergent.
HABITAT: Solitary or in small groups on ground under conifers, western North America,
not common. I have found it only once in our area, with Douglas-fir, in December.
EDIBILITY: Edible.
COMMENTS: Whether white or pink, the gills contrast sharply with the dark cap and
stem, making this a most attractive mushroom. Often the gills will develop their pinkish
tint only as they mature, leading one to wrongly assume that the developing spores are
pink! The stalk is not viscid as in H. hypothejus and H. olivaceoalbus, and the soft, clean,
waxy gills distinguish it from Clitocybe. Very similar but more common is H. camaro-
phyllus, with a dry to only slightly viscid (never slimy), streaked, grayish-brown cap and
white to grayish-tinted (never pink) gills. It also fruits under conifers, sometimes near
melting snow, and is easily mistaken for a Clitocybe. H. marzuolus grows almost
exclusively in the spring, often near snow. It has well-spaced gills and is entirely pallid
when young, but soon becomes grayish overall (including the gills).

TRICHOLOMATACEAE
THIS is by far the largest and most diverse family of pale-spored agarics. The spore print
is usually white, but ranges to buff, yellowish, pale lilac, or pinkish. A stem is normally
present, but in several of the shelflike, wood-inhabiting species it is rudimentary or even
absent. The gills are typically attached to the stem, but in some of the smaller forms they
are free. Though most of the species are terrestrial, many grow on wood—whereas other
pale-spored families are almost exclusively terrestrial. They are largely woodland fungi,
but a few, such as Marasmius oreades, grow in grass.
The best way to recognize the family is to eliminate the other pale-spored families. The
gills are not normally soft, thick, and waxy as in the Hygrophoraceae, nor shallow, blunt,
and foldlike as in the Cantharellaceae; the fleshy forms do not have noticeably dry, brittle
flesh as in the Russulaceae (and their tissue lacks sphaerocysts—a microscopic feature),
nor do they have a volva as in the Amanitaceae, and the few species with a veil do not have
the free gills typical of the Lepiotaceae. (There are some exceptions to the above, but they
are discussed under individual species or genera.)
The size and complexity of the Tricholomataceae are such that no generalizations can be
made regarding edibility—other than that some are poisonous and some are not. Many
have not been adequately tested and others are too small to be of value. The safest approach
is to learn the distinguishing characteristics of each edible species—and there are some
plentiful collectable delectables that are well worth getting to know! The most notable
are the oyster mushroom (Pleurotus ostreatus), man-on-horseback (Tricholoma flavo-
virens), honey mushroom (Armiliariella mellea), matsutake (Armillaria ponderosa),
fairy ring mushroom (Marasmius oreades), and blewit (Clitocybe nuda). Several lesser-
known species (e.g., Lentinusponderosus) are also excellent.
130 TRICHOLOMATACEAE

The Tricholomataceae embraces more genera than any other family of gilled
mushrooms, and its taxonomy is still in a state of flux. Many of the genera are defined by
esoteric chemical and anatomical (microscopic) characteristics, which presents obvious
problems when attempting to construct (or use) a key based solely on field characters. It
helps to break down the genera into three large groups: wood-inhabiting, shelflike forms
(typified by Pleurotus)\ fleshy-stalked, mostly terrestrial forms (typified by Tricholoma
and Clitocybe), and thin, fragile- or cartilaginous-stalked, terrestrial or wood-inhabiting
types (typified by Mycena, Collybia, and Marasmius).
In the following key, only the more distinctive genera are included; the more difficult or
obscure ones are then keyed out under the distinctives ones, sometimes on a species-
by-species basis.

Key to the Tricholomataceae

1. Growing on other mushrooms; gills thick and widely spaced or poorly formed to practically
absent .Asterophora, p. 200
1. Not growing on other mushrooms, or if so then gills well-developed, thin, close. 2
2. Fruiting body pinkish to salmon, orange, or yellow-orange; cap surface conspicuously reticulate
(netted or veined and pitted); cap 2-5 cm broad; stalk central or off-center, tough; spore print
pinkish; found on dead eastern hardwoods (e.g., maple); infrequent . . Rhodotuspalmatus
2. Not as above . 3
3. Stalk absent, or if present then typically off-center to lateral; usually growing on wood (or
woody material such as coffee bean waste or wood chip mulch) or on moss .4
3. Stalk present, well-developed, more or less central; growing on ground or wood .6
4. Fruiting body tough and leathery or corky; gill edges longitudinally split and cap densely hairy
or cap concentrically zoned or grooved (and often velvety) and gills often maze like (forming
elongated pockets) or wavy; found on hardwoods .(see Polyporaceae & Allies, p. 549)
4. Fruiting body fleshy, or if tough then not as above . 5
5. Edges of gills conspicuously serrated (toothed) or eroded, even when fresh.
.Lentinus & Lentinellus, p. 141
5. Not as above; gill edges usually entire (but sometimes wavy). Pleurotus & Allies, p. 132
6. Stalk arising from an underground “tuber” (the “tuber” cylindrical or bulbous, often hollow);
cap usually scaly, fibrillose, or granulose; not common .Squamanita, p. 197
6. Not as above (but stalk may have a tapered underground “tap root”) . 7
7. Cap granulose (covered with a layer of mealy or powdery granules that are sometimes washed
off by rain); veil present (check young specimens!), sometimes forming an annulus (ring) on
stalk; stalk granulose below the veil .Cystoderma, p. 198
7. Not as above; veil absent, or if present then cap and stalk not granulose . 8
8. Veil present, usually forming a distinct annulus (ring) on stalk . . . Armillaria & Allies, p. 189
8. Veil absent or rudimentary and evanescent, not forming an annulus . 9
9. Gills and stalk bruising dark gray to black (sometimes slowly) or developing such stains in age;
basidia with siderophilous granules; not common in our area . Lyophyllum & Allies, p. 173
9. Not as above . 10
10. Stalk fleshy, usually at least 5 mm thick . 11
10. Stalk usually thin and hollow or stuffed and either fragile or cartilaginous (tough), typically
5 mm thick or less (occasionally thicker but then with a tough cartilaginous outer rind) . 23
11. Fruiting body partially or completely purple, violet, or lilac when fresh (at least the gills); odor
not radishlike; usually on ground or compost . 12
11. Not as above . 13
12. Gills thickish and fairly well-spaced; stalk fibrous and/or fibrillose; spore print white or lilac-
tinged .Laccaria, p. 171
12. Gills close, not thick; spore print dull or dingy pinkish.Clitocybe & Allies, p. 148
13. Copious white mycelial mat usually present at base of stalk or in substrate; stalk and gills white or
yellowish but not gray; cap and stalk dry, dull, unpolished, often tough; stalk not hollow; spore
print white; spores with amyloid warts; found in woods or under trees Leucopaxillus, p. 166
13. Not with above features; spore print variously colored (white, buff, pinkish, etc.) . 14
TRICHOLOMATACEAE 131

14. Spore print white, yellowish, or buff (or in one case brownish) . 15
14. Spore print pinkish to pinkish-buff. 27
15. Typically growing in dense clusters in disturbed soil (along roads, paths, etc., but sometimes also
in woods); gills whitish to gray; stalk at least 1 cm thick; caps typically at least 3 cm broad; spore
print white (not buff); basidia with siderophilous granules . . . Lyophyllum & Allies, p. 173
15. Not with above features . 16
16. Gills pinkish, flesh-colored, cinnamon, or somewhat vinaceous, thickish and fairly well-spaced;
cap up to 6 cm broad; stalk rather tough and fibrous, not white; spores spiny Laccaria, p. 171
16. Not with above features . 17
17. Gills typically adnate to decurrent. 18
17. Gills typically notched, adnexed, or even free (occasionally adnate but not decurrent) ... 21
18. Gills and flesh olive-yellow to yellow to orange; cap not viscid; gills not repeatedly forked;
growing on or near wood (but wood often buried or not visible) or from roots . 19
18. Not with above features ... 20
19. Fruiting body orange to yellow-orange or with olive tones; cap and stalk smooth, without
scales; associated with hardwoods. Omphalotus, p. 146
19. Fruiting body pale yellow to yellow, often with differently colored scales or fibrils on cap or
stalk; associated mainly with conifers .Tricholomopsis, p. 144
20. Cap viscid or slimy when moist and/or gills thick, widely spaced, and clean or waxy-looking
. (see Hygrophoraceae, p. 103)
20. Not as above . .Clitocybe & Allies, p. 148
21. Growing on or near wood (sometimes very rotten or buried); flesh and gills often (but not
always!) yellow to pale yellow .Tricholomopsis, p. 144
21. Not as above .22
22. Cap smooth, without fibrils or scales, usually white to gray to grayish-brown or dark brown
when fresh and moist; gills crowded; found in many habitats but especially in grassy or land¬
scaped areas or in mountains soon after snow melts; spores amyloid . . Melanoleuca, p. 169
22. Not as above; cap variously colored (yellow, greenish, brown, grayish, white, reddish-brown,
etc.); found in woods or with trees; spores not amyloid .Tricholoma, p. 176
23. Spore print pinkish to ochre-brown; cap bell-shaped to conical when young, reddish-brown to
dark brown to blackish (the margin often paler); stalk similarly colored, minutely velvety;
odor usually strong and fishy or reminiscent of cucumber; giant cystidia present on gills; not
common in our area (but widespread) . Macrocystidia cucumis
23. Not as above . 24
24. Cap conical or bell-shaped when young (but may expand in age), often translucent-striate when
moist, margin not usually incurved when young; stalk not polished or tough Mycena, p. 224
24. Not as above .25
25. Gills purple to pinkish, flesh-colored, or dingy cinnamon, rather thick and well-spaced; cap
convex to plane or uplifted, not conical or bell-shaped; stalk tough, fibrous, not white; growing
on ground; spores usually spiny.Laccaria, p. 171
25. Not as above . 26
26. Cap small (up to 2.5 cm broad), cap and stalk golden-yellow to pale yellow and covered with
scurfy or mealy particles; gills creamy to yellow, usually adnate to decurrent; growing on logs
and sticks of hardwoods in eastern North America and on aspen in the Southwest and Rocky
Mountains .Cyptotrama chrysopeplum (-Xerulina chrysopepla)
26. Not as above . 27
27. Gills typically adnate to decurrent. 28
27. Gills notched to adnexed or free, or sometimes adnate Marasmius, Collybia, & Allies, p. 201
28. Gills and/or cap pale yellowish to yellow, orange, pinkish, or greenish when fresh (may fade!);
fruiting body small or minute (cap often less than 2.5 cm broad); stalk usually 1-3 mm thick;
cap usually depressed centrally in age; often found on logs or with grass, moss, or lichen
.Omphalina& Xeromphalina, p. 221
28. Not as above . 29
29. Stalk thin, tough, and pliant or if thick then with a tough outer cartilaginous rind; gills usually
adnate, if decurrent then usually widely spaced .Marasmius, Collybia, & Allies, p. 201
29. Not as above; gills often decurrent, usually close or crowded .Clitocybe & Allies, p. 148
132 TRICHOLOMATACEAE

PLEUROTUS & Allies

Small to large wood-inhabiting mushrooms usually growing shelflike. CAP smooth or hairy, dry
or viscid. Flesh soft, rubbery, pliant, or tough. GILLS adnate to decurrent, edges typically not
serrated. STALK absent or if present usually lateral or off-center, occasionally central. VEIL
absent (except in Pleurotus dryinus). VOLVA absent. SPORE PRINT pale (white to yellowish,
pale lilac, or pinkish). Spores smooth, not amyloid except for Panellus.

THIS is an artificial grouping of pale-spored, wood-inhabiting agarics with a consistently

off-center to lateral or absent stalk. The most common genus, Pleurotus, can be recognized
by its rather soft, fleshy fruiting body. Hohenbuehelia includes a number of species once
placed in Pleurotus but now segregated because of their semi-gelatinous to rubbery-
pliant flesh and large, thick-walled sterile cells (cystidia) on the gills. Panus and Panellus
incorporate those forms with a tough, often hairy fruiting body (the latter with amyloid
spores), while Phyllotopsis includes species with pinkish, sausage-shaped spores.
All of the above genera are small but widely distributed. The oyster mushroom
(Pleurotus ostreatus) and its relatives are among our best edible mushrooms, but most of
the other species are either too small, too tough, or too rare to be of value.
Key to Pleurotus & Allies
l. Veil present when young but often disappearing in age; flesh very thick; cap medium-sized to
large, typically with grayish fibrils, hairs; or scales(but sometimes whitish or ochre); gill edges
normally entire (not serrated); growing on hardwoods (often living) Pleurotus dryinus, p. 136
1. Not with above features . 2
2. Veil present, at least when young, sometimes forming an annulus (ring) on stalk . 3
2. Veil absent in all stages .4
3. Fruiting body small, tough, brownish; cap less than 2.5 cm broad; veil membranous; spores
amyloid; growing on dead hardwoods .Tectellapatellaris(-Panus operculatus)
3. Not as above; larger .(see Lentinus& Lentinellus, p. 141)
4. Cap minute (2-8 mm broad), bluish-gray to bluish-black or grayish-black; fruiting body shaped
like an inverted cup; gills widely spaced; stalk absent; growing on hardwoods, shrubs, vines,
etc., but not on mosses . Resupinatus applicatus
4. Not as above . 5
5. Cap small (less than 2.5 cm) and hairy, white to brownish; gills forking or veinlike and very wavy
(crisped); on hardwoods in eastern North America Trogia crispa(see Schizophyllum, p. 590)
5. Not as above . 6
6. Cap small (less than 2.5 cm), often rubbery or gelatinous, white to grayish; gills often veinlike
or even absent; usually (but not always) growing on mosses; mostly northern Leptoglossum
6. Not as above . 7
7. Cap small or minute (up to 2.5 cm broad), white to creamy or tinged purplish or pinkish . . 8
7. Not as above (larger and/or differently colored) . 11
8. Gills very widely spaced and often with veins between them; stalk usually present and darkening
in age from base upward; common along west coast . . . (stzMarasmiellus candidus, p. 206)
8. Not as above .. 9
9. Fruiting body white or whitish; gills fairly well-spaced to widely-spaced; fruiting body not
tough; spores not amyloid .Pleurotus (-Cheimonophyllum) candidissimus
9. Not as above; spores amyloid . 10
10. Gills pinkish-gray; cap usually with purplish or vinaceous tints; growing in groups on dead
hardwoods (especially birch) .Panellus ringens
10. Gills pallid to pale pinkish-gray; cap more or less same color and often with a gelatinous layer
when young; found on dead conifers (especially larch) .Panellus mitis
11. Gills usually veined (especially near stalk or base of cap) or repeatedly forked; gills orange to
yellowish or olive-yellow, or if not then lower stalk brown and velvety (see Paxillaceae, p. 476)
11. Not with above features . 12
12. Gills orange to yellow-orange, olive-yellow, or yellowish . 13
12. Gills some other color (white, gray, brownish, violet, etc., but may age or discolor yellowish) 15
Pleurotus ostreatus, gill detail. The white gills, white or lilac-tinged spores, off-center stalk, and fleshy
texture typify Pleurotus. (Dan Harper)

13. Cap densely hairy or fuzzy; stalk absent or rudimentary .Phyllotopsis nidulans, p. 140
13. Not as above; cap not hairy, or if hairy then stalk present ... 14
14. Cap hairy and taste bitter-acrid or cap viscid when moist or old and flesh white . 26
14. Cap not hairy; flesh not white .(see Omphalotus, p. 146)
15. Cap fairly large to very large (10-50 cm broad when mature); either growing on hardwoods and
cap coarsely hairy or found on or near conifers and stalk usually well-developed . 16
15. Cap smaller, or if large then not as above . 17
16. Cap coarsely hairy; growing on hardwoods .Panus strigosus, p. 140
16. Stalk well-developed; growing on or near conifers .(see Lentinusponderosus, p. 143)
17. Cap with dense brown hairs and fibrils, funnel-shaped or with a deeply depressed center; gills
decurrent, very crowded, pallid; stalk well-developed; common in the tropics and also along
the Gulf of Mexico .Panus (-Lentinellus) crinitis
17. Not as above . 18
18. Cap only 2-4 cm broad, viscid when moist, pallid to pale orange becoming pinkish to caramel-
brown; gills close to fairly well-spaced (not crowded); spores amyloid . Panellus longinquus
18. Not with above features; common. 19
19. Gills pallid (white to creamy, yellowish, or gray) . 20
19. Gills darker (tan to brown, reddish-brown, violet-tinted, etc.) . 26
20. Gills narrow (shallow) and crowded; fruiting body small to medium-sized . 21
20. Gills broad or fairly broad (deep), well-spaced to close, but not crowded; fruiting body medium¬
sized to large . 23
21. Cap pure white when fresh, but may become creamy in age.Pleurotusporrigens, p. 135
21. Not as above . 22
22. Cap dark grayish-brown to bluish-black; usually found in the wild .
.Hohenbuehelia atrocaerulea (see H. petaloides group, p. 136)
22. Cap some shade of brown or paler; found in wild or often in gardens, flower pots, etc.
.Hohenbuehelia petaloides group & others, p. 136
23. Stalk absent or rudimentary, not well-developed; cap with ridges and/or spines; taste un¬
pleasant .Hohenbuehelia mastrucatus (see H. petaloides group, p. 136)
23. Not as above . 24
24. Cap white to tan or pinkish-tinged, often breaking up into scales in age; stalk present; found
on living hardwoods (elm, etc.), often high up in the tree; widespread but not yet reported
from California; edible but rather tough .... Hypsizygus tessulatus (-Pleurotus ulmarius)
24. Not as above . 25
25. Stalk well-developed and often long (4-22 cm); gills usually adnexed or notched; cap creamy or
tinged pinkish, often with watery spots; usually on living hardwoods . Pleurotus elongatipes
25. Stalk absent, or if well-developed then gills typically decurrent; cap white, gray, brown, greenish,
etc.; very common, especially on hardwoods ..Pleurotus ostreatus & others, p. 134
134 TRICHOLOMATACEAE

26. Cap small (3 cm broad or less); gills ochre-buff to brownish to pale cinnamon or tawny-olive in
old age; taste usually acrid or bitter.Panellus stipticus, p. 138
26. Not as above; usually larger . 27
27. Cap densely hairy .Panus rudis, p. 139
27. Not as above; cap more or less smooth (or breaking up into scales) . 28
28. Cap viscid when moist or in age, yellow-green to olive-green, olive-brown, or with violet tones;
stalk absent or lateral; growing shelflike .Panellus serotinus, p. 137
28. Cap not viscid, violet to reddish-brown, tan, etc., but never greenish or yellow; stalk often well-
developed, lateral to off-center or central .Panus conchatus, p. 138

Pleurotus ostreatus (Oyster Mushroom) Color Plates 27,28

CAP 4-15 cm broad or more, oyster- or fan-shaped, convex becoming plane or sometimes
funnel-shaped; surface smooth, slightly lubricous when moist but not viscid; color
variable: white to gray, grayish-brown, tan, or dark brown (sometimes yellowish in old
age); margin inrolled when young, often wavy or lobed. Flesh thick, white, firm but soft,
tougher near the stalk; odor and taste mild. GILLS fairly close, broad, decurrent (if stalk
is present), white or tinged gray but often discoloring yellowish in old age. STALK absent
or if present usually short, stout, and off-center or lateral (but sometimes central); 0.5-4
cm long and thick; solid, firm, dry, usually hairy or downy at least at base. VEIL absent.
SPORE PRINT white to pale lilac or lilac-gray; spores 7-9 * 3-4 microns, oblong to
elliptical, smooth, not amyloid.
HABITAT: Occasionally solitary but usually in shelving masses or overlapping rows or
columns on hardwood logs and stumps; sometimes also on standing trees, rarely on
conifers; common throughout most of the northern hemisphere. Its preferred hosts
include elm, cottonwood, alder, and sycamore, but in our area it favors oak and tanoak,
producing large crops after the first fall rains and smaller crops thereafter through the
spring. I have also seen stupendous fruitings (several hundred pounds!) growing in clusters
in a treeless field where crushed coffee beans were dumped. It is easily cultivated on a wide
variety of substrates, including compressed sawdust, shredded Time magazines, and
presumably coffee grounds. If an “oyster log” is dragged home from the wild and kept
moist, it will produce crops regularly.
EDIBILITY: Edible and delicious—breaded and fried it is superb and remarkably remi¬
niscent of seafood. Be sure to check for small beetles between the gills (these can be re¬
moved by dousing the mushroom briefly in water), and of course, for maggots. The tough
stem or basal stump of tissue should be removed. Large specimens can be pounded like
abalone to make them tender. P. ostreatus and its Asian counterpart, P. sajor-cajou, are
now cultivated commercially and sold fresh under the name “tree oysters.”
COMMENTS: Pure, pale, and graceful, the oyster mushroom is easily distinguished by its
white gills, tender flesh, smooth cap, and shelflike growth habit on wood. The cap color
and position of the stem depend to some extent on the location of the fruiting body. When
growing out of the side of a log, the stem is lateral or absent, since there is no need to elevate
the cap. When growing from the top of a log, however, the stem can be central, leading to
confusion with Clitocybe. The cap is generally darker in sunlight and correspondingly paler
in dim surroundings, but distinct color forms also seem to occur, including a brown-
capped form that grows on bush lupine along the ocean and a giant thick-fleshed form
that is common on cottonwood in inland valleys. In fact, P. ostreatus has long been
recognized as a “collective” species, i.e., a group of closely related but distinct forms.
Fortunately, they all appear to be edible, so that their exact taxonomy needn’t concern you
(at least, it doesn’t concern me!). Other species: P.sapidus is now regarded as a synonym for
P. ostreatus; P. columbinus is a rare species with a bluish- or greenish-tinted cap, but is
otherwise very similar; P. cornucopiae grows in dense, upright clusters on woody debris in
So you think oyster mushrooms or “tree oysters” (Pleurotus ostreatus) always grow on trees? Left:
Clusters growing in a treeless field on decomposing coffee beans. Right: A cluster growing out of a
kitchen chair. (The owner of the chair claims to be a sloppy eater who unknowingly helped incubate
the developing mycelium by sitting in the chair. A fortuitous leak in his roof elicited this crop of
oyster mushrooms. He now waters the chair regularly!)

the Rocky Mountains and probably elsewhere. It has a lined or ridged, nearly central stalk
and depressed or funnel-shaped cap that is open (incised) on one side. It is edible when
young but often develops a bitter or unpleasant taste in age. For a smaller, thinner, white
species growing on conifers, see P. porrigens.

Pleurotus porrigens (Angel Wings)

CAP 4-8 (10) x 2-5 cm, fan-shaped to tongue- or petal-shaped; surface smooth, not viscid,
pure white to milky-white, but sometimes creamy in old age; margin at first incurved, often
lobed or wavy. Flesh very thin, pliant, white, odor and taste mild. GILLS crowded, thin,
narrow, white or yellowish, decurrent if a stalk is present. STALK absent or present only
as a narrowed, stubby white base. VEIL absent. SPORE PRINT white; spores 6-7 * 5-6
microns, nearly round, smooth, not amyloid.
HABITAT: In shelving groups or overlapping clusters on old rotting conifers, especially
hemlock; widely distributed. It is very common in the fall in the Pacific Northwest and
northern California, but I have yet to find it south of San Francisco.
EDIBILITY: Edible, but in my humble fungal opinion, bland and insubstantial. However,
some people proclaim it superior to P. ostreatus.

Left: Pleurotus porrigens, a thin white cousin of the oyster mushroom that grows on dead conifers.
Right: Pleurotus dryinus (see p. 136) resembles other oyster mushrooms, but boasts a veil, remnants
of which can be seen clinging to the margin of the cap.
136 TRICHOLOMATACEAE

COMMENTS: Also known as Pleurocybella porrigens and Pleurotellus porrigens, this

species can be distinguished from other types of Pleurotus by its thin, pliant, white fruiting
body and narrow, crowded gills. It is to rotting conifers what P. ostreatus is to rotting
hardwoods—i.e., common and cosmopolitan. The shining white fruiting bodies stand
out vividly in the forest gloom, looking so exquisitely pure and unsullied that it is easy to
see how they acquired the nickname “Angel Wings.’’Other species:/*. lignatilis(-Clitocybe
lignatilis) is a similar whitish species with narrow, crowded gills and a more prominent
stem. It frequently has a farinaceous odor and prefers hardwoods rather than conifers.

Pleurotus dryinus (Veiled Oyster Mushroom)

CAP 4-20 cm broad, broadly convex sometimes becoming plane or slightly depressed in
age, surface dry, with soft grayish fibrils or scales, but sometimes whitish or in age yellow¬
ish; margin at first inrolled. Flesh very thick, white, firm; odor mild to pungent or fragrant.
GILLS decurrent, fairly close, often veined or forking on the stalk; white, but sometimes
discoloring yellowish in age. STALK 3-10 cm long, 1-3 cm thick, usually off-center but
sometimes central; rather tough, often short, equal or tapered downward; solid, whitish.
VEIL membranous, white to grayish, forming a slight ring on stalk or leaving remnants
on cap margin or disappearing entirely. SPORE PRINT white; spores 9-12 (17) * 3.5-5
microns, elliptical, smooth, not amyloid.
HABITAT: Solitary or in small groups on hardwoods (usually living); widely distributed
but not common. Alder is a favorite host; it is also reported on oak, and I’ve found it
growing locally from the wound of a living madrone, in December.
EDIBILITY: Edible but rather tough.
COMMENTS: The thick firm flesh and soft hairs or scales on the cap are good fieldmarks;
so is the veil when it is visible (see photograph at bottom of p. 135). It might be confused
with Lentinus lepideus, which has serrated gills, or Panus strigosus, which is differently
colored. P. corticatus, Panus dryinus, and Armillaria dryina are synonyms.

Hohenbueheliapetaloides group (Shoehorn Oyster Mushroom)

CAP 2-7 x 3-7 (10) cm, spatula- to funnel-shaped or shoehorn-like when upright (i.e, split
or open on one side), fan- or petal-shaped when shelflike; tapering to a stemlike base;
surface smooth or with a whitish bloom when young and often downy toward the base,
moist to somewhat rubbery-gelatinous but not viscid except when very wet; some shade of
brown, tan, or grayish-brown; margin often lobed or wavy, at first incurved or inrolled.
Flesh pliant, usually white (sometimes watery tan). GILLS narrow, thin, crowded, deeply
decurrent, white or tinged gray, often becoming yellowish or creamy in age and often
becoming crisped (wavy) in dry weather. STALK lateral or off-center, continuous with
cap, often short (1-4 cm long), equal or tapered downward, up to 2.5 cm thick, white or
grayish; fuzzy, downy, or minutely hairy. VEIL absent. SPORE PRINT white; spores 7-9
x 4-5 microns, elliptical, smooth, not amyloid. Gills with large, thick-walled cystidia.

Hohenbuehelia petaloides group. Note the crowded gills and shoehorn-shaped fruiting body. The
white specks on the caps at left are an abnormality (probably a fungal parasite).
Hohenbueheliapetaloides group. Left: Close-up of gills. The edges often become wavy as the gills dry
out, but are not serrated (toothed). Right: These young specimens remind me of penguins. They
came up in a potting mix composed largely of wood chips.

HABITAT: Usually ingroups or small clusters on rotting or buried wood, sawdust mulch,
etc.; widely distributed. This species “complex” is common year-round in our area in
nurseries, flower pots, landscaped areas where wood chip mulch has been used, etc. It
occurs less commonly in the wild, usually on rotting conifers such as hemlock. I have seen
it in Yosemite National Park in the spring and fall.
EDIBILITY: Edible, but not choice (according to most sources); I haven’t tried it.
COMMENTS: Also known as Pleurotus petaloides, this species and its close relatives
are best distinguished by their crowded gills, brown cap, white spores, and lateral stem
or stemlike base. Terrestrial fruiting bodies are often reminiscent of upright, rolled-up
leaves or shoehorns, while those that grow shelflike on wood are usually fan-shaped as in
other oyster mushrooms. The gelatinized layer of tissue beneath the surface of the cap is
seldom evident unless the specimens are waterlogged. U nder the microscope, however, it is
often discernible. H. geogenia is a very similar species with a hazel-brown to yellow-brown
cap that differs microscopically. Like H. petaloides, it is apt to be looked for in Clitocybe,
but the cap is open or split on one side and the stalk is usually off-center. Other species:
H. atrocaerulea has a brown to bluish-black, mussel-shaped cap which is felty at least
toward the base; it grows on wood, including yucca. H. angustatus has a pale (pinkish-
buff) cap and round spores. H. mastrucatus has a grayish cap, unpleasant taste, and broad
gills that are fairly well-spaced. All of these species were formerly included in Pleurotus.

Panellus serotinus (Late Oyster Mushroom)

CAP 2.5-10 (15) cm broad, kidney- or fan-shaped; surface viscid when moist or in age,
color variable: olive-green to yellow-green, ochre, greenish-brown, or with violet tones;
margin incurved, often lobed or wavy. Flesh thick, firm, white, with a gelatinous layer
under the cuticle. GILLS adnate to decurrent, close, pale orange to ochraceous to pale
yellow, often fading in age. STALK absent or if present, laterally attached, short, and
stubby (0.5-2.5 cm long); yellow to brownish or colored like cap, but hairy or velvety. VEIL
absent. SPORE PRINT yellowish; spores 4-6 * 1-2 microns, sausage-shaped, smooth,
typically amyloid (at least in dried specimens).
138 TRICHOLOMATACEAE

HABITAT: Scattered or in shelving groups on dead hardwood logs and branches (espe¬
cially wild cherry), sometimes also on conifers; widely distributed. Fairly common in the
Pacific Northwest in the fall and winter, but I have yet to find it in our area. Like Flammu-
lina velutipes, it is a cold-weather fungus, and its appearance is usually a sign that the
mushroom season is almost over.
EDIBILITY: Edible but mediocre; it sometimes develops a bitter taste as it ages.
COMMENTS: The viscid, greenish to yellowish or violet-tinted cap and pale yellow to
orange gills, plus the short, stubby stem and growth on wood make this an easy mushroom
to recognize. According to Alexander Smith, the spores of some forms do not display
the amyloid reaction until dried out or stored in a herbarium.

Panellus stipticus
CAP 0.5-3 cm broad, spatula-, kidney-, or fan-shaped, convex to plane or depressed near
the stalk; surface dry, minutely hairy or scurfy, buff to ochre-buff, tan, brownish, or
cinnamon-brown, sometimes concentrically zoned. Flesh thin, tough, white or pale
yellowish; taste usually acrid or astringent. GILLS close, narrow, often forked, brownish
to pale cinnamon or ochre-buff; adnate to decurrent, often luminescent. STALK 0.5-2 cm
long, 3-8 mm thick, off-center to lateral, usually narrowed at base, often somewhat
flattened; same color as cap or paler (to nearly whitish). VEIL absent. SPORE PRINT
white; spores 3-5 * 1.5-3 microns, elliptical to oblong or sausage-shaped, smooth, amyloid.
HABITAT: Usually gregarious or in clusters or overlapping tiers on dead hardwoods;
widely distributed, more common in eastern North America than in the West. It occurs in
California but I have yet to find it in our area. It usually fruits in the fall, but the fruiting
bodies do not rot quickly and consequently can be found practically year-round.
EDIBILITY: Inedible due to its small size, tough texture, and bitter taste.
COMMENTS: But for its luminescent gills, this listless little wood-rotter wouldn’t
attract enough attention to merit mention. Because of the brownish gills it can be mistaken
for a Crepidotus or small Paxillus, but the spore print is white and the texture much
tougher. The species epithet refers to its use as a styptic (blood-clotter). For other listless
Panellus species, see the key to Pleurotus and Allies.

Panus conchatus (Smooth Panus; Conch Panus)

CAP 4-17 cm broad, broadly convex becoming plane or broadly depressed in age; surface
dry, smooth or minutely downy, often cracked into small scales in age; vinaceous-brown or
violet-tinted when young and moist, fading to brownish, reddish-brown, or tan in age or
as it dries; margin often wavy, at first inrolled. Flesh rather tough, firm, white; taste mild.
GILLS decurrent, fairly close, narrow, often forking near stalk, tan to buff, or when moist
often violet-tinted. STALK 2-5 cm long, 0.5-3 cm thick; off-center to lateral or sometimes
central, usually tapered downward; solid, tough, colored more or less like cap; covered with
fine hairs at least when young. VEIL absent. SPORE PRINT white; spores 5-7 * 2.5-3.5
microns, elliptical, smooth, not amyloid.
HABITAT: Solitary or in small groups or clusters on hardwood logs, stumps, and fallen
branches; widely distributed but rather infrequent in our area, where it fruits in the winter.
EDIBILITY: Tough but apparently harmless.
COMMENTS: Also known as P. torulosus, this mushroom is almost entirely violet when
young and moist, but fades in age to tan or reddish-tan. The color, plus the growth on
wood, decurrent gills, and non-hairy cap are good fieldmarks. The stem is sometimes
Panus conchatus (-P. torulosus). These specimens are brownish, but were distinctly violet or vina-
ceous when younger (just like P.rudis). Note smooth (not hairy!) cap and growth on wood.

central, leading to confusion with Clitocybe, and it is consequently keyed out under that
genus. The cap is not hairy as in P. rudis, nor do the gills have serrated edges as in
Lentinus.

Panus rudis (Hairy Panus)

CAP 2.5-10 cm broad, fan-shaped or wedge-shaped to somewhat irregular in outline,
convex becoming plane or depressed; surface dry, covered with dense, coarse, stiff,
velvety hairs, reddish-brown to tan, but often violet when fresh and wet; margin incurved,
often lobed. Flesh tough, thin, white; taste slightly bitter. GILLS decurrent, close, nar¬
row, edges entire; white, creamy, or colored like cap. STALK a short, stout plug of tissue
up to 2 cm long, off-center to lateral or sometimes central; tough, solid, hairy like the cap
and more or less same color. VEIL absent. SPORE PRINT white or yellowish; spores
5-7 * 2-3 microns, elliptical, smooth, not amyloid.
HABITAT: Usually in groups on rotting hardwood stumps and logs, widely distributed.
I have seen it several times in the fall on tanoak, but it is rather rare in our area.
EDIBILITY: Edible but very hairy. You’d do better to brush your teeth with it than eat it.

Panus rudis grows on hardwood stumps and logs. The cap is hairy, the stalk off-center to lateral.
Young specimen at left has a violet cap, mature individuals at right are tan.
140 TRICHOLOMATACEAE

COMMENTS: The hairy cap, tough texture, white spores, and short lateral to off-center
stem set this singular fungus apart. As in P. conchatus, fresh wet caps are a gorgeous deep
violet, but soon fade to reddish- or pinkish-brown.

Panus strigosus (Giant Panus)

CAP 10-40 cm broad or more, fan-shaped to broadly convex, plane, or slightly depressed;
surface dry, with coarse hairs, white to buff or creamy, discoloring yellowish in old age or
when dried. Flesh thick, rather tough, white or yellowish, taste mild. GILLS broad, white
to buff or even tinged lilac or brownish, becoming yellowish in old age; usually decurrent,
edges typically entire. STALK 2-15 cm long, 1-4 cm thick, usually off-center or lateral;
solid, tough, white to buff or aging yellowish; equal or thicker below, coarsely hairy
especially toward base. SPORE PRINT white; spores 10-13 x 3-5 microns, oblong,
smooth, not amyloid.
HABITAT: Solitary or clustered (but rarely more than four together), usually in wounds
of living hardwoods; widely distributed but rare, at least in the West. It favors maple and
birch, but occurs in Arizona on walnut. Three gigantic local specimens were brought to
me, but the collector didn’t note the host. Each cap was more than two feet in diameter!
EDIBILITY: Reportedly edible, but too rare and tough to be of consequence.
COMMENTS: But for the hairy cap this humongus fungus might be mistaken for a giant
oyster mushroom (Pleurotus ostreatus). Lentinusponderosus rivals it in size but grows on
conifers and is not as hairy. The species epithet means “strigose,” which means hairy.

Phyllotopsis nidulans
CAP 2-8 cm broad, more or less fan-shaped to scallop-shaped in outline, broadly convex
to plane; surface dry, often covered at first with a white chamois-like, cottony pubescence,
otherwise pale orange to orange-buff, yellow-orange, or fading to buff, and densely hairy
or fuzzy; margin at first inrolled. Flesh colored like cap or paler; odor typically strong and
disagreeable (like sewer gas or rotten eggs), but sometimes mild. GILLS close, narrow,
orange-buff to orange-yellow or pale orange. STALK absent or rudimentary. VEIL
absent. SPORE PRINT pale pinkish to apricot-pink to pinkish-brown; spores 5-8 x
2-4 microns, sausage-shaped, smooth, not amyloid.
HABITAT: In groups or shelving masses on rotting logs and stumps(of both hardwoods
and conifers); widely distributed. In our area it is common on dead oaks in the fall and
Phyllotopsis nidulans. A common wood-inhabitor, easily told by its hairy or fuzzy cap, orangish to
yellow-orange gills, and obnoxious odor. Note the absence of a stalk.
PHYLLOTOPSIS 141

winter, and in the Sierra Nevada and Rocky Mountains I have seen it on aspen.
EDIBILITY: Unknown. The odor is so disgusting that only a zealot with the iron con¬
stitution of Charles Mcllvaine would consider eating it.
COMMENTS: Formerly known as Claudopus nidulans and Panellus nidulans, this
rather attractive pale orange shelving mushroom is easily recognized by its peach-fuzz¬
like cap and obnoxious odor (the latter feature, however, is lacking in some collections).
Paxillus panuoides is somewhat similar, but has yellowish-buff spores and veined or
forked gills. Crepidotus species have brown spores, Panellus species have white to yel¬
lowish spores, while Claudopus species have pinkish spores but do not have orange gills.

LENTINUS & LENTINELLUS

Small to medium-sized or very large fungi usually growing on wood. CAP often hairy or scaly.
Flesh firm or tough. GILLS usually adnate to decurrent, edges usually toothed, serrated, or ragged.
STALK absent to lateral, off-center, or central. VEIL absent or sometimes present and forming
a slight annulus (ring) on stalk. VOLVA absent. SPORE PRINT white to yellowish or buff. Spores
smooth or rough, amyloid (Lentinellus) or not amyloid (Lentinus).

THESE pale-spored, wood-inhabiting agarics can usually be recognized by their ragged

or serrated gill edges (see photo on next page). Other white-spored wood-inhabitors do
not normally have serrated gills unless they are very old or weathered. In Lentinellus the
fruiting body is small to medium-sized and the spores are amyloid, while in Lentinus the
fruiting body is sometimes gigantic and the spores are not amyloid. In both genera the stalk
can be central, leading to confusion with Tricholomopsis, Clitocybe, Armiliariella, and
other wood-rotters, but more often than not it is off-center to lateral or even absent.
Both genera are small and neither is particularly common, at least in our area. Some
of the fleshier Lentinus species are edible, but by and large this group is not one to tempt
the “toadstool-tester.” A notable exception is Lentinus (-Tricholomopsis) edodes,
the renowned shiitake or “black mushroom” of Oriental cuisine. For centuries it has been
grown on log “teepees” in Japan, and more recently on sawdust mixtures. It is now being
grown commercially in North America and is sold fresh or dried in many markets and
specialty shops. Shiitake “logs” (cultivation kits) are even available for those who want to
grow their own (see photo on p. 31). In North America the shiitake has not yet been found
in the wild, but it may very well escape cultivation and establish itself on native oaks (or
other hardwoods). For this reason it is keyed out below, along with several other species.
Key to Lentinus & Lentinellus
1. Stalk typically well-developed and central to off-center or sometimes lateral .2
1. Stalk typically absent or present only as a stubby lateral point of attachment to wood, not
well-developed .Lentinellus ursinus & others, p. 144
2. Veil present when young, often leaving remnants on stalk and/or cap in age . 3
2. Veil absent (check young specimens if possible) . 6
3. Found on hardwoods along Gulf of Mexico and in tropics; fruiting body Collybia-like but
tough; gills usually staining reddish-brown when bruised .Lentinus detonsus
3. Not as above .4
4. Usually growing on conifers (including fence posts and railroad ties); cap whitish to buff or
yellow, often with darker scales .Lentinus lepideus, p. 142
4. Not as above; growing on hardwoods or cultivated commercially .5
5. Cap with a dense coating of dark brown to black hairs or small scales which become sparser
in age, revealing the whitish to buff background; veil leaving a slight ring on stalk or remaining
intact (covering the gills and never breaking) .Lentinus tigrinus
5. Cap brown to dark brown, often with whitish veil remnants (especially near margin); native to
Asia, cultivated in U.S. but not yet naturalized (see photo on p. 31) .Lentinus edodes
Gill detail in Lentinusponderosus. The serrated edges are characteristic of Lentinus and Lentinellus.

6. Cap more than 8 cm broad when mature; stalk over 1 cm thick; fruiting body tough or hard;
found on or near conifers (but wood sometimes buried!) .Lentinusponderosus, p. 143
6. Not as above; smaller . 7
7. Growing in clusters on hardwoods, stalks often fused; caps some shade of brown, often
irregular or misshapen and deeply depressed in age; not uncommon in eastern North America,
much rarer in the West .Lentinellus cochleatus
7. Not as above . 8
8. Stalk usually grooved and stuffed (inside) with soft whitish tissue; cap and stalk reddish-brown
to pinkish-brown, smooth; taste usually acrid; found on ground or woody debris; spores
amyloid .Lentinellus omphalodes
8. Not with above features . 9
9. Cap 3-8 cm broad, pinkish-tan to tan; found on conifers (e.g., Sitka spruce) along the Pacific
Coast .L entinus kauffmanii
9. Cap up to 4 cm broad, brown to orange-brown or cinnamon; found on hardwoods; widely
distributed .Lentinus sulcatus

Lentinus lepideus (Train-Wrecker)

CAP 5-15 (20) cm broad, convex to plane; surface dry or slightly viscid, whitish to buff or
pale yellow, but usually with darker (brownish) scales; margin sometimes beaded with
droplets when young. Flesh thick, tough, white, but often aging or bruising yellow; not
decaying readily, odor usually distinctive (pungent or fragrant). GILLS usually decurrent
but sometimes notched or adnate, whitish to buff or in one form yellow, often bruising
brownish and/or yellowish in age; edges entire when young but often serrated in age.
STALK 3-5 cm long, 1-3 cm thick, central to somewhat off-center, tapered at base; solid,
tough and hard, colored more or less like cap, usually with brownish to reddish-brown
scales or fibrils below ring. VEIL membranous, forming a pallid, superior to apical ring
on stalk which may be slight or disappear in age. SPORE PRINT whitish; spores 9-12
* 4-5 microns, almost cylindrical, smooth, not amyloid.
HABITAT: Solitary, scattered, or in small groups on conifer logs, stumps, fence posts, and
other lumber, sometimes also on oak; widely distributed and fairly common in cool
weather in the coniferous forests of the West, but rare in our area and less frequent than
L. ponderosus in the Sierra Nevada. It used to be common on railroad ties, resulting
in derailments and the common name, “train-wrecker.” It causes a brown rot in its host.
EDIBILITY: Edible and quite good, but the tough flesh requires thorough cooking.
Use only young caps—older specimens may have an unpleasant taste.
COMMENTS: The scaly cap, serrated gills (at least in age), membranous veil, white
spores, and growth on wood (sometimes buried!) set apart this species. Like L. ponde¬
rosus, it decays slowly, and old or weathered specimens can be difficult to recognize.
Several variants occur, including a yellow one.

142
Lentinus ponderosus. Left: Mature specimens growing from a log buried by a landslide. Exposed caps
are apt to be scalier than sheltered ones. (Bob Winter) Right: Young specimens. Note decurrent gills.
See p. 142 for close-up of the serrated gills, and p. 43 for a picture of a clump growing in a lake!

Lentinus ponderosus (Ponderous Lentinus)

CAP 10-50 cm broad or more, convex to plane or somewhat depressed; surface dry or
slightly tacky, at first smooth but in most cases soon breaking up into large scales, revealing
the white flesh beneath; color variable depending on age and exposure: white to tan,
yellowish, brownish, or pinkish-brown, usually discoloring yellowish or orangish in
age. Flesh thick, tough, not readily decaying, white (but may age or bruise yellowish); odor
often fragrant. GILLS typically decurrent, fairly close, white to yellowish, but often
developing orangish to rusty-brown stains in age; edges serrated or torn, at least at
maturity. STALK 5-20 cm long, 2-8 cm thick, central or off-center, usually with a nar¬
rowed, rooting base; solid, hard, tough, whitish aging yellowish to brown or rusty-orange,
often with brownish patches or scales. VEIL absent. SPORE PRINT white; spores 8-12
x 3-5.5 microns, elliptical, smooth, not amyloid.
HABITAT: Solitary or in groups or clusters on or near dead conifers (but often appearing
terrestrial) in the late spring, summer, and early fall; known only from western North
America. Although not common in most areas, it is very conspicuous because of its large
size. I have never seen it at low elevations, but it is often abundant in the Sierra Nevada
in the summer on lodgepole pine. I have also collected it on ponderosa pine in the South¬
west. It produces a brown rot in its host.
EDIBILITY: Edible and choice, but thorough cooking or parboiling is required because
of its toughness. Biologist Bob Winter of Fresno says that it is avidly sought byJapanese-
Americans as a shiitake- and matsutake-substitute, perhaps because of its chewy texture.
COMMENTS: This large, tough mushroom rivals Catathelasma imperialis and Clito-
cybe gigantea for the title of “Most Humongus Gilled Fungus Among Us.” The former,
however, has a veil and the latter has a fragile cap, and both are terrestrial, whereas L.
ponderosus lacks a veil and grows on or near wood. Panus strigosus can also be very large,
but grows on hardwoods and has a hairy cap and non-serrated gills. The fragrant odor of
L. ponderosus is sometimes reminiscent of the matsutake (Armillaria ponderosa), but
that species is terrestrial and has a prominent veil. Smaller specimens can be confused
with L. lepideus, which is rather similar in overall aspect but also has a veil. The cap color
and degree of scaliness vary considerably from specimen to specimen (depending on age,
temperature, and exposure to direct sunlight), but the size, toughness, and decurrent
gills with serrated edges are distinctive.
Lentinellus ursinus. Note hairy cap, ragged or serrated gill edges, and near absence of stalk.

Lentinellus ursinus
CAP 3-10 x 2-5 cm, kidney- to fan-shaped in outline, broadly convex becoming plane;
surface dry, dark brown to brown, yellow-brown, or reddish-brown, with sparse to dense,
brown to dark brown pubescence (fine hairs), at least toward the stalk; margin usually
smooth, often paler and lobed, at first incurved. Flesh thin; taste slowly acrid or bitter.
GILLS decurrent (if stalk present), close, broad, dingy white to pinkish-brown with ragged
or coarsely toothed edges. STALK absent or rudimentary. VEIL absent. SPORE PRINT
white; spores 2.5-5 x 2-3.5 microns, nearly round, with minute amyloid spines.
HABITAT: On rotting logs and stumps, usually in groups or shelving clusters; widely
distributed. It grows on both hardwoods and conifers but is not common in our area. I
have found it in the late fall and winter on Douglas-fir and live oak.
EDIBILITY: Inedible due to the bitter or acrid taste.
COMMENTS: This flaccid, fleshless, featureless fungus could carelessly be mistaken for
for a decrepit oyster mushroom (Pleurotus ostreatus) were it not for the ragged gills and
hairy cap. L. flabelliformis is a similar but slightly smaller species with whitish pubesence
on the cap. Another widespread species, L. vulpinus, also has whitish pubescence (at least
at the base of the cap), but is often ribbed or reticulate and sometimes has a stalk; it favors
hardwoods. Still another species, L. montanus, can be told by its well-spaced gills and
tendency to fruit on dead conifers, usually at higher elevations after the snow melts.

TRICHOLOMOPSIS
Medium-sized mushrooms usually found on or near rotting wood. CAP smooth orscaly, not viscid.
Flesh often yellow. GILLS attached, usually yellow. STALK typically central, fleshy. VEIL absent
or evanescent. VOLVA absent. SPORE PRINT white. Spores smooth, not amyloid. Cystidia
abundant on the edges of the gills.

TH IS is a small genus of wood-inhabiting agarics formerly distributed among Tricholoma,

Clitocybe, and Collybia. In most species the fruiting body is largely yellow, but in T.
platyphylla it is white to grayish-brown. Tricholomopsis may occasionally appear

144
TRICHOLOMOPSIS 145

terrestrial, but the yellow gills and yellow flesh, absence of a veil, and central, fleshy stalk are
distinctive. None of its members are particularly good eating. Of the species keyed below,
only T. rutilans is common in our area.
Key to Tricholomopsis
l. Lower portion of stalk dark rusty-brown to blackish-brown and velvety from a coating of
minute hairs; usually growing in tufts or clusters. (see Flammulina velutipes, p. 220)
1. Not as above . 2
2. Cap and stalk yellow, or yellow beneath a layer of colored fibrils or scales; flesh and gills pale
yellow to yellow . 3
2. Not as above . T. platyphylla & others, p. 146
3. Cap grayish to black at the center or with small grayish-brown to olive-brown to blackish
scales. T. decora (see T. rutilans, below)
3. Not as above . 4
4. Cap and/or stalk with reddish to purple-red scales or fibrils . Tricholomopsis rutilans, below
4. Cap basically yellow, without differently colored scales or fibrils, at least when young (but
may have brownish fibrils or streaks in age) T. sulfureoides & others (see T. rutilans, below)

Tricholomopsis rutilans (Plums and Custard)

CAP 3-12 cm broad, convex becoming plane; surface dry, yellow, but covered with dark red
to purple-red scales or fibrils which become sparser in age or toward the margin. Flesh
thick, firm, pale yellow, odor mild. GILLS adnate or notched, close, yellow to pale yellow.
STALK 5-10(18) cm long, 1-2.5 cm thick, equal or slightly thicker below, dry, yellow with
reddish or reddish-purple scales like those on the cap (but usually sparser and sometimes
entirely yellow in old age. VEIL absent. SPORE PRINT white; spores 5-7 * 3-5 microns,
elliptical, smooth. Cystidia on gill edges numerous, club-shaped.
HABITAT: Solitary, tufted, or in small groups on or near rotting conifers, wood chips, and
humus rich in lignin; widely distributed. Fairly common in our area in cool weather (late
fall, winter), but rarely fruiting in large numbers. I find it most often with redwood and pine.
EDIBILITY: Edible, but according to Chroogomphus-connoisseur Ciro Milazzo—
who was born and raised in Brooklyn—“it tastes like rotting wood.” Since I’ve never tasted
rotting wood, I cannot attest to the validity of this statement. I wasn’t born and raised in
Brooklyn either.

Tricholomopsis rutilans is a beautiful wood-loving agaric with dark red scales on the cap and stalk.
The scales are not always as dense or prominent as those on the young specimens pictured here. Also,
the cap tends to broaden with age. T. decora( not illustrated) has olive-brown to blackish scales that
are much sparser than those of T. rutilans.
146 TRICHOLOMATACEAE

COMMENTS: A real beauty when fresh, this mushroom is easily recognized by the dark
red or purple-red fibrillose scales on a yellow background (hence its British name, “plums
and custard”). This color combination is practically unique among fleshy-stemmed,
white-spored agarics. If you find what looks to be a small T. rutilans with an entirely
yellow stem, you probably have T. flammula, a questionably distinct species. T. decora
is also quite similar, but has gray to brownish or black cap center and/ or scales. I find it
occasionally on rotting redwood, but it is more common farther north. Finally, there are
several entirely yellow species also found on rotting conifers, including: T.flavissima, with
a fibrillose, fringed cap margin; and T. sulfureoides, partial to hemlock, with an evanescent
veil, and a cap that develops small brownish scales or streaks in age. T. rutilans was origi¬
nally placed in Tricholoma, T. decora in Clitocybe, and T. sulfureoides in Pleurotus.

Tricholomopsis platyphylla (Broad-Gill)

CAP 4-12 cm broad, convex to plane or centrally depressed; surface smooth, not viscid,
often streaked; dark brown to grayish-brown, or sometimes pallid with a darker center
and/or fibrils. Flesh pallid, thin. GILLS adnate or more often notched, well-spaced, very
broad (deep), often splitting or with eroded edges in age, white or grayish. STALK 6-12
cm long, 1-3 cm thick, equal or thicker below, white or flushed cap color, hollow in age with
a tough outer rind; base usually with white mycelial cords attached. VEIL absent. SPORE
PRINT white; spores 7-9 * 4-7 microns, elliptical, smooth. Cystidia abundant on gill edges.
HABITAT: Solitary or in small groups on or near rotting logs and stumps, especially of
hardwoods, widely distributed. In eastern North America it is common in the spring and
early summer when few other mushrooms are out and about, and it is also said to be quite
common in Arizona. In our area, however, it is rather rare and fruits in the fall and winter.
EDIBILITY: Not recommended. Some people are adversely affected by it and the flavor
is poor. Also, it is not particularly easy to identify.
COMMENTS: This mushroom has few obvious relatives and has consequently been
placed in several different genera, including Collybia and Oudemansiella. It is somewhat
reminiscent of Pluteus cervinus, but does not have pinkish spores. If not clearly growing on
wood it can be mistaken for a Tricholoma or robust Collybia, but the very broad, fre¬
quently eroded gills and white mycelial cords (rhizomorphs) are distinctive. The latter
may only be evident if the mushroom is dug out carefully and completely, and even then are
sometimes absent. T.fallax is a closely related species with yellowish-tinted gills and stalk,
found on conifers in the Rocky Mountains.

OMPHALOTUS
Golden-yellow to olive-yellow to bright orange mushrooms growing from hardwood trees, stumps,
and roots; often clustered. CAP smooth. GILLS well-developed, with acute edges, typically
decurrent, often luminescent when fresh. STALK central or off-center, fleshy. VEIL and VOLVA
absent. SPORE PRINT white or tinged yellow. Spores smooth, not amyloid.

POPULARLY known as “jack-o-lantern mushrooms,” these are brightly colored agarics

with a fleshy stem and decurrent gills that often glow in the dark. Their luminescence is best
seen by sitting alone in a dark closet with the mushroom while eating a grilled cheese
sandwhich. Unless you are a voracious eater, this method helps combat boredom while
allowing your eyes to adjust to the darkness. After a few minutes an eerie silvery-green
glow will become visible, growing gradually brighter with each bite (of the cheese sand¬
which) until each gill is clearly outlined. Fresh, actively-sporulating specimens glow the
brightest, but even they will not always cooperate.
OMPHALOTUS 147

Jack-o-lantern mushrooms are strictly wood-inhabitors, but frequently appear terres¬

trial because they like to grow on roots or old, buried stumps. Two to four species occur in
N orth America, but only one on the west coast. They contain muscarine and are poisonous.

Key to Omphalotus
1. Fruiting body pumpkin-colored (bright yellow-orange to orange); common in eastern North
America, Mexico, possibly the Southwest .O. olearius(see O. olivascens, below)
1. Fruiting body golden-yellow to yellow-orange, but usually toned with olive (sometimes
other colors also present); restricted to the west coast.O. olivascens, below

Omphalotus olivascens Color Plates 40,41

(Jack-O-Lantern Mushroom; Western Jack-O-Lantern Mushroom)
CAP 4-16 (25) cm broad, broadly convex becoming plane or depressed; surface smooth,
not viscid, color variable: bright golden-yellow to orange with olive tones often present
also, varying to dull orange, brownish-orange, olive, or slightly reddish. Flesh rather
thin, pliant, colored more or less like cap; odor mild. GILLS olive to bright yellow-orange
(often yellow with olive tints), decurrent, close, usually luminescent when fresh. STALK
4-20 cm long, 1-4 (8) cm thick, central to off-center, equal or tapered downward, solid,
dry, more or less colored like cap or gills, or dingier olive. SPORE PRINT white to yellow¬
ish; spores 6-8 x 5.5-7 microns, elliptical to nearly round, smooth.
HABITAT: In tufts or clusters or occasionally solitary on or around hardwood trunks,
stumps, and buried wood; known only from the west coast, but replaced elsewhere by
O. olearius (see comments). Common in our area from fall through early spring, especially
on oak, manzanita, madrone, and chinquapin.
EDIBILITY: Poisonous! Profuse sweating and gastrointestinal distress are typical symp¬
toms; muscarine is one of the toxins (see p. 894). It is sometimes eaten under the mistaken
impression that it is a chanterelle.
COMMENTS: The bright yellow-orange to olive color, decurrent gills, pale spores, and
tendency to grow in clusters distinguish this handsome mushroom. The chanterelle
(Cantharellus cibarius) is somewhat similar but has thick, shallow, blunt, foldlike gills and

Omphalotus olivascens often—but not always—grows in clusters. The gills are well-developed,
decurrent, and have thin edges. Entire fruiting body is golden-yellow to orange or olive (see color
plates), including the flesh.
148 TRICHOLOMATACEAE

white flesh; the false chanterelle (Hygrophoropsis aurantiacus) is smaller and has oranger,
repeatedly forked gills, while Gymnopilus species have dark orange to rusty-brown spores.
All lack the olive tones characteristic of mature O. olivascens.
The common jack-o-lantern mushroom of eastern North America is essentially the same
as O. olivascens except that it is pumpkin colored (bright orange to yellow-orange, without
any olive tones). Formerly called Clitocybe illudens, it is now called O. olearius (or O. il-
ludens by those who consider it distinct from the O. olearius of Europe). According to one
report, its luminescence is sometimes bright enough to read a newspaper by. And then
there’s the tale of the shipwrecked sailor on an uninhabited island, who wrote a last message
by the light of a jack-o-lantern mushroom, using the ink from a shaggy mane and the stalk
of an Agaricus as a pen. Unfortunately, he starved to death because he was afraid to eat any
of the mushrooms he found!

CLITOCYBE & Allies

Fairly small to large mushrooms found mostly on the ground, sometimes on rotten wood. CAP
convex to plane or often depressed to funnel-shaped at maturity, rarely viscid. GILLS usually
adnate to decurrent and usually white to gray pr buff. STALK central, usually fleshy, but often
slender. VEIL and VOLVA absent. SPORE PRINT white to buff, yellowish, or dull pinkish.
Spores smooth or roughened but not ridged or angular; usually not amyloid.

THIS is a large and complex group of soft, fleshy, pale-spored mushrooms with no veil
and a central, usually fleshy stem. The spore color is typically white, buff, or yellowish, but
in some species—at one time honored with their own genus, Lepista—it is dull or pale
pinkish. How, then, can you separate Clitocybe from other pale-spored mushrooms?
Mostly by a process of elimination: the gills are not soft and waxy as in the Hygrophora-
ceae, nor orange as in Hygrophoropsis and Omphalotus, nor thick, blunt, shallow, and
foldlike as in Cantharellus; the flesh is not granular and brittle as in Russula, there is no
latex as in Lactarius, and the white-spored species do not have the notched gills
characteristic of Tricholoma—though this is a somewhat capricious character, since the
attachment depends to some extent on the age and shape of the cap; the “Lepistas” are easily
confused with the Entolomataceae (particularly the genus Entoloma), but the latter have
deeper, more vividly colored spores which are angular or longitudinally ridged under the
microscope; the wood-inhabiting Clitocybes generally have a central stalk, thus
eliminating Pleurotus, Panus, and other shelflike types; the smaller Clitocybes with slender
stems are apt to be mistaken for Omphalina, in which the fruiting body is very small, and
Colly bia, which has a cartilaginous stem of a different texture from the cap, and adnexed to
adnate but not decurrent gills; finally, there are a number of small but common genera
(e.g., Laccaria, Leucopaxillus, Lyophyllum) that are best distinguished by learning the
individual species. Whew!!
Except for the blewit (C. nuda) and its close relatives, Clitocybe is a lackluster group
whose aesthetic and gustatory value is practically nil. Confirmed Clitocybe experts will
be the first to admit that the anonymous throngs of white to grayish Clitocybes that litter
our wintertime woods are exceedingly difficult to differentiate. Clitocybes are most preva¬
lent in coniferous forests, but also occur under hardwoods and in grass or manure. Like
the Tricholomas, they are largely cold weather fungi, most abundant in our area from
December through February.
Though the blewit is a safe and popular edible mushroom, several Clitocybes are
poisonous, including the small grass-inhabiting species, C. dealbata. The larger forms
are slightly easier to identify but may be just as difficult to digest. Several have a disagree-
CLITOCYBE & ALLIES 149

able odor, notably C. nebularis and C. robusta. On the other hand, the anise-scented types
(e.g., C. deceptiva and C. odora) are edible and quite good.
Over 200 species of Clilocybe occur in North America. About 50 are listed for California,
but a diligent effort to catalog our Clitocybes would probably double that number. Only
some of the more easily identified species are described here, including a few smallish
types with amyloid spores that are now placed in the genera Myxomphalia, Clitocybula,
and Cantharellula. If your “Clitocybe” does not key out satisfactorily, check the Hygro-
phoraceae (if the spores are white) or the Entolomataceae (if the spores are pinkish).

Key to Clitocybe & Allies

1. Odor distinctly licorice- or aniselike .2
1. Not as above (but odor may be sweet, e.g., like root beer) .. 3
2. Fruiting body entirely or partially blue-green to greenish or grayish-green . . C. odora, p. 161
2. Fruiting body lacking blue or green tints .C. deceptiva & others, p. 162
3. Spore print pink or pinkish; fruiting body often (but not always) purple .4
3. Spore print white to yellowish or buff, or tinged lilac or brownish; fruiting body not commonly
purple, and if purple then growing on wood . 13
4. Stalk 3-7 mm thick; growing in grass, manure, straw, etc.C. tarda, p. 152
4. Stalk thicker, or growing in woods . 5
5. Spore print bright pink; cap and stalk dingy cinnamon to vinaceous-brown; gills decurrent;
growing in woods (especially pine), mainly in eastern North America . C. martiorum
5. Not as above; spore print dull pinkish to pinkish-buff or flesh-colored . 6
6. Gills (and usually rest of fruiting body) distinctly bluish-purple to purple or pale purple (lilac)
when fresh (but often fading in age); very common . C. nuda, p. 153
6. Not as above; gills not distinctly purple . 7
7. Stalk distinctly purple to lilac-tinged when fresh. C. saeva(see C. nuda, p. 153)
7. Not as above . 8
8. Fruiting body with subtle vinaceous or lilac tints when fresh .
. C. glaucocana& C. graveolens(see C. tarda, p. 152)
8. Not as above .9
9. Fruiting body white when fresh, soon developing rusty to reddish stains .
.(see Collybia maculata, p. 217)
9. Not as above . 10
10. Cap pinkish to orangish to orange-brown or reddish-brown; spores angular in end view ....
.(see Entolomataceae, p. 238)
10. Not as above; cap white to watery brown, tan, hazel, buff, pinkish-buff, gray, etc. 11
11. Typically growing in clusters .C. subconnexa group & others, p. 155
11. Typically growing scattered to gregarious or in rings . 12
12. Cap watery brown to tan, hazel, buff, or whitish; growing in pastures, lawns, or sometimes
in woods (especially oak) .C. brunneocephala & others, p. 154
12. Cap white becoming pinkish-buff to dingy buff or tan in age; growing in woods; widespread
.C. irina(see C. brunneocephala, p. 154)
13. Growing on burnt ground or debris; cap small (less than4 cm), blackish to grayish-brown to gray
or olive-brown .Myxomphalia maura & others, p. 165
13. Not growing in burned areas, or if so, then very differently colored . 14
14. Gills forked repeatedly and usually orange or odor very fragrant (somewhat like root beer)
.(see Paxillaceae, p. 476)
14. Not as above . 15
15. Growing on wood (sometimes very rotten or buried) or on coffee grounds . 16
15. Growing on ground . 24
16. Gills bright yellow to orange; fruiting body small . (see Omphalina& Xeromphalina, p. 221)
16. Not as above . 17
150 TRICHOLOMATACEAE

17. Fruiting body entirely or partially vinaceous, purplish, or reddish-tinged when fresh and moist
.(seePleurotus& Allies, p. 132)
17. Not as above . 18
18. Cap open or incised on one side; gills very crowded, narrow . (seePleurotus& Allies, p. 132)
18. Not as above . 19
19. Gill edges serrated or eroded and/ or fruiting body tough, hard, fairly large to very large (cap
10-50 cm broad when mature; stalk 2-5 cm thick) .(seeLentinus& Lentinellus, p. 141)
19. Not as above . 20
20. Cap ochre-brown to pinkish-brown or cinnamon when fresh, not typically growing in clumps or
dense clusters .C. americana& others (see C. inversa, p. 156)
20. Not as above; cap differently colored (brownish to gray or white), or growing in clumps . 21
21. Stalk white, usually tough and short, at least 1 cm thick; cap without hairs or scales, often pale;
spore print frequently tinged lilac .(seePleurotus ostreatus, p. 134)
21. Not as above . 22
22. Typically growing in clumps or dense clusters on wood; cap not white(or if watery whitish, then
stalk usually rather long and less than 5 mm thick) . 23
22. Typically growing solitary, scattered, or in small groups but not clumps; cap variously colored
.24
23. Stalk typically less than 5 mm thick; growing on rotting conifers .
.(see Clitocybulafamilia& C. abundans under Collybia acervata, p. 215)
23. Stalk generally thicker than above; found in southern United States, usually on hardwoods .
. (see A rmillariella tabescens under A. mellea, p. 196)
24. S pore print brownish; fruiting body small and white or grayish; cap often somewhat hairy, espe¬
cially toward margin; not common Ripartites (R. tricholoma is the most widespread species)
24. Not as above . 25
25. Stalk thin (usually 1-3 mm thick); fruiting body small; cap white or yellow, pinkish, vinaceous,
or tinged faintly gray.26
25. Not as above; stalk thicker or fruiting body differently colored.27
26. Growing on ground in groups or troops under pine or other conifers or growing on logs, sticks,
berry canes, etc.; gills very widely spaced .(steMarasmius, Collybia, & Allies, p. 201
26. Not as above .(see Omphalina& Xeromphalina, p. 221)
27. Cap orange-buff to orange-brown to reddish-brown, cinnamon, purplish-brown, pinkish-
brown, pinkish-tan, or tan when fresh . 28
27. Cap white to buff, grayish, olive-gray, olive-brown, greenish, brown, or darker .34
28. Gills widely spaced . (see Camarophylluspratensis, p. 110)
28. Gills fairly close to crowded . 29
29. Fruiting body medium-sized to large; stalk 1 -4 cm thick; cap usually 7 cm or more broad when
mature and at that stage usually depressed or funnel-shaped .C. maxima, p. 157
29. Fruiting body smaller, thinner, or differently shaped; not as above . 30
30. Usually growing in grassy areas or lawns; stalk typically 5 mm thick or less; cap not usually
depressed .(ste Calocybe carnea, p. 176)
30. Not as above; growing in woods . 31
31. Cap vinaceous-red to purplish-red, gills yellow-ochre; growing at high altitudes under conifers
. (see Calocybe onychina under C. carnea, p. 176)
31. Not as above . 32
32. Gills white to pale buff; cap tan to pinkish-tan, flesh-colored, etc.33
32. Gills often pale pinkish-cinnamon or colored like cap in age; cap orange to orange-brown,
cinnamon, reddish-brown, reddish-tan, etc.C. inversa & others, p. 156
33. Stalk white, buff, or tinged only slightly with the cap color .C. gibba, p. 157
33. Stalk colored like cap or darker. C. squamulosa (see C. gibba, p. 157)
34. Spore print pale yellowish to buff; odor rancid; stalk at least 1 cm thick . 35
34. Spore print white, or if yellowish to buff, then not as above . 36
35. Cap white .C. robusta(see C. nebularis, p. 159)
35. Cap grayish to buff or brownish .C. nebularis, p. 159
CLITOCYBE & ALLIES 151

36. Growing in grass, straw, or compost, often in groups or rings but not in massive clusters; cap
usually less than 5 cm broad, white to grayish, buff, or tinged pinkish . 37
36. Not as above; growing in woods or under trees, or differently colored . 38
37. Cap broadly convex to plane or umbonate; gills usually notched but sometimes adnate or very
slightly decurrent; spores amyloid; not often growing in rings .... (seeMelanoleuca, p. 169)
37. Cap convex becoming plane or depressed; gills adnate to decurrent, grayish-white to buff or
pinkish-buff; spores not amyloid; often growing in rings .C. dealbata & others, p. 163
38. Cap white or whitish . 39
38. Not as above .41
39. Typically growing in large clusters along roads and paths in the Pacific Northwest and Rocky
Mountains .C. dilatata, p. 159
39. Not as above .40
40. Cap medium-sized to very large (8 cm broad or more) . . C. candidate C. gigantea, p. 158)
40. Cap smaller, generally less than 8 cm broad . . C. variabilis & others (see C. albirhiza, p. 161)
41. Cap and/ or gills greenish to bluish-green, more than 2 cm broad; growing in woods in eastern
U.S. or under western mountain conifers .... C. aeruginosa & others (see C. odora, p. 161)
41. Not as above .42
42. Growing in tight clumps (occasionally solitary) from a fleshy mass of tissue which is often buried
. C. sclerotoidea, p. 164
42. N ot growing in clumps from a fleshy mass of tissue .43
43. Base of stalk or surrounding humus with conspicuous white mycelial threads; common under
mountain conifers, especially as or just after the snow melts .C. albirhiza, p. 161
43. Not as above .44
44. Gills gray; cap with a hoary bloom when young; growing under mountain conifers, usually near
melting snow . (seeLyophyllum montanum, p. 175)
44. Not as above .45
45. Base of stalk (or flesh in base) pinkish to pale orange . . (see Tricholoma saponaceum, p. 184)
45. Not as above .46
46. Gills forked repeatedly; cap grayish to grayish-brown; gills sometimes reddish-stained; usually
growing in moss in northern and eastern North America .
.Cantharellula umbonata(see Clitocybe cyathiformis, p. 164)
46. Not as above .47
47. Stalk very thick (2-5 cm); gills whitish to buff to dingy tan or yellowish, but not gray .... 48
47. Not as above . 50
48. Cap 10-40 cm broad, white becoming buff or dingy brownish in age, thin and easily broken at
maturity, margin often obscurely ribbed . C. gigantea, p. 158
48. Not as above .49
49. Odor strongly unpleasant; cap grayish to dingy tan C. septentrionalis(see C. gigantea, p. 158)
49. Not as above; cap more or less grayish-brown . C. crassa(see C. nebularis, p. 159)
50. Stalk less than4 mm thick; cap small(up to 5 cm broad but usually less than3 cm), dark greenish
to olive-brown, sooty-brown, ashy-gray, or blackish . 51
50. Not as above; usually larger .53
51. Cap minutely scaly or scurfy at center .... C. epichysium(see Myxomphalia maura, p. 165)
51. Not as above . 52
52. Cap and gills yellow-green to green, dark green, olive, etc.
.C. atroviridis & others (see C. odora, p. 161)
52. Not as above; not greenish . Omphaliaster & Fayodia spp. (see Myxomphalia maura, p. 165)
53. Stalk clothed with dark scurfy scales; gills well-spaced; cap dark brown to blackish; growing on
rotten wood or in rich humus.Clitocybula atrialba(see Clitocybe cyathiformis, p. 164)
53. Not as above; stalk not clothed with dark scurfy scales . 54
54. Cap typically incised or open on one side; gills crowded and narrow (shallow) .
.(see Hohenbueheliapetaloides group, p. 136)
54. Not as above . 55
152 TRICHOLOMATACEAE

55. Typically growing in clusters; stalk usually 1 cm thick or more .56

55. Not as above . 57
56. Cap tan to brown or purple-brown; spore print off-white to pale buff; known from Alaska . .
.C. polygonarum (see C. subconnexa group, p. 155)
56. Cap gray, brown, tan, etc.; spore print white; widespread . (seeLyophyllum& Allies, p. 173)
57. Gills distinctly grayish to grayish-brown, at least at maturity . . v.58
57. Gills white to buff or yellowish . 59
58. Stalk 1 cm thick or more at apex; cap6-15 cm broad or more C. harperi(seeC.nebularis,pA59)
58. Stalk usually less than 1 cm thick; cap 2-5 (7) cm broad .C. cyathiformis, p. 164
59. Stalk 1 -3 cm thick at apex; usually growing on or near rotting wood .
. C. avellaneialba (see C. clavipes, p. 160)
59. Not as above; stalk up to 1.2 cm thick at apex; growing on ground.60
60. Cap brown to grayish-brown or olive-brown . 61
60. Cap paler .C. coniferophila & others (see C. albirhiza, p. 161)
61. Gills distinctly decurrent; cap usually depressed at maturity .... C. clavipes & others, p. 160
61. Gills only very slightly decurrent if at all; cap broadly convex to plane or slightly umbonate
.(seeMelanoleuca, p. 169)

Clitocybe tarda
CAP 1-6 (9) cm broad, convex with an incurved margin, then plane to broadly funnel-
shaped or at times umbonate; surface smooth, not viscid, flesh-colored to brownish or
grayish with a faint lilac or vinaceous tinge when moist, fading as it dries. Flesh thin, odor
mild or slightly fragrant. GILLS adnate to slightly decurrent or at times notched, close,
grayish to brownish-buff or pinkish-buff, often with a lilac tint when fresh. STALK 2-6 cm
long, 3-8 mm thick, usually slender, equal or slightly thicker below, colored more or less
like cap or paler, fibrillose. SPORE PRINT dingy pale pinkish; spores 6-8 * 3-5 microns,
elliptical, finely roughened.
HABITAT: Scattered to gregarious or clustered or sometimes in rings in grass, dung,
manure, straw heaps, old fields, compost piles, etc. Widespread and not uncommon in
our area after heavy rains, late fall through spring.
EDIBILITY: Edible but thin-fleshed and not particularly easy to identify. I haven’t tried it.
COMMENTS: Also known as Lepista tarda and Tricholoma sordidum, this is a smaller,
slimmer version of the blewit. It isn’t purple, but when fresh and moist it often has a slight
lilac or vinaceous hue—especially the gills. It can easily be mistaken for a Melanoleuca,
but the spore print is pale pinkish. The cap color is difficult to characterize but is generally
some shade of buff, brown, or even gray. Two other pinkish-spored “Lepistas” with subtle
purple tints are: C. graveolens, with a strong, disagreeable odor (like “moldy hay”), and
C. glaucocana. Both of these are much larger and more robust than C. tarda, and in fact
they closely resemble the blewit (C. nuda) but for their subtler color (older specimens can
scarcely be distinguished). Both species are rare and fortunately, not poisonous.

Clitocybe tarda is a slim, trim version of the blewit, but has only a slight violet tinge (if any), and is
thus easily confused with other grayish or brownish Clitocybes. The spore print, however, is pinkish.
Clitocybe (-Lepista) nuda, the blewit, has a characteristic shape that is hard to describe but easy
to recognize. Note stocky build and inrolled margin when young. The monstrosity on the left is a stem
which kept growing after the cap was cut off.

Clitocybe nuda (Blewit) Color Plate 32

CAP 4-14 (18) cm broad, convex with an inrolled margin when young, becoming broadly
umbonate to plane, or with an uplifted, often wavy margin in age; surface smooth, lubri¬
cous when moist but not viscid, often somewhat lustrous when dry; purple, or purple
shaded with brown or gray when fresh, soon fading to brownish, flesh-color, tan, etc.,
but the margin often retaining purple tones well into maturity. Flesh thick, rather soft, pur¬
plish to lilac-buff; odor faintly fragrant when fresh(like frozen orangejuice), taste pleasant
to slightly bitter. GILLS close, adnate to adnexed or notched, or sometimes decurrent;
purple or pale purple to bluish-purple or grayish-purple when fresh, fading to buff, pinkish-
buff, or brownish in age. STALK 2.5-7 (10) cm long, 1-2.5 (3) cm thick at apex, equal or
more often with an enlarged base; dry, fibrillose, purple to pale purple or colored like the
gills; base often covered with downy purple mycelium. SPORE PRINT dull pinkish to
pinkish-buff; spores 5.5-8 x 3.5-5 microns, elliptical, roughened.
HABITAT: Scattered to gregarious, often in rings or arcs—in woods, brush, gardens,
compost piles, i.e., wherever there is organic debris; widely distributed. It seems to favor
cool weather but is common in our area throughout the mushroom season. A favorite
abode is in brambles under live oak, often in the company of chanterelles; it is also common
under pine and cypress, and I have found it on Ano Nuevo Island in beach grass and
elephant seal dung. A single mycelium will produce several crops a year, so check your
patches regularly. I know of one fairy ring sixty feet in diameter that produces about 200
blewits each time it fruits! Known as a“trash inhabitor” because of its fondness for virtually
any type of decomposing organic matter, it can be grown on a wide range of substrates,
including shredded newspapers and compost.
EDIBILITY: Edible and very popular—a favorite with beginners and gourmets alike,
and one of the most plentiful edible wild mushrooms in our area. It has the dubious dis¬
tinction of being one of the few purple foods that actually tastes good. It is even popular
in fungophobic England and Scotland, where it is sometimes sold in markets.
COMMENTS: The ubiquitous blewit is the quintessential embodiment of spunk and
persistence—cut one down and two will grow back! Decapitated stems will often continue

153
154 TRICHOLOMATACEAE

to grow as if nothing had happened—a new cap will not form, and a grotesque (but edible)
cancerous-looking pale purple growth will take its place. The blewit’s trademarks are its
beautiful purple to bluish-purple color with inrolled cap margin when young, stout stature,
absence of a veil, faintly fruity fragrance, and dull pinkish spores. The cap has a character¬
istic lubricous feel when moist, but may look quite different—polished and silvery-violet—
when dry. The amount of purple present varies considerably depending on the age and
moisture content of the mushroom, and possibly the habitat or geographical area (some
forms, such as the one commonly found under cypress, tend to be quite pale, with only a
slight violet tinge). Old faded blewits are barely recognizable, but by that stage are usually
bug-bitten anyway.
Other purple mushrooms include: Inocybe lilacina, with brown gills (when mature),
brown spores, and a small umbonate cap; many Cortinarius species, with a cobwebby veil
when young and rusty-brown spores; Mycena pura, small and slender with white spores;
the Laccaria amethystina group, with white or lilac-tinged spores and a long, tough,
fibrous stem. Of these, only the Inocybe and possibly the Mycena are poisonous. There are
also several bluish Entoloma and Leptonia species, but they are not nearly as purple.
Synonyms for the blewit are almost as numerous as the blewit itself. They include:
Tricholoma nudum, Rhodopxillus nudus, Lepista nuda, and incorrectly, Tricholoma
personatum. “Blewit,” incidentally, is a corruption of “blue hat”—though the blewit is
more purple than blue. In Europe the blewit is often called the“ wood blewit,” to distinguish
it from the “field blewit” or “blue-leg,” C. saeva (-Lepista saeva, Tricholoma personatum).
The latter is very similar to C. nuda in shape and stature and is equally delicious, but shows
purple only on the stem—the cap and gills being grayish to pinkish-buff to watery tan (or
the gills tinged vinaceous). Also, it tends to grow in pastures or grass rather than in the
woods. It is infrequent in North America but has been reported from California. See also
C. tarda, and the species discussed under it.

Clitocybe brunneocephala
CAP 4-13 cm broad, convex with an inrolled margin becoming broadly umbonate to plane
or uplifted; surface moist or lubricous but not viscid, smooth, watery brown to tan, hazel,
buff, or even whitish (usually darker when young). Flesh thick, pallid, odor mild or
pleasant. GILLS usually notched but often adnate or slightly decurrent, close, buff to
grayish-buff or pale brown, then dusted pinkish with spores. STALK 2-5(10) cm long, 1-3
(4) cm thick (usually about 2); equal or enlarged below, often stout and relatively short,
solid, dry, smooth, buff or colored like the cap (but usually paler). SPORE PRINT rosy-
buff or dull pinkish; spores 5-8 * 3-4 microns, elliptical, minutely roughened.
HABITAT: Scattered to gregarious, often forming fairy rings, late fall through early
spring, mainly in lawns and pastures, but also at the edges of woods or under trees (cypress,
oak, etc.); known only from California. It was very abundant in our area during the warm
and wet winter of 1977-78, but has been rather rare since.
EDIBILITY: Edible and quite good—I have tried it. Be sure not to confuse it with
poisonous Entolomas, however, or C. olesonii (see comments below).
COMMENTS: Listed in the first edition as “Lepista sp. (unidentified),” this interesting
relative of the blewit has recently been rechristened C. brunneocephala by clitocybiologist
Howard Bigelow. The shape, stature, and characteristically lubricous feel of the cap when
moist are very reminiscent of the blewit and blue-leg, but there is no purple anywhere on
the fruiting body. The gills are usually notched, but as in most “Lepistas” their attachment
varies considerably. I nearly always find it growing in grass, but a very similar species with
a pungent or unpleasant odor, C. olesonii, is common under oak in southern California
Clitocybe brunneocephala is built like a blewit but is never purple. It is sometimes common locally
in lawns and pastures as well as under oaks.

and the Sierra Nevada foothills. Both C. brunneocephala and C. olesonii (whose edibility
I haven’t determined) are larger and stouter than C. tarda, and do not grow in clusters
like C. subconnexa and C. densifolia. They can be separated from most Entolomas by their
duller spore color, non-angular spores, and in the case of C. brunneocephala, by the grass¬
land milieu. Other pinkish-spored species: C. praemagna, a western prairie and sagebrush
species, is quite similar but has a white cap when young that becomes tan to dull brownish
in old age; C. (-Lepista) irina is a widespread woodland species that often has a fragrant
blewit-like odor. Its cap is white to pinkish-buff, dingy buff, or pale tan, and its spores
are slightly larger than those of C. brunneocephala.

Clitocybe subconnexa group

CAP 3-10 cm broad, convex with an incurved margin becoming plane or with uplifted
margin; surface smooth, dry, satiny white at first, often discolored or spotted slightly in
age. Flesh thick, whitish, rather brittle; odor usually mild or faintly pleasant. GILLS
adnate to decurrent, crowded, narrow, pallid soon becoming buff, then dull pinkish as
the spores ripen. STALK 3-10 cm long, 0.5-2 (3) cm thick, equal or thicker below, smooth,
dry, dull grayish to buff with a whitish silky-fibrilose coating. SPORE PRINT pinkish-buff
or flesh-colored; spores 4.5-6 * 3-4 microns, elliptical, minutely roughened.
HABITAT: In groups on ground, usually tufted or clustered, widely distributed. In our
area both this species and C. (see comments) are fairly commonfromfall through
early spring in woods or at their edges, brushy areas, and open places.
EDIBILITY: Edible, but not recommended. It is good when fresh according to some
sources, but can develop an unpleasant astringent taste in age. There is also the possibility
of confusing it with C. dilatata or a poisonous Entoloma.
COMMENTS: This common species can be identified by its whitish color, adnate to
decurrent gills, dull pinkish spores, and tendency to grow in clusters—though solitary
fruiting bodies can be found. A very similar and equally common species, C. densifolia, also
grows in clusters, but has smaller spores, narrower gills, and a whitish cap that becomes
dingy buff to grayish in age. Both species, like the blewit, seem to grow almost anywhere.
They are larger and fleshier than Clitopilus prunulus and do not smell as nice, nor have
they longitudinally ridged spores. They differ from Clitocybe brunneocephala in their
more typically decurrent gills and clustered growth habit, and from C. dilatata in their
pinkish spores. Other species: C.fasciculata(-Tricholomapanaeolum var. caespitosus) is

155
Clitocybe subconnexa group gives a dull pinkish spore print, has adnate to decurrent gills, and
grows in clusters in a wide variety of habitats.

also very similar, but has a rancid-farinaceous odor and taste and tends to grow in clusters
along roads; C. subalpina is a brown to dark brown clustered species known from the
Pacific Northwest; C. polygonarum has a tan to brown or purple-brown cap, but has
off-white to pale buff spores; it also grows in clusters and is common in Alaska.

Clitocybe inversa
CAP 2-10 cm broad, broadly convex or centrally depressed with an incurved margin,
becoming broadly depressed or even funnel-shaped in age; surface dry, dull orange to pale
orange-brown, orange-tan, tan, reddish-tan, ochre-buff, or cinnamon-brown; margin
often paler. Flesh thin; odor mild or sharp. GILLS distinctly decurrent, close, buff to pale
pinkish-cinnamon or colored like cap but paler. STALK 3-10 cm long, 4-8 mm thick, equal
or thickened below, typically rather slender and often curved, colored like cap or paler,
smooth or with whitish hairs at base. SPORE PRINT white to creamy-yellowish; spores
4-5 x 3.5-4 microns, nearly round, minutely prickly (or appearing smooth).
HABITAT: Scattered to gregarious or tufted on ground in woods; widespread but
particularly common on the west coast, from Alaska to southern California. In our area it is
fairly common in the late fall and winter in mixed woods and under oak or pine, sometimes
in large fairy rings.
EDIBILITY: Not recommended

Clitocybe inversa is a cheerful orange-brown to cinnamon color. Left: Several typical examples.
Right: Close-up of the decurrent gills.
CLITOCYBE 157

COMMENTS: Also known as Lepista inversa, this species can be told by its cheerful
color, depressed cap, decurrent gills, and white or pale spores. C. gibba is rather similar
but differently colored, while Rhodocybe nuciolens has pinkish spores. The common form
of C. inversa in our area has a slightly yellowish spore print and sharp, spicy or pepperlike
odor. It may actually be C. flaccida, a very similar species. In eastern North America C.
gilva is quite common; it differs in having a plane to only slightly depressed, yellowish to
dull pinkish cap. Other similarly- colored Clitocybes include: C. sinopica, in humus or on
burnt ground, with an orange-brown to rusty-cinnamon cap, yellowish spores, and
farinaceous odor and taste; C. ectypoides, northern, growing on rotting conifers, with a
minutely scaly ochre-brown cap; and C. americana, the most common and widespread
wood-inhabiting Clitocybe, with a watery brown to pinkish-brown or cinnamon cap that
fades to whitish as it dries, growing on hardwood stumps and logs.

Clitocybe gibba (Funnel Cap) Color Plate 34

CAP 3-8 cm broad, plane or with a central depression, soon becoming funnel-shaped;
surface smooth, not viscid, tan to pinkish-tan, flesh-colored, or pinkish-cinnamon, but
fading in age; margin often wavy. Flesh thin, whitish; odor mild or faintly sweet (like
cyanide). GILLS deeply decurrent, crowded, white or pale buff. STALK 3-8 cm long, 0.4-1
cm thick, usually rather slender, equal or thicker below, smooth or faintly fibrillose; whitish
to buff, the base often with white down. SPORE PRINT white; spores 5-8 x 3.5-5 microns,
elliptical, smooth.
HABIT AT: S olitary to scattered or in small groups on ground in woods, especially under
oak but also with conifers; widely distributed. In coastal California it fruits in the winter
and early spring but is not common; in New Mexico it is common in August along with
C. squamulosa(see comments).
EDIBILITY: Edible and excellent, but not a good mushroom for beginners—too many
species of unknown edibility resemble it. Many mushroom books rate it as mediocre, but
it’s probably a case of each author taking another’s word for it.
COMMENTS: The pale pinkish-tan cap which is funnel-shaped at maturity plus the
crowded, whitish, decurrent gills and pallid slender stem render this species distinct.
C. infundibuliformis is a lengthy synonym. It is paler than C. inversa, especially in its gill
color, and more slender than C. maxima. C. squamulosa is a very similar species in which
the stalk is the same color as the cap or darker. It is a prominent feature of our western
coniferous forests, particularly at higher elevations in the spring and fall. It is interesting
to note that C. gibba releases hydrogen cyanide gas into the atmosphere. So does the
common fairy ring mushroom, Marasmius oreades, as well as several other mushrooms
and plants such as lupine and almond. The gas is not released in sufficient quantity to
harm humans, however.

Clitocybe maxima (Large Funnel Cap)

CAP (4) 7-20 (30) cm broad, at first broadly convex or plane with an inrolled margin,
becoming depressed or broadly funnel-shaped; surface dry, usually smooth, pale pinkish-
tan or flesh-colored varying to reddish-tan, sometimes spotted; margin sometimes paler
and sometimes lobed or radially ribbed. Flesh thin, white, firm becoming flaccid; odor mild
or rather unpleasant. GILLS soon deeply decurrent, close, white to pale buff or tinged
pinkish-buff. STALK 3-10 (15) cm long(l) 1.5-3 (4) cm thick, equal or with an enlarged
base; solid, fairly firm, whitish to buff or becoming colored like cap, fibrillose-striate, base
usually with white down. SPORE PRINT white; spores 6-9 * 4-6 microns, elliptical,
smooth.
Clitocybe maxima is essentially an overgrown version of C. gibba (see color plate). Along with its
close relative C. geotropa, it is fairly common under conifers in the Rocky Mountains and Pacific
Northwest. Note deeply decurrent gills and depressed or vase-shaped cap.

HABITAT: Solitary or more often in groups or rings on ground in woods or clearings;

widely distributed and fairly common in the late summer and fall in the mountains of
western North America. I’ve see large fruitings near Mr. Rainier in Washington and in
the Sacramento M ountains of southern New Mexico, but I have not found it in California.
EDIBILITY: Said to be edible; I haven’t tried it.
COMMENTS: This large, handsome Clitocybe is easily told by its thick stem, decurrent
whitish gills, and pinkish-tan, broadly funnel-shaped cap. It is essentially an overgrown
C. gibba, and some consider it a variety of that species. C. geotropa is a very similar edible
species with more or less round spores. It tends to be longer-stemmed and more leather-
colored when young, and its cap often has a broad central umbo, even when funnel-
shaped. It is also widely distributed, but rare; I have seen it only in the southern Rockies.

Clitocybe gigantea (Giant Clitocybe)

CAP (5) 8-35 (45) cm broad, convex becoming plane and then broadly funnel-shaped;
surface smooth, not viscid; white becoming buff or pale tan, especially at the center; easily
broken when mature; margin at first inrolled, often obscurely ribbed in age. Flesh white,
in age becoming fragile; odor and taste pleasant to slightly disagreeable. GILLS close,
pallid soon becoming pale buff or creamy-buff, often pale dingy tan in old age; crowded,
decurrent, at least some forked. STALK 4-10 cm long, 2-4 (6) cm thick, often rather stout,
equal or enlarged slightly at either end; white, or in age pallid with darker fibrils. SPORE
PRINT white; spores 6-8 * 3-4.5 microns, elliptical, smooth, weakly amyloid.
HABITAT: Solitary to scattered or gregarious, often in arcs or huge rings, in open woods
and grassy (but usually wooded) areas; widely distributed, but most common in the Pacific
Northwest and Rocky Mountains. I have seen it and the very similar C. Candida (see com¬
ments) in the mountains of northern New Mexico and southern Arizona. Neither species
has been reported from California, but they may well occur.
EDIBILITY: Temptingly large and handsome, but poor in flavor and difficult to digest.
CLITOCYBE 159

COMMENTS: This mammoth mushroom is often so wide in proportion to its thickness

that the cap breaks or crumbles if handled carelessly. Its size and fragility, plus the buff-
colored decurrent gills, pale cap, and white spore print are diagnostic. Young, compact
specimens can be mistaken for C. irina, C. praemagna, or C. robusta, but have white
spores. Older individuals resemble Leucopaxillus, but are not nearly so tough, and lack
the white mycelial mat at the stem base. C. Candida is a very similar but more common
species with a slightly “smaller” cap (6-30 cm) that remains white until old age. Another
large species, C. septentrionalis, has a dull grayish to dingy tan cap, strong unpleasant odor,
and short thick stalk. I have seen it in groups or clumps under ponderosa pine in both
Oregon and New Mexico. Along with C. gigantea and C. Candida it is placed in the genus
Leucopaxillus by some mycologists because of its weakly amyloid spores. There are also
numerous large grayish Clitocybes with white, non-amyloid spores. The most common in
our area is C. harperi (see comments under C. nebularis).

Clitocybe dilatata
CAP 2-15 cm broad, convex to plane or often somewhat misshapen; surface dry, smooth,
gray when young, but soon white or chalky-white, sometimes with buff areas; margin at
first incurved, often wavy in age. Flesh white to grayish, firm; odor mild, taste typically
somewhat sour or disagreeable. GILLS adnate to decurrent, whitish to buff, close. ST ALK
5-12 cm long, 0.5-3 cm thick, equal or enlarged below, whitish, fibrillose, fibrous, often
curved. SPORE PRINT white; spores 4.5-6 x 3-3.5 microns, elliptical, smooth. Basidia
lacking siderophilous granules.
HABITAT: In densely-packed groups or clusters in sandy or gravelly soil along roads,
trails, etc.; very common in the fall in the Pacific Northwest and Y ukon. I have not seen it in
California, but it may occur in the northern part of the state, and Chuck Barrows reports it
from New Mexico.
EDIBILITY: Probably poisonous—it is thought to contain muscarine.
COMMENTS: The white spores and growth habit in clusters (often quite large) in dis¬
turbed ground is unusual for a Clitocybe and serves to distinguish this species from its
brethren. It is most likely to be confused with the Lyophyllum decastes group (espe¬
cially L. connatum), which also grows along roads, and with the C. subconnexa group,
which has pinkish spores. C. cerussata var. difformis is a synonym.

Clitocybe nebularis (Cloudy Clitocybe)

CAP 6-25 cm broad or more, convex becoming plane or depressed; surface dry, finely
fibrillose or often with a hoary bloom, gray to grayish-brown to buff, often darker at the
center, sometimes with watery spots or appearing streaked; margin at first incurved, often
wavy or lobed in age. Flesh thick, white; odor rancid and disagreeable. GILLS adnate to
decurrent, close, whitish becoming dingy yellowish or buff. ST ALK 6-15 cm long, 1.5-4 cm
thick, base often enlarged and covered with white down; whitish, or with dingy brownish
fibrils, firm but easily broken. SPORE PRINT pale buff to yellowish; spores 5.5-8.5 x 3.5-
4.5 microns, elliptical, smooth.

HABITAT: Scattered to gregarious, often in large rings, under both hardwoods and
conifers; widely distributed but especially common on the west coast. It is fond of cold
weather and in our area seldom appears before December.
EDIBILITY: Edible but far from incredible. It is indigestible unless thoroughly cooked,
its flavor is said to be poor, and its rank odor doesn‘t exactly make you want to rush home
and throw it in the frying pan.
Clitocybe nebularis is a common large cold-weather Clitocybe. Note decurrent gills and grayish cap
which is broadly convex at first but becomes depressed in age (the ones on left look darker than they
are because of shadow). Cap margin is often frilled or lobed and the skunklike odor can’t be missed.

COMMENTS: The outstanding feature of this large, drab, cold-weather Clitocybe is the
unpleasant odor, which has been likened to that rancid flour, rotting cucumbers, skunk
cabbage, mice cages, and beer barf. In size and stature it is reminiscent of Leucopaxillus
albissimus, but is grayer, decays much more rapidly, and has pale yellowish spores.
C. robusta(-C. alba) is a closely related species with the same stature, spore color, and
rancid odor. However, it has a pure white, sometimes lustrous cap. I have found it several
times in our area in mixed woods but it is more common in eastern North America. There
are also a number of large, undistinguished grayish Clitocy bes that have white spores and a
more or less mild odor. These include: C. harperi, with darker (grayish) gills, fairly
common in our area and throughout the West under conifers; and C. crassa, a springtime
Rocky Mountain species with a thick, massive stalk.

Clitocybe clavipes (Club-F oot)

CAP 2-10 cm broad, broadly convex or plane becoming depressed or funnel-shaped; sur¬
face more or less smooth, not viscid, brownish to grayish-brown or olive-brown, often
paler toward the margin. Flesh pallid, odor mild or often somewhat fragrant or fruity.
GILLS decurrent, fairly close, at first white, then yellowish-buff in age. STALK 2-7 cm
long, 0.5-1.2 cm thick at apex, typically club-shaped (enlarged below), pallid with grayish
or sordid olive-buff fibrils; base often spongy and covered with white down. SPORE
PRINT white; spores6-8.5 * 3-5 microns, elliptical, smooth.
HABITAT: Solitary, scattered, or gregarious under conifers and in mixed woods, late
fall and winter; widely distributed. It is particularly abundant in pine plantations in
eastern North America, rather infrequent in California (although several similar species
occur). I have not seen it in our area.
EDIBILITY: Edible, but not recommended. Not only is there the danger of confusing it
with poisonous species, but it has reportedly caused coprine-like poisoning when
consumed with alcohol (see p. 896).
COMMENTS: I must often strain to find something interesting to say about Clitocybes,
and this species—the type of the genus—is no exception. The drab grayish-brown cap,
decurrent gills, white spores, and club-shaped stem are characteristic, but there are many

160
Clitocybe clavipes has a swollen stem base, decurrent gills, and grayish-brown cap, but there are
dozens of similar, difficult-to-differentiate Clitocybes.

more or less similar species. One of these, C. avellaneialba, is fairly common in the Pacific
Northwest and northern California in humus or near decayed logs. It is somewhat larger,
with a darker cap, white gills and stem, and elongated spores (8-10 microns long). Another
western species, C. leopardina, is also similar but has a viscid, watery-spotted cap and
more or less equal stalk. It has been found by Greg Wright in southern California.

Clitocybe albirhiza (S nowmelt Clitocybe)

CAP 2-10 cm broad, convex becoming plane to broadly umbonate, then depressed; surface
smooth or with whitish down, or sometimes with riverlike lines; watery brown to pale buff
to cinnamon-, pinkish-, or grayish-buff. Flesh thin, colored like cap; odor unpleasant or
mild. GILLS adnate to decurrent, close, pale buff or colored more or less like cap. STALK
3-8 cm long, 0.4-2 cm thick, equal or tapered at either end, often hollow in age, sometimes
fluted; colored more or less like cap, the base with a dense mass of white mycelial threads.
SPORE PRINT white; spores 4.5-6 * 2.5-3.5 microns, elliptical, smooth, not amyloid.
HABITAT: Scattered to densely gregarious, often in small clusters or rings, on ground
under conifers; common in the mountains of western North America, especially in the
spring shortly after the snow melts (sometimes found with Lyophyllum montanum).
EDIBILITY: Unknown.
COMMENTS: This species is merely one of a metagrobolizing myriad of mundane white
to buffy-brown or grayish Clitocybes that are exceedingly difficult to distinguish. In other
words, they are better neglected than collected! It can be separated from its numerous look-
alikes, however, by the dense mass of white mycelial threads (rhizomorphs) at the base of
the stem and in the surrounding humus. Its growth in the spring is also distinctive. Other
smallish Clitocybes include: C. cerussata, with whitish cap; C. variabilis, with pure white
cap when fresh and spores 5.5-8 microns long, found under conifers; C. candicans, also
with a white cap, but favoring hardwoods; and C. coniferophila, one of several conifer-
loving species with a dingy buff cap. All of these lack the rhizomorphs characteristic of
C. albirhiza, and in California are more likely to fruit in the fall or winter than the spring.

Clitocybe odora (Blue-Green Anise Mushroom)

CAP 2.5-10 cm broad, convex to plane or somewhat depressed; surface smooth, not viscid,
bluish-green to dingy greenish, gray, or grayish-brown with a slight blue-green tinge, to
nearly whitish in dry weather. Flesh whitish or tinged cap color; odor strongly fragrant,
anise-like, at least when fresh. GILLS adnate to decurrent, close, blue-green to greenish or
dark green in oneform, whitish to buff or pinkish-buff in another. STALK 2-6 (9) cm long,
0.5-1.5 (3) cm thick, equal or thicker at either end; smooth, white to buff or cap-colored.
SPORE PRINT pinkish-cream or buff; spores 6-8 * 3-5 microns, elliptical, smooth.
Anise mushrooms. Left: Clitocybe deceptiva (formerly C. suaveolens), a young specimen. Right:
Mature specimens of Clitocybe deceptiva, a whitish to buff species, and Clitocybe odora, a partially
or completely greenish or bluish-green species (far right). Both smell like anise when fresh.

HABITAT: Scattered or in groups in woods; widely distributed. It favors oak in eastern

North America, but in the Rocky Mountains and Pacific Northwest it is often abundant
under conifers. In our area it is rare—I have seen it only twice, under oak in the winter.
EDIBILITY: Edible, but best used as a flavoring agent because of the strong taste.
COMMENTS: The blue-green to dull greenish color combined with the anise or licorice
odor immediately identify this mushroom. The form with blue-green tints in the gills and
stalk is var. pacificus, while the typical variety shows the color mainly in the cap. The other
anise-scented Clitocybes(C. deceptiva, etc.) are never blue-green. C. aeruginosa of eastern
North America has a greenish cap, but has white spores and does not smell like anise; a very
similar but unidentified greenish species with a distinctive (but not anise-like) odor occurs
in the Sierra Nevada under conifers. There are also several small (cap 1-2.5 cm broad, stalk
1-4 mm thick) greenish species that connect Clitocybe to Omphalina. These include: C.
atroviridis (-Omphalina chlorocyaneal), with a green to blackish-green to olive-gray cap
and paler gills, found in grass, moss, or on lichens; Omphalina grossula, greenish-yellow
to olive-yellow, favoring lichens and moss; and O. wynniae, with a greenish-yellow to
olive-brown cap and yellow to greenish-yellow gills, usually found on rotting conifers.

Clitocybe deceptiva (Anise Mushroom)

CAP 1-6 cm broad, broadly convex becoming plane or depressed; surface smooth, not
viscid, color variable: pale brown to grayish-brown fading to buff or watery whitish,
sometimes with a darker marginal band; margin at first incurved. Flesh thin, pallid, odor
distinctly fragrant and anise-like when fresh. GILLS adnate to slightly decurrent, close,
white or cap-colored or slightly darker. ST ALK 2-7 cm long, 1.5-5 (7) mm thick, equal or
thicker at base, smooth, more or less colored like cap, usually rather slender. SPORE
PRINT pale pinkish-buff; spores 6.5-9 * 4^4.5 microns, elliptical, smooth.
HABITAT: Scattered to gregarious in damp places under conifers; western North
America. In our area this species and its close relatives(see comments) are often common in
the fall, winter, and early spring under redwood and pine, or occasionally oak.
EDIBILITY: Edible, but best used as a flavoring agent (in cakes, breads, cookies, etc.),
because of its strong, anise-like (but not sweet) flavor. Be sure each and every specimen
smells like anise—there are numerous look-alikes without the anise odor, some of
which grow in the same habitats and are poisonous!
COMMENTS: Better known as C. suaveolens, the distinct anise odor of this species and
its close relatives separate them from the dozens of other small, nondescript white to
grayish or buff-colored Clitocybes (see C. albirhiza). Similar anise-smelling species in¬
clude: C.fragrans, practically identical, but with a paler cap and white spores, and growing

162
CLITOCYBE 163

under hardwoods as well as conifers; C. oramophila, with a dull pinkish or flesh-colored

cap when moist and pale creamy spores; and C. obsoleta, with a strongly fragrant but not
aniselike odor.

Clitocybe dealbata (Sweat-Producing Clitocybe)

CAP 1-4 (5) cm broad, convex becoming plane or depressed; surface smooth, not viscid,
white to dingy white, grayish, or buff, sometimes with a pinkish tinge in wet weather,
sometimes with watery spots. Flesh thin, grayish to white; odor mild. GILLS grayish-white
to buff or pinkish-buff, close, adnate to decurrent. STALK 1-4 cm long, 2-7 mm thick,
equal or slightly thicker at either end, colored like cap, smooth, rather tough. SPORE
PRINT white or rarely creamy; spores 4-5.5 * 2 A microns, elliptical, smooth.
HABITAT: Scattered to gregarious or in rings, in pastures and other grassy places; widely
distributed and common locally in the fall and winter. In the Pacific Northwest it fre¬
quently mixes company with Marasmius oreades, but in our area is more often found
with Agaricus campestris and A. cupreobrunneus.
EDIBILITY: Poisonous—and potentially fatal to small children in the “grazing” stage! It
produces profuse sweating, salivation, diarrhea, etc. Muscarine is the main toxin (see
p. 894 for details).
COMMENTS: The dingy grayish-white color, close decurrent gills, white spores, small
size, and growth in grass distinguish this drab, undistinguished mushroom, also known
as C. sudorifica. In our area it frequents pastures while the edible and somewhat similar
fairy ring mushroom (Marasmius oreades) grows mainly on lawns; in other regions, how¬
ever, they may grow together. H owever, the fairy ring mushroom has a browner, frequently
umbonate cap and well-spaced gills that are never decurrent. The edible Clitopilus
prunulus is also similar, but has pinkish spores. There are numerous woodland look-alikes
in the genus Clitocybe, and all should be strictly avoided. Closely related, poisonous
grass-inhabiting species include: C. augeana, very similar but with a farinaceous odor,
sometimes rivulose in age (developing riverlike lines or cracks on the cap), and growing in
manure and compost as well as in grass; C. rivulosa, with cap often rivulose and tinged
flesh-color, growing in grass; and C. morbifera, with a grayish-brown cap, in grass.

Clitocybe dealbata is a small poisonous grass-inhabiting mushroom with closely spaced, adnate to
decurrent gills. Overall color is dingy gray to buff, leading to confusion with the edible fairy ring
mushroom (Marasmius oreades).
Clitocybe sclerotoidea. Note growth habit in small, tight clumps. Each clump arises from a mass of
tissue thought to be aborted Helvetia lacunosa.

Clitocybe sclerotoidea (Parasitic Clitocybe)

CAP 0.5-4 (5) cm broad, convex to plane or slightly depressed centrally; surface dry,
unpolished, with a fine whitish fibrillose coating which rubs off; pallid to sordid buff to
brownish or grayish, sometimes with darker watery spots. Flesh whitish, odormild. GILLS
adnate or slightly notched becoming decurrent, pale buff or pinkish-buff darkening to
gray, olive-gray, or grayish-brown. ST ALK 2-4 (8) cm long, 3-10 mm thick, equal or thicker
at either end, solid, colored like cap or paler from soft, matted, downy white hairs; arising
from a fleshy mass of tissue which is often at least partially buried. SPORE PRINT
white; spores 8-11 * 3-4 microns, subfusiform (elongated), smooth.
HABITAT: Typically in small, tight clumps on ground under pine, known only from the
Pacific Coast; common at times in our area in the winter and spring. The fluted black
elfin saddle (Helvetia lacunosa) is often found nearby.
EDIBILITY: Unknown, and like myself, likely to remain so.
COMMENTS: I can find hidden virtues in almost any fungus, but this drab Clitocybe
defies me—its mediocrity is downright stupefying. It serves as a compelling reminder that
organisms do not exist for our enjoyment alone, and should not be judged accordingly. It
can be recognized by its small size, dingy color (or lack of color), and habit of growing in
compact clumps from a fleshy mass of tissue. S mall, aborted individuals are usually present
in each clump, and occasionally large, solitary specimens are encountered. The tissue mass
from which they arise (“sclerotium”) is actually composed of hyphae from both C.
sclerotoidea and Helvetia lacunosa—suggesting that the Clitocybe is parasitic on the
Helvetia. It is interesting to note, however, that the Helvella occurs with both hardwoods
and conifers, while C. sclerotoidea is confined to pine, at least in my experience.

Clitocybe cyathiformis
CAP 2.5 -8 cm broad, centrally depressed with an inrolled margin, becoming funnel-
shaped in age; surface smooth, not viscid, dark brown to dark gray-brown, but fading in
age to grayish or paler brown. Flesh thin, pallid; odor mild. GILLS at first adnate but soon
deeply decurrent, pallid becoming grayish or grayish-brown, close. STALK 5-12 cm long,
0.4-1 cm thick, equal or thicker below, often rather long, colored like cap or paler, fibrillose,
often with whitish down at base. SPORE PRINT white; spores 7-11 * 5-6 microns,
elliptical, smooth, amyloid.

164
CLITOCYBE 165

HABITAT: Solitary or in small groups in humus or on rotten logs, in woods and at their
edges; widely distributed. Not uncommon in our area in the fall and winter, especially
under redwood.
EDIBILITY: Not recommended. It is said to be edible, but there are many very similar
species of unknown edibility.
COMMENTS: This species is one of numerous small to medium-sized, grayish, white-
spored agarics with decurrent gills and a depressed to funnel-shaped cap. Because of its
amyloid spores it has been placed in several different genera, including Cantharellula and
Pseudoclitocybe. Similar species with amyloid spores include: Cantharellula umbonata,
common in moss beds in northern and eastern North America, with crowded, narrow,
whitish, forked gills that stain reddish in age; and Clitocybula atrialba, cap 2-10 cm broad
and blackish-brown, gills well-spaced and white to grayish, stalk clothed with dark scurfy
scales, growing in rich humus or on rotting hardwoods in the Pacific Northwest. The first of
these were formerly placed in Cantharellus, the second in Clitocybe. Also see Myx-
omphalia maura and Clitocybe albirhiza.

Myxomphalia maura
CAP 1-3.5 (5) cm broad, convex or centrally depressed with an incurved margin, be¬
coming plane or centrally depressed; surface viscid when moist but soon dry and often
shiny, smooth, dark grayish-brown or olive-brown to blackish-brown, fading to gray or
paler as it dries. Flesh thin, white to grayish. GILLS adnate to slightly decurrent, close,
white to pale grayish (usually paler than cap). ST ALK 2-6 cm long, 2-5 (6) mm thick, more
or less equal, smooth, colored more or less like cap or slightly paler, but not fading as
quickly. SPORE PRINT white; spores 4.5-6.5* 3.5-4.5 microns, broadly elliptical
to nearly round, smooth or very minutely ornamented, amyloid.
HABITAT: Solitary, scattered, or in groups on burnt soil and debris, especially under
conifers; widely distributed and fairly common in our area in the appropriate habitat, fall
through spring. I often find it with Pholiota brunnescens and Psathyrella carbonicola.
EDIBILITY: Unknown, but too puny to be of value.

Left: Clitocybe cyathiformis, mature specimens. Right: Myxomphalia maura, a small mushroom
that favors recently burned areas. Note the adnate to decurrent gills.
166 TRICHOLOMATACEAE

COMMENTS: The habitat on burnt soil distinguishes this undistinguished mushroom

from a host of similar grayish-brown look-alikes. Lyophyllum atratum(-Collybia atrata)
is another species that grows in burned-over areas; it is quite similar but has darker (gray)
gills and non-amyloid spores (the two are sometimes found growing together). Fayodia
anthracobia is a minute charcoal-lover (cap up to 1 cm broad) with round, warted, amyloid
spores. Omphaliaster asterosporus and O. borealis have warted or starlike spores, but
are larger (cap 2-5 cm broad), have grayish to grayish-brown gills, and prefer moss (under
conifers) to burnt soil. Another small widespread species, Clitocybe (-Omphalina)
epichysium, has a dry cap about 2 cm broad that is dark sooty-brown to olive-brown or
ashy and minutely scaly or scurfy at the center. It is fairly common in our area in mixed
woods and under conifers, but not in burnt soil. See also C. cyathiformis.

LEUCOPAXILLUS

Medium-sized to large terrestrial mushrooms. CAP convex to plane or depressed, margin usually
inrolled when young; surface dry, unpolished. Flesh rather tough and dry. GILLS attached, close,
usually white or yellowish. STALK typically central, fleshy, tough, often with a conspicuous white
mycelial mat at base. VEIL and VOLVA absent. SPORE PRINT white. Spores amyloid, rough.

THIS is a small but very conspicuous group of robust mushrooms with a dry, unpolished
cap, white to pale yellow gills, a tough, fleshy stem, and white, amyloid spores. In some
species the gills peel rather easily from the cap (as in the brown-spored genus Paxillus). The
flesh is rather dry and brittle as in Russula, but the stature of the fruiting body is quite
different and the fibrous stalk does not snap open cleanly like chalk. Also, most Leuco-
paxillus species have a copious white mycelium that permeates the surrounding duff and
frequently adheres to the base of the mushroom when it is plucked—a character not found
in Russula.
Confusion with Clitocybe and Tricholoma is also likely (Leucopaxillus species were
originally placed in those genera), but they are not so tough and rot more readily. In
contrast, Leucopaxillus species are remarkably resistant to bacterial decay—antibiotic
substances have been isolated in some of them and their fruiting bodies, like those of
Laccaria, persist for weeks and thus appear to be more common than they actually are.

Leucopaxillus albissimus, mature and dried-up specimens. Note the tremendous variation in size,
shape, and stature. Some of these are quite old and shrivelled.
Leucopaxillus albissimus. Young white specimens. (Ralph Buchsbaum)

Less than ten species of Leucopaxillus are known from North America. The two
common ones in our area fruit prolifically and are likely to be among the first agarics you
encounter. They are strictly woodland fungi and may be mycorrhizal. Though alluringly
large and firm, they are difficult to digest because of their toughness. However, they are
not known to be poisonous, so intrepid toadstool-testers may opt to experiment with
the mild-tasting forms. They have one compelling advantage—a single large specimen
can fill a basket!

Key to Leucopaxillus
1. Gills pale clear yellow when young; cap 9-30 cm broad, buff to dingy tan or yellow-brown;found
under hardwoods in eastern North America and southern Arizona . L. tricolor
1. Not as above; gills not yellow when young (but may be dingy yellowish in old age) . 2
2. Cap brown to reddish-brown, medium-sized; taste bitter. L. amarus, p. 168
2. Cap white, buff, pale tan, yellowish, etc; medium-sized to very large; taste mild or bitter ... .
.L. albissimus & others, below

Leucopaxillus albissimus (Large White Leucopaxillus)

CAP 5-20 (40) cm broad or more, convex with an inrolled margin when young, becoming
plane or depressed in age; surface dry, dull, unpolished, white becoming buff, yellowish,
or even tan (at least toward center) in age, often cracked or splitting in age or dry weather;
margin sometimes obscurely ribbed. Flesh thick, tough, white; odor unpleasant or frag¬
rant, taste also variable (mild or bitter). GILLS white to slightly yellowish in old age,
attached (typically decurrent at least by lines, but ranging to adnate or even adnexed);
close. STALK 5-20 cm long, 1-5 cm thick, tough, solid, equal or enlarged at the middle or
below (often with a narrowed base); white, discoloring slightly in age; base usually im¬
bedded in a white mycelial mat; smooth or often scaly (especially in mid-portion). SPORE
PRINT white; spores 5.5-8.5 * 4-6 microns, elliptical, warty, amyloid.
HABITAT: Solitary, scattered, or gregarious (often in large rings) in woods, mainly
under conifers; widely distributed. It is common in our area in the cool winter months,
particularly under redwood, but also with other trees, including eucalyptus. It is slow to
grow and even slower to decay.
168 TRICHOLOMATACEAE

EDIBILITY: A tempting specimen, but I don’t recommend it—even the non-bitter forms
are coarse and difficult to digest. Thorough cooking would be necessary.
COMMENTS: Several varieties of this widespread species have been described based on
differences in cap color, taste, and spore characters. The above description is based
primarily on the variety common in our area, var. paradoxus (-L. paradoxus). It has a
rather strong but not unpleasant odor and a distinctive but not usually bitter taste. It is
white when young, but usually becomes dingy yellowish, buff, or pale tan as it ages or
dries out. A white mycelial mat can usually be seen permeating the duff around the mush¬
room or adhering to its base. However, its most outstanding attributes are the large size,
tenacious toughness, and resolute resistance to decay, as evidenced by the following two
“tried and true” methods for distinguishing it from Clitocybe nebularis and other large
look-alikes:
1) Choose a firm (not waterlogged) specimen and throw it against a wall, housemate’s
head, or other dense, hard, thick object. If it (the mushroom) remains more or less intact,
it is probably L. albissimus; if it shatters, chances are it was a Clitocybe.
2) Choose a firm (not waterlogged) specimen and leave it out on the porch, or hide it
in your housemate’s closet. If after one week it (the mushroom) shows no visible signs of
decay, it is probably L. albissimus; if unchanged after one month it is definitely L. albis¬
simus. If, on the other hand, it shows visible or smellable signs of decay (i.e., has begun
to rot or is reduced to a heap of writhing maggots), chances are, once again, that it was a
Clitocybe. (Note: L. albissimus can behave like a Clitocybe if too old or maggot-riddled.)
Other species: L. albissimus var. lentus is quite similar to var .paradoxus, but is usually
smaller (stalk only 0.8-1.5 cm thick and smooth); var. piceinus is similar in color, but has
a bitter taste; var. typicus also has a bitter taste but is pure white; L. laterarius is a hard¬
wood-loving species with a very bitter taste, pinkish-buff-tinged cap, and round spores; it
is particularly common in eastern North America.

Leucopaxillus amarus (Bitter Brown Leucopaxillus) Color Plate31

CAP 5-12 (15) cm broad, broadly convex with an inrolled margin when young, becoming
plane or slightly depressed; surface dry, unpolished, smooth, dark brown to brown, pecan-
brown, or reddish-brown, evenly colored or paler at the margin, sometimes cracked or
faded in age; margin often obscurely ribbed. Flesh thick, firm, dry, white; odor mild or
pungent; taste very bitter. GILLS typically adnate but ranging from notched to slightly
decurrent by lines; close, white. STALK 4-10 cm long, 0.5-2 (4) cm thick, equal or en¬
larged below, smooth, dry, solid, white or sometimes discolored brownish below; base
imbedded in a copious white mycelial mat. SPORE PRINT white; spores 4-6 x 3-5 microns,
nearly round, warted, amyloid. Cystidia numerous on gill edges.
HABITAT: Scattered to gregarious (often in rings) under conifers and oaks; common and
widely distributed. In our area it is often abundant in the fall and winter.
EDIBILITY: Unequivocally inedible—it smells like “creepy crawlers” and tastes like a
mildewed army tent. If you’re lost in the woods and have nothing to eat, you’d do better
to follow the example of Charlie Chaplin and stew your boots before venturing to make a
meal of this mushroom.
COMMENTS: Also known as L. gentianeus, this mundane mushroom can usually
be recognized by its dull, boring brown to reddish-brown cap, white gills, bitter taste,
absence of a veil, and moldy-looking white mycelium that usually permeates the sur¬
rounding humus and frequently adheres to the stem when it is plucked. The latter feature
helps to distinguish it from brown-capped Tricholomas (e.g., T. imbricatum). Hygro-
phorus roseibrunneus can also be similar in color, but has soft and waxy gills. As with L.
albissimus, several varieties and forms have been described. In addition to the color plate,
L. amarus is shown on p. 918.
 169

MELANOLEUCA
Small to medium-sized terrestrial mushrooms. CAP usually smooth, broadly convex to plane or
often umbonate; often hygrophanous. GILLS close or crowded, attached (usually adnate or
notched), usually white. STALK central, typically rather stiff, straight, semi-cartilaginous; often
slender. VEIL and VOLVA absent. SPORE PRINT white or creamy. Spores minutely warted or
roughened, amyloid. Cystidia often present on gill edges.

THESE rather unimposing whitish to gray or brownish mushrooms have been aptly
characterized as “overgrown Colly bias,” for the stature of the fruiting body is intermediate
between that of Tricholoma and Collybia. The straight, semi-cartilaginous stem, close to
crowded (and often notched) gills, and smooth, dull-colored, frequently umbonate cap
are the principal fieldmarks.
Melanoleucas can grow almost anywhere, but in our area are found most often on lawns
or shaded areas in pastures and parks. They are also quite common under conifers in the
Sierra Nevada and other mountain ranges. I can find no mention of poisonings attributed
to Melanoleuca, but the North American species are not well known. Two widespread
species are described here, and several others are keyed out.

Key to Melanoleuca
1. Gills tan to ochre, creamy-ochre, or pale pinkish-cinnamon, at least in age; cap 5-13 cm broad;
stalk usually quite tall .M. cognata, p. 170
1. Not as above; gills typically whitish, sometimes with a slight yellow or pinkish tinge .2
2. Cap viscid when moist, 4-8 cm broad, white or in age often tinged or stained yellow; stalk white,
usually scurfy, 0.7-1.5 cm thick; fairly common in central and southern California in a variety
of habitats .M. lewisii
2. Not as above . 3
3. Cap yellowish-brown fading to yellowish-white; fairly common under hardwoods in eastern
North America .M. alboflavida
3. Not as above; cap some shade of brown or gray when moist (but may fade in age) .4
4. Growing under mountain conifers, usually shortly after the snow melts . 5
4. Growing in lawns, pastures, etc., sometimes also in woods, but not typically as above.
.M. melaleuca group, below
5. Stalk fleshy, usually more than 1 cm thick; cap medium-sized to fairly large.
.M. evenosa group, p. 171
5. Stalk usually less than 1 cm thick; cap medium-sized to fairly small .
.M. graminicola & others (see M. melaleuca group, below)

Melanoleuca melaleuca group

CAP 2-7 (10) cm broad, broadly convex to plane to shallowly depressed, often with a low
broad umbo; surface smooth, hygrophanous but not viscid; dark brown to gray to grayish-
brown when moist, often fading to buffy-tan, gray, or even paler in sunlight(or as it dries).
Flesh thin, whitish, odor mild. GILLS narrow, close or crowded, white, usually notched
but varying to adnate or even very slightly decurrent. STALK 2-8 cm long, 3-6(12) mm
thick, equal or with a slightly swollen base, rather slender, stiff, whitish or with darker
(brown) fibrils, apex sometimes minutely scaly or scurfy. SPORE PRINT white; spores
6-8 x 4-5.5 microns, elliptical, minutely warted, amyloid. Harpoonlike cystidia typically
present on gill edges.
HABITAT: Scattered to gregarious on ground in open places(lawns, pastures, etc.), under
trees, along roads and trails, in straw and wood chips, and also in the woods; common and
widely distributed, but usually fruiting at lower elevations. Common in our area
throughout the mushrom season but most numerous in the winter and early spring.
Melanoleuca melaleuca group is common in grassy areas, but also grows under trees and in the woods.
Cap is usually dark when young and moist (see specimen on right), but may fade considerably in age
or sunlight. Note crowded white gills.

EDIBILITY: Edible. The caps are delicious fried in butter, but since it is not an easy
mushroom to recognize, I hesitate to recommend it. Also, the edibility of closely related
species has not been adequately ascertained.
COMMENTS: The above description will actually fit a number of closely related
Melanoleucas whose exact identities are best left to Melanoleuca-masters. As a group they
are characterized by a smooth, hygrophanous, dark brown to grayish (or paler) cap, close
white gills, straight stalk, and strongly amyloid, warted spores. Specimens growing in
the open are generally much paler than their counterparts growing in the shade. Related
species include: M. brevipes, common on lawns, with a short brown to whitish stalk and
abundant cystidia on the gills; and M. polioleuca, with a hoary bloom on the cap when
fresh. There are also several species that fruit prolifically in the spring and early summer
in the coniferous forests and alpine meadows of western and northern North America. The
most common of these, M. graminicola, is similar to M. melaleuca in color and stature,
but lacks cystidia on the gills. Another, M. evenosa, is larger and more robust (see descrip¬
tion). Although probably harmless, none of these species should be eaten until better
known. M. vulgaris is a synonym for the tongue-twisting M. melaleuca.

Melanoleuca cognata
CAP (5) 7-13 cm broad, broadly convex to plane, usually with a broad umbo, sometimes
becoming slightly depressed in age; surface smooth and sometimes shiny, dry or slightly
viscid, brown to ochre-brown, fading to pale tan in age, the center sometimes darker. Flesh
whitish; odor often slightly sweet, rancid, or “peculiar” (Smith). GILLS pallid becoming
tan, creamy-ochre, deep ochre, or pale pinkish-cinnamon; crowded, attached (usually
notched). STALK 6-12 cm long, 1-2 cm thick, equal or with a swollen base, longitudinally
lined or twisted-striate, straight, colored more or less like cap or paler, the base some¬
times brownish-stained. SPORE PRINT creamy or yellowish; spores7-10 * 4.5-6 microns,
elliptical, minutely warted, amyloid. Cystidia abundant on edges and faces of gills.
HABITAT: Solitary, scattered, or in small groups on ground in mixed woods and under
conifers in the spring, summer, and early fall; widely distributed but not common. It does
not occur in our area but is to be looked for in the Sierra Nevada. It is fairly frequent in the
spruce-fir-aspen forests of the southern Rocky Mountains and Southwest.
EDIBILITY: Edible. I haven’t tried it.
COMMENTS: In contrast to many of its kin, this Melanoleuca is fairly easy to recognize.
The broad, frequently umbonate brown to ochre-tan cap, tall straight stem, and brown to
tan or ochre mature gills are good fieldmarks.

170
MELANOLEUCA 171

Melanoleuca evenosa group (Robust Melanoleuca)

CAP 5-19 cm broad, broadly convex to plane or wavy; surface smooth, dry to slightly
viscid when moist, pallid to brown to grayish-brown or in age sometimes slightly ochre.
Flesh firm, thick, white; odor often rather pleasant. GILLS crowded, white (or in age
sometimes brownish- or ochre-stained); usually notched or adnexed. STALK 3-7 cm long,
(1) 1.5-3 (3.5) cm thick, equal or slightly thicker below, very firm, stiff, solid; white with
brownish or cap-colored fibrils; apex or upper portion usually dandruffy or minutely
scaly (or sometimes dandruffy throughout). SPORE PRINT white; spores 8-11 * 4-5
microns, elliptical, minutely warted, amyloid.
HABITAT: Solitary, widely scattered, gregarious, or even clustered in duff under conifers
or in grassy clearings, at the edges of forests, etc.; fairly common in the mountains of
western North America during the spring and early summer.
EDIBILITY: Unknown. Several Melanoleucas are edible and this one is fleshy enough
to warrant cautious experimentation, but be sure of your identification!
COMMENTS: Anyone who regularly hunts the Cascades and Sierra Nevada during the
springtime will come across this species or species “complex,” which also goes under the
name M. subalpina. Because of its robust stature (see photo on p. 897) it is likely to be
mistaken for a Tricholoma. However, the crowded white gills, scurfy stalk, smooth
cap, amyloid spores, and springtime growth under conifers should distinguish it from
most Tricholomas. Several other Melanoleucas grow under mountain conifers, but
are more slender (see comments under the M. melaleuca group). M. cognata can be
fairly large, but is usually taller and slimmer with tan to ochre gills in age.

LACCARIA
Small to medium-sized terrestrial mushrooms. CAP convex to plane, centrally depressed, or
uplifted; not viscid. GILLS attached, rather thick, slightly waxy, pinkish to flesh-colored, cinna¬
mon, purple, or lilac. STALK tough and fibrous, elastic, often slender, more or less central. VEIL
and VOLVA absent. SPORE PRINT white to pale lilac. Spores usually spiny, not amyloid.

THIS small but common genus can be distinguished in the field by its thick, purple to
pinkish or flesh-colored gills and tough, fibrous stem. The gills may be somewhat waxy-
looking as in the waxy caps (Hygrophoraceae), but the tough stalk is distinctive and the
spores are usually spiny under the microscope. I n small individuals the stalk may be slender
but is not noticeably fragile, and the cap is not conical or bell-shaped as in Mycena.
Laccarias form mycorrhizae with many kinds of trees and shrubs, and are among the
few mycorrhizal species that are easily raised to fruition in the laboratory. Their omni¬
presence in coniferous forests has caused them to be called “mushroom weeds.” However,
they also grow with hardwoods. They seem particularly fond of sandy, boggy, or other
poor soils, and are a conspicuous fungal feature of our coastal pine forests. They persist
for weeks without decaying, and thus appear to be more plentiful than they actually are.
About 16 species of Laccaria occur in North America. M ost are quite variable in shape
and color and thus difficult to distinguish in the field, but they split nicely into two groups
—the ones that are purple and the ones that aren’t. All are thought to be edible and most
are fairly good, or at least better than some authorities would have us believe. Two repre¬
sentative species are described here.

Key to Laccaria
1. Downy mycelium at base of stalk and/ or gills purple to lilac when fresh (may fade in age!) 2
1. Violet tones completely absent, even when fresh (but may be vinaceous-tinged) .
.L. laccata & others, p. 172
172 TRICHOLOMATACEAE

2. Growing in sand (often buried); spores smooth L. trullisata(see L. amethystina group, below)
2. Not as above (but may grow in sandy soil); spores spiny .3
3. Fruiting body medium-sized to large (cap 5-20 cm broad; stalk 1-3 cm thick); common under
hardwoods (especially oaks) in eastern North America .L. ochropurpurea
3. Not as above; small to medium-sized (cap usually less than 7 cm broad; stalk usually less than
1 cm thick, but at times up to 1.5 cm); widespread in many habitats .4
4. Gills distinctly purple when fresh .L. amethystina group, below
4. Gills with only a tinge of violet (if any) when fresh L. bicolor (see L. amethystina group, below)

Laccaria laccata (Lackluster Laccaria) Color Plate 29

CAP 1.5-6 cm broad, convex becoming plane to centrally depressed or sometimes even
with a hole in the center, the margin often uplifted in age; surface not viscid, often minutely
scaly; color variable: flesh-colored to orangish, brownish-cinnamon, reddish-tan,
or pinkish-brown when moist, much paler as it dries; margin often wavy or irregularly
lobed in age. Flesh thin, tinged cap color, odor mild or sometimes radishlike. GILLS thick,
well-spaced, somewhat waxy, pale pinkish to flesh-colored or reddish-tan, dusted white by
spores at maturity; attachment variable but typically adnate to slightly decurrent. ST ALK
2-10 cm long, 3-10 mm thick, more or less equal, tough and fibrous, elastic, often fibrillose,
often somewhat twisted orcompressed; same coloras moist cap or darker(reddish-brown),
usually rather slender; downy mycelium at base white (if present). SPORE PRINT white;
spores 7-10 * 6-9 microns, round or nearly round, spiny.
HABITAT: Scattered or in groups or troops in woods or near trees, especially in poor or
sandy soil or in boggy areas; very common and widely distributed. In our area it can be
found almost anywhere at anytime, but favors cool weather and pine. L. proximo and
L. altaica(see comments) are also common locally.
EDIBILITY: Edible and fairly good, especially if seasoned; the tough stems should be
discarded. Be certain of your identification, however—there are many similarly-colored
mushrooms!
COMMENTS: This cosmopolitan mushroom has a knack of turning up in droves when
one is seeking more exotic or unusual species, and is thus spurned and scorned as a
mushroom “weed.” To be sure, it is as unostentatious as it is ubiquitous, without the lovely
color of the L. amethystina group, yet in its own unexciting way it is quite beautiful. Like
the honey mushroom, it is so vexingly variable in size, color, and shape that even veteran
collectors have trouble recognizing some forms! The telltale traits are the overall color
(pinkish to orangish or dull cinnamon), thick well-spaced gills, white spores, and tough
or fibrous stem. L. proximo is a very similar, widespread species. It tends to be slightly
larger with a more fibrillose stalk and scalier cap, but can only be told with certainty by
its broadly elliptical (rather than round) spores. There are also several similarly-colored
but smaller species (cap 0.5-4 cm broad) that usually grow in wet or boggy areas and have
striate caps when moist. They include: L. ohiensis andL. altaica( both of which have been
called L. striatula), with larger spores (8-13 microns), the latter especially common in
northern latitudes; andL. lortilis, a very small species with only a few( 10-15) gills and even
larger spores. See also L. bicolor(\mder L. amethystina group), which has violet mycelium.

Laccaria amethystina group (Amethyst Laccaria) Color Plate 30

CAP (1) 2-4 (7) cm broad, convex becoming plane, centrally depressed (sometimes with a
hole in the middle), or with an uplifted margin in age; surface often minutely scurfy or scaly,
not viscid; purple to brownish-purple when fresh and moist, fadingto brown, gray, buff, or
whitish (see comments!) as it loses moisture; often cracked in dry weather; margin often
wavy or lobed in age. Flesh thin, violet-tinged; odor mild. GILLS well-spaced, thick, some-
LACCARIA 173

what waxy, deep or bright amethyst-purple when fresh, gradually fading to dull purple or
grayish-purple and eventually dusted white by spores; attachment variable but usually
adnate to slightly decurrent. STALK 5-12 cm long, 0.3-1 (1.5) cm thick, more or less equal,
typically rather long and slender, often curved or twisted; tough and fibrous, elastic,
usually conspicuously fibrillose, colored more or less like moist cap or browner, fading as
it dries; base usually covered with downy purple or white mycelium. SPORE PRINT
white or tinged lilac; spores 7-11 microns, round, spiny (but see comments!).
HABITAT: Scattered to densely gregarious onground in forests and at their edges; wide¬
spread and common. In our area this species and its look-alikes (see comments) are par¬
ticularly abundant under coastal pines. They can fruit most any time but are partial to cold
weather and are often abundant when other collectable delectables are scarce.
EDIBILITY: Edible, and a good choice for beginners because of its distinctive color and
convenient availability. It has a nice texture but not much flavor, so try mixing it with
potatoes and seasoning it with garlic. The tough, hairy stems should be discarded.
COMMENTS: The lovely amethyst-purple gills when fresh combine with the rather long
and tough, fibrous-hairy stem to set apart this cosmopolitan mushroom and its close rela¬
tives. The color fades rather dramatically as the mushroom loses moisture, but the gills
usually retain their purple tint well into maturity. The common11 Amethyst Laccaria” along
the west coast has recently been given a new name,L. amethysteo-occidentalis, based on
its broadly elliptical (rather than round) spores and the tendency of its cap to fade to brown
rather than buff or gray as in the “true” L. amethystina (or L. amethystea). Our local
variety, however, sometimes fades to buff or even white, so that microscopic examination
is often necessary to distinguish it from L. amethystina (the latter may occur in our area,
but if so is uncommon). The specimens in the color plate are probably L. amethysteo-
occidentalis, but are labeled L. amethystina group because their spores were not examined.
To muddle matterseven more,L. bicolor is also common in our area, especially under pine.
It has broadly elliptical spores, but has only a slight violet or vinaceous tinge (if any)
when fresh, except for the violet downy mycelium at the base of the stalk (which may,
however, fade to white in age, leading to confusion with L. laccata). Other purple-gilled
species: L. trullisata has long( 16-22 microns) smooth spores and grows only in sand or sand
dunes (often buried!). L. ochropurpurea is a robust eastern species (see key to Laccaria).

LYOPHYLLUM & Allies

Small to medium-small mushrooms, or if larger then usually growing in dense clusters. CAP
typically some shade of gray or brown, but sometimes white; not viscid. GILLS attached or
sometimes free, usually white or gray. STALK fleshy or thin, central, usually white, gray, or brown.
VEIL and VOLVA absent. SPORE PRINT white. Spores smooth or spiny, not amyloid. Cystidia
on gills typically inconspicuous or absent. Basidia with siderophilous (carminophilous) granules.

LYOPHYLLUM is a nondescript amalgamation of white-spored mushrooms at one time

dispersed among Tricholoma, Collybia, and Clitocybe. They are puzzling to the amateur
because the various species appear to have little in common aside from their drab color—
some are slender and Collybia- or Mycena-like; others are robust and Tricholoma- or
Clitocybe-like. The unifying feature is rather esoteric: the basidia contain particles which
darken dramatically when heated in acetocarmine! (Wouldn’t you?)
Lyophyllum is a fairly sizable genus but only the L. decastes group, which fruits in large,
dense clusters along roads and trails, is conspicuous. The rest qualify as“LBM’s” and are
treated here only perfunctorily (they are differentiated largely on microscopic
characteristics anyway). Also included here is one species of Calocybe—a small, rare genus
which, like Lyophyllum, has siderophilous granules in the basidia, but typically has a more
brightly colored cap.
174 TRICHOLOMATACEAE

Key to Lyophyllum & Allies

l. Cap pinkish-tan to reddish (sometimes fading to tan); stalk usually 5 mm thick or less .
. Calocybe carnea & others, p. 176
1. Not as above; cap grayish to brown, black, or white . 2
2. Gills and/or stalk staining gray or black where bruised (sometimes slowly), or developing
grayish to black spots in age.:. L. semitale & others (see L. montanum, p. 175)
2. Not as above . 3
3. Growing on burnt ground or debris; stalk thin (up to 3 mm); cap small, dark brownish to black
.L. atratum (see Myxomphalia maura, p. 165)
3. Not growing on burnt ground and/ or stalk thicker.4
4. Stalk fairly thick (1-2.5 cm thick), usually white; typically growing in dense clusters (rarely
solitary) along roads and paths or less commonly in undisturbed woods . 5
4. N ot as above; stalk usually slender . 6
5. Caps white, often somewhat misshapen; taste usually sour or disagreeable; known only from
western North America .(see Clitocybe dilatata, p. 159)
5. Not as above; caps not white, or if white then taste more or less mild L. decastes group, below
6. Fruiting in spring in western mountains, usually near melting snow; fruiting body grayish
when fresh; gills gray; cap usually with a hoary bloom at first .. . L. montanum, p. 175
6. Not as above . 7
7. Found in sphagnum bogs; cap usually with an umbo; stalk thin, fragile, often long L palustre
7. Similar to above but not growing in bogs .L. rancidum & others

Lyophyllum decastes group (Fried Chicken Mushroom)

CAP 3-12 cm broad, convex to plane or with slightly uplifted margin in age; surface
smooth, often with a soapy (lubricous) feel when moist, but not viscid; color variable: dark
brown to grayish-brown to yellowish-brown, watery tan, or paler; margin often lobed.
Flesh firm, white; odor mild. GILLS adnate to slightly decurrent or often notched,
close, white or pallid but sometimes becoming straw-colored in age. STALK 3-10 cm
long, 1 -2.5 cm thick, solid, equal or tapering downward, often curved, smooth, dry, white,
sometimes discolored brownish in age, especially at base. SPORE PRINT white; spores
4-6 microns, round or nearly round, smooth. Basidia with siderophilous granules.
HABITAT: Gregarious on the ground, usually in large compact clumps and often half-
hidden by leaves and grass; fruiting mainly in disturbed areas—along roads and beaten
paths (sometimes even forcing its way up through asphalt), in waste places, around old
sawdust piles, or sometimes in the woods; widely distributed. It is common along the west
coast from Alaska to southern California and fruits in our area from early fall through
spring or even summer. It was the single most abundant mushroom on the University of
California, Santa Cruz campus (a very “disturbed” place!) during the 1976-77 drought. A
couple hundred pounds could easily have been harvested.
EDIBILITY: Edible and quite popular in some regions, but it hardly tastes like chicken as
its common name implies. I’ve found it to be crunchy, but bland. Its abundance and habit
of fruiting when other good edibles are scarce make experimentation worthwhile. Be
careful, however—this is a species “complex,” and some mild cases of poisoning have been
attributed to it. These may simply have been “allergic” reactions, or may have resulted
from confusion with poisonous species (e.g., Clitocybe dilatata).
COMMENTS: Also known as Clitocybe multiceps and Tricholoma aggregatum, this
exceedingly common species “complex” is best recognized by its fondness for disturbed
places and its growth in dense clusters which may contain a hundred or more individuals
and weigh as much as 15 pounds! Within the L. decastes group at least three species have
Lyophyllum decastes group is especially common along roads and paths. Note clustered growth
habit, white stalk, and absence of a veil. Another clump is shown on p. 898.

been recognized on the basis of cap color: L. connatum has a white or whitish cap; L. lori-
catum has a blackish-brown to dark brown cap when young, often with a hoary sheen or
metallic luster, and a thick cartilaginous cap cuticle; as it ages, however, it fades to paler
brown or tan, and is then indistinguishable from “typical” L. decastes, which has a brown
to grayish-brown to tan cap. L. loricatum is the most common of the three in our area,
but L. decastes also occurs. A fourth, unidentified species is fairly common in our
live oak woodlands. It has a grayish cap and grayish gills and its edibility is unknown.
Still another species grows under mountain conifers soon after the snow melts.
Though all of these species tend to grow in clusters, solitary individuals occasionally
occur, and these are apt to baffle the beginner. Since the cap color and gill attachment are
so variable, it is best only to eat those growing in large clumps in disturbed ground. Make
sure the spore deposits on the lower caps of mature clusters are white. Poisonous Ento-
lomas sometimes grow in clusters, but have deep pinkish spores. The Clitocybe sub-
connexa group also grows in clusters and has pinkish spores, while C. dilatata is white-
spored with a whitish cap and disagreeable taste. Most other fleshy, clustered terrestrial
mushrooms have darker spores and/ or a veil.

Lyophyllum montanum (Snowbank Lyophyllum)

CAP 2-6.5 cm broad, convex to broadly umbonate or plane; surface not viscid, gray to
lead-colored beneath a hoary (whitish) bloom which may wear off; often becoming some¬
what yellower (to dingy yellow-brown) in old age. Flesh thin, brownish to whitish, odor
mild. GILLS dark gray to gray, close, adnate to nearly free (usually adnexed). STALK 3-7
cm long, 0.5-1.5 cm thick, equal or thicker below, hoary and colored more or less like cap
when fresh, or slightly browner; base often with white mycelial down. SPORE PRINT
white; spores 6.5-8 * 3.5-4 microns, elliptical to oblong, smooth. Basidia with siderophilous
granules.
HABITAT: Solitary, scattered, or in small groups on ground under conifers (particularly
spruce and fir), usually near melting snow; fairly common in the spring and early summer
in the mountains of western North America.
EDIBILITY: Unknown.

175
176 TRICHOLOMATACEAE

COMMENTS: There are a number of nondescript grayish Lyophyllums that are very
difficult to identify. This one, however, can be told by its snowbank milieu and hoary cap
surface when young (see photo on p. 46). Other species: L. semitale is one of more than 50
difficult-to-differentiate Lyophyllums that stain gray, black, or brown when bruised
(often slowly). Its cap and gills are grayish and it is widespread under conifers or less com¬
monly hardwoods. L. infumatum also blackens, but has whitish gills when young.

Calocybe carnea
CAP 1.5-4 cm broad, convex to plane or slightly umbonate; surface dry, more or less
smooth, usually pinkish to pinkish-brown, but varying to dark reddish or fading to pale
tan. Flesh thin, whitish. GILLS crowded, narrow, white, adnate to slightly decurrent or
notched. STALK 1.5-4 cm long, 2-5 mm thick, equal, colored more or less like cap,
smooth or finely fibrillose. SPORE PRINT white; spores 4-6 * 2-3 microns, elliptical,
smooth. Basidia with siderophilous granules.
HABITAT: Scattered or in groups in lawns, grassy clearings in woods, and other open
places; widely distributed but not common. I have found it only once, on a lawn in Santa
Cruz, California, during the seventh inning stretch of the fourth game of the 1978
World Series (I don’t remember the date).
EDIBILITY: Not recommended. It is said to be edible but is easily confused with
poisonous species.
COMMENTS: Formerly known as Clitocybe socialis and Lyophyllum carneum, this
pretty little lawn-lover can be recognized by its pinkish cap, crowded white gills, and white
spores. It appears to have a wide distribution but nowhere does it seem to be common.
A similar but slightly larger species with a purple-red cap and yellow-ochre gills, C.
onychina, occurs in the spruce-aspen forests of the Rocky Mountains in the summer.

TRICHOLOMA
Medium-sized to large terrestrial, mostly woodland fungi. CAP viscid or dry, smooth, fibrillose,
or scaly. GILLS typically notched or adnexed, occasionally adnate. ST ALK central, fleshy. VEIL
absent (except in T. zelleri and a few others). VOLVA absent. SPORE PRINT white. Spores
smooth, not amyloid. Gills only rarely with cystidia.

TRICHOLOMA is a large and prominent, well-defined group of terrestrial white-spored

mushrooms with notched gills and a central, fleshy stem. It corresponds in stature to
Entoloma (deep pinkish spores) and Hebeloma (brown spores), but has little else in
common with those fungi. Among the white-spored genera, Collybia differs in its carti¬
laginous stalk, Tricholomopsis grows on or near wood, Hygrophorus has gills that are soft
and waxy, Russula has dry, brittle flesh, and Clitocybe has adnate to decurrent gills and/ or
pinkish spores. Melanoleuca, Leucopaxillus, and Lyophyllum differ microscopically and
are keyed out in Tricholoma, while Armillaria and Catathelasma have a veil.
Tricholoma contains several excellent edible species as well as some poisonous ones.
The man on horseback, T. flavovirens, is the only species I consider safe for beginners. The
brown-capped and gray-capped species should be strictly avoided until you are intimately
familiar with each and every one of them. Then, and only then, should species like T. por-
tentosum become part of your culinary repertoire. Tricholomas, incidentally, form the
bulk of the wild mushrooms served by the Czarnecki family at their fabulous wild
mushroom restaurant in Reading, Pennsylvania.
TRICHOLOMA 177

Tricholomas are almost exclusively woodland fungi. Most species are mycorrhizal—
especially with pine and oak, but also with spruce, aspen, fir, and other trees. They are
partial to cold weather and in our area reach their peak in December or January. In colder
regions they may continue to fruit after there is snow on the ground!
Though Tricholoma is an easy genus to recognize, its species are perplexing even to the
professional. Several kinds are known only from a single locality and microscopic charac¬
teristics are not as helpful as in, say, Mycena. Over 100 species occur in North America
and perhaps 50 in California. Fifteen are described here.
Key to Tricholoma
l. Cap yellow to greenish-yellow, at least at margin, or yellow overlaid with purple-red fibrils 2
1. Cap not yellow at margin . 8
2. Stalk sheathed with cottony or shaggy scales . (seeArmillaria& Allies, p. 189)
2. Not as above . 3
3. Cap and/ or stalk with reddish-purple fibrils or scales; flesh pale yellow to yellow .
. (see Tricholomopsis rutilans, p. 145)
3. Not as above .4
4. Flesh yellow; odor typically unpleasant and pungent (like coal-tar gas); gills yellow, widely
spaced; cap not viscid .T. sulphureum, p. 179
4. Not as above . 5
5. Cap with blackish to dark brown to purple-gray or grayish center and/ or radiating fibrils . 6
5. Cap lacking dark radiating fibrils, entirely yellow or yellow at the margin and reddish-brown to
brown toward the center. 7
6. Cap yellow to greenish-yellow with blackish to dark brown center and radiating fibrils .
. T. sejunctum & others, p. 180
6. Not as above; cap streaked with grayish to purple-gray fibrils .T. portentosum, p. 180
7. Gills yellow . T. flavovirens, p. 179
7. Gills white .T. leucophyllum (see T. flavovirens, p. 179)
8. Veil present, usually forming a distinct annulus (ring) on stalk .9
8. Veil absent or evanescent, not forming an annulus ... 11
9. Cap small (up to 5 cm broad), gray to brownish-gray or bluish-gray; associated mainly with
willow; odor usually farinaceous . T. cingulatum
9. Not as above (but may have farinaceous odor) . 10
10. Stalk rather slender and fragile; veil fibrillose, usually forming only a slight ring on stalk . .. .
.(see Limacella glioderma, p. 291)
10. Stalk usually at least 1 cm thick; veil membranous, forming a persistent ring T. zelleri, p. 188
11. Stalk belted with rusty-orange scales or scurf up to a well-defined line near apex, sometimes
also beaded with orange droplets; cap viscid when moist, yellow-orange to tawny, orange,
rusty-brown, or greenish .T. aurantium & others, p. 187
11. Not as above . 12
12. Base of stalk(or interior at base) usually pale pinkish to pinkish-orange; cap not viscid, its color
variable but usually greenish to olive-gray, sometimes shaded with brown, or grayish at center
and pallid at margin; surface smooth or cracking, but without differently colored fibrils or
hairs . T. saponaceum, p. 184
12. Not as above . 13
13. Stalk with a tapered “tap root” extending deep into humus; stalk usually with a tough or carti¬
laginous outer rind .(see Caulorhiza umbonata&others, p. 218)
13. Not as above . 14
14. Odor pungent and very unpleasant(like coal-tar gas); cap pallid; gills widely spaced; associated
with conifers .T. platyphyllum (see T. sulphureum, p. 179)
14. Not as above . 15
15. Cap white to creamy when fresh (but may discolor or age yellow, reddish, tan, brown, etc.);
stalk lacking small yellow dandruffy scales or granules at apex . 16
15. Cap distinctly colored or at least with colored scales or fibrils, even when young, or if cap white
then stalk with yellow scales or granules at apex or growing under mountain conifers soon
after the snow melts .. 21
178 TRICHOLOMATACEAE

16. Fruiting body often huge (15-75 cm broad!); known from Florida .T. titans
16. Not as above . 17
17. Fruiting body soon developing reddish or rusty stains; taste often bitter; stalk usually longer
than width of cap.(see Collybia maculata, p. 217)
17. Not as above . 18
18. Cap with at least a few scattered grayish scales or a grayish center . T. pardinum, p. 183
18. Not as above . 19
19. Cap viscid when moist, white or in age often tinged or stained yellow,4-8 cm broad; stalk usually
scurfy-scaly; spores amyloid .(see M elanoleuca, p. 169)
19. Not as above; spores not amyloid . 20
20. Stalk usually hollow and as long or longer than width of cap, often slender; usually growing
under redwood (in California) . (see Flygrophoraceae, p. 103)
20. Not as above; stalk not hollow; occurring with various trees but not redwood .
.T. resplendens & others, p. 183
21. Found under mountain conifers in spring; odor strongly cucumber-like or fishy .
. (see Armillaria & Allies, p. 189)
21. Not as above . 22
22. Cap black to gray, grayish-brown, or purplish-gray, or whitish with gray to blackish scales or
fibrils . 23
22. Not as above . 32
23. Cap viscid when moist (but may dry out), often streaked . . . T. portentosum & others, p. 180
23. Cap not viscid . 24
24. Cap whitish with scattered pale gray to dark gray scales, at least at center; stalk thick (1.5-3 cm)
and fleshy . T. pardinum, p. 183
24. Not as above; cap usually darker ... 25
25. Taste acrid . T. acre (see T. virgatum, p. 181)
25. Taste not acrid . 26
26. Cap densely hairy, scaly, or fibrillose-scaly (scales sometimes sparse in age); cap not usually
umbonate at maturity; veil sometimes present when very young . 27
26. Cap smooth to radially fibrillose or streaked, sometimes conical or umbonate at maturity; veil
absent . 29
27. Fruiting body fairly small (cap usually less than 8 cm broad; stalk less than 1.5 cm thick) . 28
27. Fruiting body fairly robust (cap 5-18 cm broad; stalk typically at least 1 cm thick).
.T. atroviolaceum & others (see T. virgatum, p. 181)
28. Stalk with small grayish to blackish scales . . T. squarrulosum (see T. terreum group, p. 182)
28. Stalk lacking grayish to black scales .T. terreum group & others, p. 182
29. Growing on or near wood, or if not then stalk usually attached to white mycelial cords that
arise from wood; gills broad (deep) .(see Tricholomopsisplatyphylla, p. 146)
29. Not as above; typically terrestrial . 30
30. Cap streaked with radiating fibrils, conical when young, often with a pointed umbo in age . .
.T. virgatum, p. 181
30. Not as above; cap without fibrils, broadly convex to plane or with a low umbo . 31
31. Cap with a hoary bloom when young; gills grayish; growing under mountain conifers, usually
near melting snow; spores not amyloid . (see Lyophyllum montanum, p. 175)
31. Not with above features (but may grow near snow); gills usually white or whitish .32
32. Gills crowded, pale; spores amyloid, or if not then odor often heavy and sweet .
.(see Marasmius, Collybia, & Allies, p. 201)
32. Not as above . 33
33. Fibrillose veil present when young, sometimes forming a slight ring on stalk . 34
33. Veil absent . 35
34. Cap at least slightly viscid when moist; growing with both hardwoods and conifers .
. (see Limacella glioderma, p. 291)
34. Cap dry, not viscid; associated with conifers (mainly pine and spruce) . . . T. vaccinum, p. 186
35. Stalk apex with small yellow scales or granules .
.T. acerbum & others (see T. pessundatum group, p. 185)
35. Not as above . 36
TRICHOLOMA 179

36. Cap viscid when moist, dark to medium reddish-brown to reddish-tan, the margin often paler;
gills often developing reddish-brown spots or stains in age . 37
36. Cap not viscid (but may be colored as above) .39
37. Associated with poplar or cottonwood, usually in sandy soil. T. populinum, p. 185
37. Not as above; associated with other trees . 38
38. Odor distinctly farinaceous or cucumberlike .T. pessundatum group & others, p. 185
38. Odor more or less mild .T. ustale (see T. pessundatum group, p. 185)
39. Spore print creamy to yellowish; gills tan to ochre or pale pinkish-cinnamon in age; stalk
straight, usually long; cap smooth, without scales; spores amyloid (seeMelanoleuca, p. 169)
39. Not as above; spore print white; spores not amyloid .40
40. Typically growing in disturbed soil (along roads, paths, etc.); cap smooth, grayish-brown to
dark brown to brown or tan, etc. (but never reddish-brown) .(see Lyophyllum, p. 173)
40. Not as above; growing in woods .41
41. Cap grayish-olive-brown . T. sp. (unidentified) (see T. virgatum, p. 181)
41. Cap brown to reddish-brown, cinnamon-brown, pinkish-brown, flesh-colored, etc.42
42. Stalk 1 -3 cm thick; cap often smooth when young and scaly in age, especially toward the margin;
margin naked . T imbricatum, p. 186
42. Stalk usually less than 1.5 cm thick; cap scaly or fibrillose-scaly even when young; margin
usually with hairy or woolly veil remnants when young .T. vaccinum, p. 186

Tricholoma sulphureum (The Stinker)

CAP 2-8 cm broad, convex to umbonate or sometimes plane; surface dry, smooth, yellow
or sometimes olive-yellow, or tinged brownish to grayish-brown at the center. Flesh rather
thin, yellow; odor usually strong and repulsive, like coal tar gas. GILLS typically adnexed
or notched, broad, thick, well-spaced, yellow. STALK 4-10 cm long, 0.5-1 cm thick, often
rather long in relation to cap; more or less equal, smooth, dry, yellow to olive-yellow or
with darker fibrils. VEIL absent. SPORE PRINT white; spores 8-12 * 5-6 microns,
elliptical, smooth.
HABITAT: Scattered to gregarious on ground under both hardwoods and conifers,
widely distributed. It is fairly common in the Pacific Northwest under conifers and also
occurs in northern California in the late fall and winter, but I have yet to find it in our area.
EDIBILITY: Indisputably inedible because of the obnoxious odor; possibly poisonous.
COMMENTS: While several Tricholomas have a distinctly disagreeable odor, this
species has a downright disgusting one. By this trait alone it can be distinguished from the
edible man on horseback, T. flavovirens, which it rather resembles in color, and from
T. sejunctum as well. Other species; Another mushroom with the same odor is T. platy-
phyllum (-T. inamoenum?), a white to creamy species also found under conifers in the
Pacific Northwest.

Tricholoma flavovirens (Man On Horseback) Color Plate 33

CAP 4-15 (20) cm broad, convex becoming plane or with margin uplifted in age; surface
viscid when moist, smooth, entirely yellow or brown to reddish-brown toward the center
and yellow at the margin, or sometimes olive-yellow; margin at first inrolled. Flesh thick,
firm, white; odor farinaceous or sometimes like coconut. GILLS close, broad, notched or
adnexed, yellow. STALK 3-10 cm long, 1-3 (4) cm thick, equal or enlarged slightly at either
end, dry, smooth, solid; white to pale yellow or sometimes with darker stains at base. VEIL
absent. SPORE PRINT white; spores 6-8 * 4-5 microns, elliptical, smooth.
HABITAT: Scattered to densely gregarious under pines (rarely other conifers), often
partially buried or visible only as “mushrumps” in the duff; widely distributed. Common in
180 TRICHOLOMATACEAE

our area in the late fall and winter, usually in grassy, sandy, or shrubby areas with pine
present; however, a bright yellow form is found occasionally with madrone, and in the
Southwest it is quite common under aspen.
EDIBILITY: Edible and excellent—one of the least appreciated and most flavorful of our
fleshy fungi, though a few people are adversely affected by it. The viscid cap should be
brushed clean in the field or the skin peeled off, and the sand removed (if it is present).
COMMENTS: Formerly known as T. equestre, this delectable mushroom is as depen¬
dably yellow as the blewit is purple. It lacks the blackish radial fibrils at the center of the cap
characteristic of T. sejunctum, and it also lacks the veil characteristic of Armillaria albo-
lanaripes and various yellow Cortinarius species. T. sulphureum is similarly colored but
has a dry rather than viscid cap and usually smells awful. The yellow color plus the sticky
cap (when moist), absence of a veil, white spores, and habit of hiding under pine needles
combine to make it one of the safest—as well as tastiest—of gilled mushrooms. None of
which explains the misnomer “Man On Horseback”—it doesn’t look anything like a horse,
and most of the horseback riders I see are women ...
One local patch of T. flavovirens produces crops that sometimes have a very strong
coconut odor and taste, and at other times have the usual mealy(farinaceous) odor. A very
similar edible species, T. leucophyllum, is common under aspen in the Rocky Mountains
and Southwest, and may actually mingle with T. flavovirens. It differs only in having white
rather than yellow gills.

Tricholoma sejunctum
CAP 3-8 (10) cm broad, convex to plane or broadly umbonate; surface slightly viscid or
tacky when moist, smooth, yellow or greenish-yellow with dark innate (flattened) fibrils
or streaks radiating from the blackish to brown center; sometimes with small scales in age.
Flesh white or tinged yellow; odor farinaceous, taste often bitter or nauseating, but in some
forms mild. GILLS fairly close, typically notched; at first whitish or creamy-white, but
often becoming yellow near the margin of the cap or occasionally yellowish throughout.
STALK 5-8 (12) cm long, 1-2(3) cm thick, more or less equal or somewhat swollen below,
firm, smooth, whitish, but often developing yellowish tints. VEIL absent. SPORE PRINT
white; spores 5-7 * 4-5.5 microns, broadly elliptical or elliptical, smooth.
HABITAT: Scattered or in groups under both hardwoods and conifers, fruiting mainly
in the fall; widely distributed. It is fairly common in the Pacific Northwest, and I have seen
luxuriant fruitings in California in mixed woods and under manzanita.
EDIBILITY: Not recommended—it is insipid at best and poisonous at worst.
COMMENTS: This species is likely to be mistaken for the edible Tricholoma flavovirens
because of its yellowish cap color, but the radiating blackish or dark brown fibrils or
streaks at the center of the cap and the tendency of the gills to show yellow only near the cap
margin should distinguish it. Other species: T. cheilolamnium is very similar but has a
dry cap; it is fairly common in the Pacific Northwest under conifers.

Tricholoma portentosum (Streaked T richoloma)

CAP 4-12 cm broad, convex to obtusely umbonate or plane, or with uplifted margin in
age; surface viscid when moist, smooth but with a streaked or radially fibrillose
appearance, pale gray to dark gray, brownish-gray, or purplish-gray, the center sometimes
nearly black and margin often paler; yellow tints occasionally present also, especially in
age. Flesh white or tinged gray, fairly thick; odor and taste mild to farinaceous. GILLS
adnexed or notched, at first white, becoming grayish or sometimes pale yellow in age;
Tricholoma portentosum has a dark (purple-gray to blackish) streaked viscid cap, whitish gills, white
stalk, and white spores. Note how the gills are notched.

fairly close. STALK 5-10 cm long, 1-2.5 cm thick, more or less equal, firm, smooth,
dry, white or sometimes tinged yellow. VEIL absent. SPOREPRINT white; spores5-7 * 3-
5 microns, elliptical, smooth.
HABITAT: Scattered to gregarious on ground in woods, widely distributed. In most
regions it occurs with conifers, particularly pine, but in our area it favors live oak and
tanoak. It fruits in the late fall and winter and is one of the last Tricholomas to appear.
EDIBILITY: Edible and excellent, with a strong hearty flavor—but be sure of your
identification before eating it!
COMMENTS: The viscid cap separates this species from a multitude of grayish, dry-
capped Tricholomas, some of which are poisonous (see T. pardinum and T. virgatum).
The notched gills, fleshy white stalk, white spore print, absence of a veil, and gray to purple-
gray streaked cap are also important fieldmarks. T. sejunctum is somewhat similar, but
much yellower as a rule. Entoloma madidum is also similar, but has pinkish spores (and
mature gills) and usualy has a darker (cap-colored) stalk. T. niveipes is a very similar
edible pine-loving easterner; it never develops yellow tones and has narrower spores.

Tricholoma virgatum
CAP 3-8 (10) cm broad, conical to broadly conical to nearly plane with a pointed umbo;
surface dry, grayish to grayish-brown or grayish-purple (the center often darker, margin
paler), streaked with radiating fibrils or fibrillose scales. Flesh thin, white becoming
grayish; odor mild or earthy, taste usually sharp or acrid. GILLS adnexed or notched,
white to grayish, close. STALK 6-12 (15) cm long, (0.5) 1-2 cm thick, more or less equal,
solid, smooth or fibrillose, white or tinged gray. VEIL absent. SPORE PRINT white;
spores 6-7.5 * 5-6 microns, elliptical, smooth.
HABITAT: Solitary or scattered to gregarious in mixed woods and under conifers, widely
distributed; occasionally found in our area in the winter.
EDIBILITY: Not recommended—it may be poisonous, and it resembles T. pardinum,
which is definitely poisonous.
COMMENTS: The dry, fibrillose-streaked grayish cap which is conical when young
affords a good means of recognizing this species, which is also known as T. subacutum. The
cap is never viscid as in T. portentosum, and the fibrils and/or scales are radially arranged,
in contrast to the T. terreum group. Other species: T. atroviolaceum is a large (cap 5-18
cm), robust species with a convex to plane (not conical), densely fibrillose-scaly, blackish

181
182 TRICHOLOMATACEAE

to dark grayish-brown or violet-tinted cap. It has grayish or pinkish-tinged gills, a thick,

brownish, sometimes bulbous stalk, and a farinaceous odor; it is not uncommon in
California and the Pacific Northwest under conifers. A similar robust, unidentified species
is common in our area under hardwoods; it has a slightly paler (olive, dark gray, or brown)
cap and paler stalk. T. acre, fairly common under hardwoods in eastern North America
and the Rocky Mountains, is also similar but slightly smaller (cap 3-9 cm), with a grayish
fibrillose cap, white to grayish gills, and a distinctly acrid taste.

Tricholoma terreum group (M ouse Tricholoma)

CAP 2-5 (7) cm broad, conical or convex-umbonate becoming more or less plane in age;
surface dry, hairy or felty from a dense layer of mouse-colored (gray to black) scales; scales
often sparser and color often paler (silvery-gray or even white) in age and/ or toward
margin. Flesh thin, fragile, white to gray; odor and taste mild to slightly farinaceous.
GILLS adnate to adnexed or notched, fairly close, white to gray. STALK 2-5 (8) cm long,
0.5-1 cm thick, usually rather slender, equal or slightly thicker below, white or tinged
gray, smooth(but see comments), dry. VEIL absent or cobwebby and evanescent. SPORE
PRINT white; spores 6-8 * 3.5-5.5 microns, elliptical, smooth.
HABITAT: Scattered to densely gregarious onground under conifers; widely distributed.
Fairly common in our area in the late fall and winter under pine and Douglas-fir. The
largest fruiting I’ve seen was on a lawn under a huge pine.
EDIBILITY: N ot recommended. The flavor of our local variety is good, but this is a species
“complex” and is best avoided until better known. There is also the danger of confusing
larger specimens with T. pardinum or T. virgatum.
COMMENTS: The small size (for a Tricholoma), fragile flesh, and dry, scaly or “furry”
mouse-colored cap characterize a complex of confusing, closely related species best left to
the specialists. It is probable that they are all edible, but we can’t be sure until they are better
known. The most common North American member of the club is said to be T. myomyces,
which has an evanescent cortina (cobwebby veil) and smaller spores (the “true” T. terreum
lacks a cortina). Other species in the group include: T. squarrulosum, also very common in
our area, but with mouse-colored fibrils or scales on the stalk as well as the cap (see photo
below), and several slightly larger species, e.g., T. orirubens, with gills that redden in age;
T. argyraceum (-T. scalpturatum?), which favors hardwoods and has gills that yellow in
age, and T. acre (see comments under T. virgatum).

Two small, common, mouse-colored conifer-lovers. Left: Tricholoma squarrulosum (see comments
above) has a fibrillose-scaly cap and stalk. Right: Close-up of the smooth white stalk of T. terreum
(left hand specimen) and the scaly stalk of T. squarrulosum (on right).
TRICHOLOMA 183

Tricholoma pardinum (T iger T richoloma)

CAP 5-16 (25) cm broad, convex to plane; surface dry, whitish with small pale gray to dark
gray fibrillose or spotlike scales, at least at center. Flesh thick, firm, white; odor farina¬
ceous. GILLS notched or adnexed, close, white (rarely flushed pinkish), not stained or
spotted gray. STALK 4-15 cm long, 1.5-3 cm thick, equal or enlarged below, white or
sometimes tinged gray, smooth, firm, solid. VEIL absent. SPORE PRINT white; spores
7-10 * 5-6.5 microns, elliptical, smooth.
HABITAT: Solitary to scattered or gregarious cn ground in woods, widely distributed in
northern North America. It is sometimes abundant under conifers in the Pacific North¬
west and Rocky Mountains, but in our area it grows with tanoak and madrone in the
winter. It is sporadically common, i.e., abundant once every several years but otherwise
infrequent to rare.
EDIBILITY: Poisonous! It causes severe and persistent gastroenteritis that may require
hospitilization.
COMMENTS: Anyone tempted to eat Tricholomas should learn to recognize this
poisonous species. It is larger and fleshier than other grayish Tricholomas (T. terreum,
T. virgatum, etc.), and often paler. At times it is nearly white with a few very pale grayish
scales, and could then be mistaken for one of the white Tricholomas (see T. resplendens).
The stalk is not pinkish-orange in the base as in T. saponaceum.

Tricholoma resplendens (White Tricholoma)

CAP 4-10 cm broad, convex becoming plane or with margin uplifted; surface viscid when
moist, often shiny when dry; smooth, white or with yellowish center, often discoloring
slightly brownish in age, especially toward center. Flesh firm, white, odor mild. GILLS
notched or adnexed to occasionally adnate, white, close. STALK 3-8 cm long, 1-2.5 cm
thick, equal or narrowed at base, white, smooth, dry, solid. VEIL absent. SPORE PRINT
white; spores 5-7.5 * 3.5-4.5 microns, elliptical, smooth.
HABITAT: Solitary or scattered or in small groups in woods, especially with oaks; of
questionable occurrence in California, but I have encountered a very similar species under
live oak in the winter. In eastern North America this species or one very similar is some¬
times common.
EDIBILITY: Not recommended—it is said to be edible, but is easily confused with poorly
known or poisonous species (see comments below).
COMMENTS: This species is one of several white or whitish Tricholomas, and can be
distinguished by the viscid cap when moist (however, see Melanoleuca lewisii in the key to
that genus, for it also has a viscid cap when moist, but tends to age yellowish and has
amyloid spores). Tricholomas with a non-viscid, whitish cap include: T. sulphurescens,
which stains or discolors yellow quite readily; T. venenata, poisonous and common under
hardwoods in eastern North America, with cap, gills, and stalk that discolor brownish-
buff in age or where injured, and a bitter taste; T. album, taste also bitter or sharp, but not
discoloring so much; T. columbetta, edible, cap often spotted with blue, green, or pink
in age, gills crowded, and odor mild; and T. gambosum(-T. georgii, Calocybegambosa),
edible, large and fleshy, with a white to creamy-buff cap, crowded gills, and strongly
farinaceous odor, favoring grassy areas and open woods. Whether any of these species
occur in California is uncertain, but they are to be expected. They might be confused with
certain Hygrocybe species (e.g., H. subaustralis, H.fornicata), but the latter have hollow
or stuffed stems and slightly waxy gills, and are most common under redwood in our area.
Also see T. manzanitae(under the T. pessundatum group), which can be whitish in youth.
Tricholoma saponaceum is an extremely variable species. Cap at top is greenish-gray, while the small
one at the bottom is brown. In all specimens, however, the flesh in the stalk base is pinkish to orange.

Tricholoma saponaceum (S oapy Tricholoma)

CAP 4-12 (18) cm broad, convex to plane or with uplifted, often wavy margin; surface dry
or moist but not viscid, smooth or cracking into scales in dry weather; color variable: olive
to greenish-gray, gray, yellowish-olive, brownish-olive, grayish-brown, coppery, or with
rusty tints, or sometimes dingy gray at center and pallid toward the margin. Flesh thick,
white, but may stain slowly yellowish or pinkish when bruised; odor and taste mild to
farinaceous, soapy, or “of wash rooms.” GILLS adnate to adnexed or notched, well¬
spaced, rather thick, white or tinged olive or yellowish, sometimes stained reddish. ST ALK
5-12 (20) cm long, 1 -3 cm thick, shape variable but often thickest in the middle and tapered
below to a somewhat rooting base; solid, smooth or with small scales, white or tinted
variously with the cap color; base usually with pale pinkish to pinkish-orange interior.
VEIL absent. SPORE PRINT white; spores 5-6 * 3-4 microns, elliptical, smooth.
HABIT AT: S olitary, scattered, tufted, or in groups or troops under both hardwoods and
conifers, widely distributed. Common under conifers (especially spruce) throughout
much of the West, but in our area found under live oak, tanoak, and madrone in the late
fall and winter.
EDIBILITY: Inedible—it has an insipid or soapy taste and may actually be poisonous.
COMMENTS: As evidenced by the lengthy description, this is a vexingly variable species
and is rather difficult to recognize when greenish shades are not evident on the cap. One
fairly infallible (or less unreliable) feature, however, is the pinkish-orange color of the flesh
at or near the base of the stalk. This feature is normally visible in the majority of specimens
from any group, and serves to distinguish them from other Tricholomas. The gills are
rather soft and well-spaced, leading to confusion with Hygrophorus, but the cap is not truly
viscid. The cap is most often some shade of grayish-olive or yellowish-green, but may
develop brown or coppery tones, especially in dry weather. It may crack into scales, but
lacks the fibrillose scales of T. virgatum, T. pardinum, and others. In the Pacific Northwest
and Rocky Mountains T. saponaceum is at times so overwhelmingly abundant that it has
been called a mushroom “weed.” In our area it is not quite so prevalent and its beauty
is more readily appreciated. One especially attractive form has a bluish-green cap.

184
TRICHOLOMA 185

Tricholoma pessundatum group Color Plate 35

CAP 5-14(18) cm broad, convex then plane or with slightly uplifted margin; surface viscid
when moist, entirely reddish-brown to reddish-tan, or often with a paler (or even whitish)
margin; smooth or sometimes finely scaly in age; margin often lobed, at first
inrolled, sometimes faintly ribbed. Flesh thick, firm, white; odor strongly farinaceous or
like linseed oil. GILLS white, but often developing sordid reddish or reddish-brown spots
and stains; typically notched or adnexed, but at times adnate or even free; close. STALK
4-10 (14) cm long, 1-3 cm thick, equal or swollen or tapered below, solid, firm; whitish or
developing sordid reddish or brownish stains or fibrils, especially over lower portion. VEIL
absent. SPORE PRINT white; spores 4-6 * 2.5 A microns, elliptical, smooth.
HABIT AT: Scattered to densely gregarious in forests or under trees, widely distributed. It
is partial to conifers in most areas (including the Pacific Northwest), but is locally common
in the late fall and winter with live oak, and less commonly pine.
EDIBILITY: To be avoided, as evidenced by the following severely censored excerpt from
an erstwhile colleague’s memoirs: “This common, viscid, red-brown Tricholoma is
delicious when stewed slowly with zucchini and served steaming hot on rice with chicken
chow mein and white wine. Suffering from an acute attack of overconfidence, M. Henis
and C. Cole tried it in this manner one winter evening in order to determine its edibility.
They subsequently staggered thrugh an all-night ordeal of nausea, vomiting, and diarrhea,
in which not only the mushroom, but everything else, was expelled . .
It is probable, then, that this common, viscid, red-brown Tricholoma is poisonous,
although a violent allergic reaction on the part of M. Henis and C. Cole cannot be ruled out
entirely. It is suggested that those foolish enough to try it (or any other mushroom of
unknown edibility) should do so in extremely small amounts—without the rice, chicken
chow mein, and white wine, and by all means, regardless of one’s nutritional needs, culinary
quirks, or dietary deficiencies, without the zucchini.
COMMENTS: Several robust, viscid, reddish-brown Tricholomas with reddish-spotted
gills will more or less fit the above description, and I leave it to licensed tricholomatologists
to decide whether or not ours is the “true” T. pessundatum. Closely related species include:
T. albobrunneum, said to have a weaker odor and cap finely streaked with darker lines;
T. ustaloides, with a transient cortina (hairy veil) and sharply defined white zone at the
stalk apex; T.flavobrunneum(-T.fulvum), with pale yellow gills when young and yellow-
tinted flesh in the stem; and T. ustale, which lacks a farinaceous odor. (See also T. popu-
linum.) All of the above have viscid caps when moist, and none have the belted scales on
the stalk or the veil characteristic of T. aurantium and T. zelleri respectively. Other
species: T. manzanitae is a manzanita- and madrone-loving Californian with pale yellow
granules or dandruffy scales at the stalk apex and a viscid cap that ranges from white( when
young) to pinkish, orangish, or brown (often with reddish stains). T. acerbum is also said
to have a yellow-dandruffy stalk, but has a strongly inrolled, ribbed cap margin (at least
until maturity) and a fragrant odor and/ or sharp taste. N one of the above should be eaten.

Tricholomapopulinum (The Sandy; Poplar Tricholoma)

CAP 5-16 cm broad, convex with an inrolled margin becoming plane or with uplifted
margin; surface viscid when moist, then dry, often radially streaked or with watery spots;
smooth, dull reddish-cinnamon to pale dingy reddish-brown, the margin usually paler or
whitish. Flesh firm, white, thick; odor and taste strongly farinaceous. GILLS typically
adnexed or notched, close, white, developing reddish-brown spots and stains, especially on
edges. STALK 2.5-7.5 cm long, 1-3 cm thick, equal or enlarged below, solid, firm, dull
whitish, developing dingy reddish-brown stains in age or after handling. VEIL absent.
SPORE PRINT white; spores 5-6 * 3.5-4 microns, elliptical, smooth.
186 TRICHOLOMATACEAE

HABITAT: Scattered to densely gregarious, frequently fruiting in large rings or dense

masses in sandy soil or along rivers, apparently always in association with poplar or
cottonwood. It fruits in cool weather and is widely distributed in western North America.
I have found it only once in our area, in December, but it is commoner inland. It is said
to be abundant in the J ohn Day country of eastern Oregon in the fall and early winter, and
I have seen very old specimens (perhaps from the previous fall?) in the spring near Pecos,
New Mexico, while looking for morels and wild asparagus.
EDIBILITY: Edible and popular in the Pacific Northwest, but be absolutely sure it is
associated with cottonwood—the similar T. pessundatum group is poisonous!
COMMENTS: The association with cottonwood and somewhat paler cap distinguish
this species from other viscid, red-brown Tricholomas (see the T. pessundatum group).
Its penchant for growing in densely-packed masses or long arcs is also distinctive.

Tricholoma vaccinum
CAP 4-7 (10) cm broad, broadly conical to convex, becoming umbonate or plane; surface
dry, covered with dark reddish-brown to rusty-cinnamon-brown to pale pinkish-brown,
tan, or flesh-colored fibrils or scales on a buff background; often darker at center; margin
with hairy veil remnants at least when young, often splitting in age. Flesh white or pallid;
odor usually farinaceous but sometimes mild. GILLS adnate becoming notched, close,
whitish or buff when young, but usually tinged flesh-color to pale cinnamon in age; some¬
times also with darker stains. STALK 3-8 cm long, 0.8-1.5 cm thick, equal or thicker at
either end, dry, smooth or with brownish to reddish-brown fibrils or small scales, usually
hollow at least in age. VEIL woolly-fibrillose, not forming an annulus (ring) on stalk, but
usually leaving traces on cap margin. SPORE PRINT white; spores (4) 6-7.5 * 4-5
microns, elliptical, smooth.
HABITAT: Scattered or in small tufts, groups, or large troops under conifers, especially
pine and spruce; common and very widely distributed, fruiting from late summer through
early winter. I have seen enormous fruitings under spruce in the Rocky Mountains
and under pine on the northern California coast.
EDIBILITY: Listed as mildly poisonous by some authors. Like cheap coffee and frozen
French fries, it is best avoided.
COMMENTS: One of the commonest Tricholomas of the coniferous forests of North
America, this species often fruits with T. imbricatum, but is apparently replaced by that
species in our local coastal pine forests. The two are quite similar, but T. vaccinum has a
scalier cap, frequently hollow stalk, and woolly veil which normally leaves hairs on the
cap margin. Several color forms occur, ranging from dark reddish-brown to pale pinkish-
brown, and the size is also variable. Usually it is smaller and more slender than T. imbrica¬
tum, but in northern California and Oregon a fairly robust, reddish-brown form occurs.

Tricholoma imbricatum
CAP 4-12 (20) cm broad, convex with an inrolled margin, becoming convex-umbonate
to plane or uplifted; surface dry, dark brown to brown or cinnamon-brown, with flattened
fibrils that may break up into scales in age, especially toward margin (which may be ob¬
scurely ribbed). Flesh thick, firm, white; odor mild or faintly farinaceous. GILLS adnexed,
notched, or even adnate; close, white or tinged flesh-color, often discoloring brown in age,
especially on the edges. STALK 4-12 cm long, 1-3 cm thick, solid, firm, dry, equal or
swollen below with a tapered, sometimes rooting base; white or buff becoming brownish in
age, especially over lower portion (apex usually pallid); fibrillose or minutely scaly in age.
VEIL absent. SPORE PRINT white; spores 5-7 * 3.5-5 microns, elliptical, smooth.
Tricholoma imbrication, young specimens. This very common conifer-lover has a dull brown dry cap.
In age the cap often flattens out or becomes wavy. For close-up of gills, see photo on next page.

HABITAT: Solitary to scattered or densely gregarious under conifers, particularly pine

and spruce(often hidden by needles); widely distributed. Itisaprominentfungalfeature of
our coastal pine forests in the winter and early spring.
EDIBILITY: Reportedly edible, but not recommended because it is easily confused with
members of the poisonous T. pessundatum group. It is rather tough anyway.
COMMENTS: This common Tricholoma can be told from other members of the genus by
its dry, dull brown cap, solid stem, absence of a veil, and generally coarse, robust
appearance, though slender individuals also occur. The species epithet, which means
“shingled,” is somewhat misleading, since the cap is usually quite smooth in youth and only
somewhat scaly in age (it is rarely truly “shingled” in the way that the tooth fungus Hydnum
imbricatum is). Its closest relative, T. vaccinum, has a scalier cap, evanescent veil, and
hollower stem, while the T. pessundatum group has a viscid cap (at least when moist). It
looks somewhat like a Russula, but has a tough, fibrous stem. Leucopaxillus amarus also
has a brown cap, but its gills remain white and it has a bitter taste and amyloid spores.

Tricholoma aurantium
CAP 4-10 cm broad, convex becoming obtusely umbonate or plane; surface viscid when
moist, smooth or breaking into small scales (especially at center); color variable: yellow-
orange to tawny, bright rusty-orange, orange-brown, orange-tan, or even orange-red,
sometimes splashed with olive-green or in one form entirely deep olive-green when young;
margin at first inrolled, sometimes beaded with orange droplets when moist. Flesh thick,
white; odor and taste strongly farinaceous and disagreeable (like rancid oil or cucumber).
GILLS adnate to adnexed or notched, close, whitish, often developing rusty-brown or
reddish-brown spots and stains. STALK 3-8 cm long, 0.8-2 cm thick, equal or thicker at
either end, solid, firm, belted with rusty-orange scales or scurfy flakes up to a well-defined
line near apex, pallid above the line; in wet weather sometimes beaded with orange droplets
near the line. VEIL absent or very rudimentary. SPORE PRINT white; spores 4-6 * 3-5
microns, elliptical to nearly round, smooth.
HABITAT: Solitary or scattered to gregarious on ground in woods, widely distributed.
Throughout most of the West it is common under conifers or sometimes aspen; in our area
it can be found under madrone in the winter, but is fairly rare.
EDIBILITY: Indisputably unpalatable due to the obnoxious odor and taste.
COMMENTS: This species strongly resembles T. zelleri but lacks a membranous veil. Its
sharply defined line near the stalk apex, however, is suggestive of a veil and probably
represents a rudimentary one. The viscid, orange to orange-brown or olive-splashed cap,
rusty-spotted gills, and strong odor help distinguish it. Our local form(var. olivascenf!)
187
Left: Tricholoma imbricatum, close-up of gills. Right: Tricholoma aurantium, showing charac¬
teristic belts of scales or granules on stalk.

is frequently a deep olive-green when young, and when beaded with orange droplets is
quite striking. T. aurantio-olivaceum is similar in many respects, but is smaller and odor¬
less, and often has olive-stains on the gills in age; it occurs in the Pacific Northwest.

Tricholoma zelleri
CAP 4-15 cm broad, convex becoming plane or broadly umbonate; surface viscid when
moist, bright orange to yellow-orange, or orange-brown, or sometimes splashed with
olive-green; margin at first hung with veil remnants. Flesh thick, white, slowly bruising
orange-brown; odor and taste strongly rancid-farinaceous. GILLS white, developing
rusty-orange-brown stains, close, adnate or notched. STALK 4-13 cm long, 1-3 cm thick,
usually tapered downward, solid, dry, pallid above the ring, usually somewhat scaly or
with orange or brown stains below. VEIL white, membranous, forming a flaring or ragged,
median to superior ring on stalk which frequently collapses in age. SPORE PRINT white;
spores4-5.5 x 3-4 microns, elliptical, smooth, not amyloid.
HABITAT: Scattered to gregarious on ground in woods, northern North America.

Tricholoma (-Armillaria) zelleri has the stature of a typical Tricholoma, but possesses a well-
developed membranous veil that usually forms an annulus (ring) on the stalk.
TRICHOLOMA 189

Extremely abundant under conifers in the Pacific Northwest (often in the same areas as the
matsutake, Armillaria ponderosa), but rather rare in our region and fruiting mainly in
tanoak-madrone woods at higher elevations in the coastal mountains (like A. ponderosa),
in the late fall and early winter. I have also seen it fruiting in large numbers with A rmillaria
caligata under spruce in the southern Rockies.
EDIBILITY: Not edible because of the unpleasant taste and smell; however, it is a good
matsutake-indicator!
COMMENTS: This is essentially a veiled version of T. aurantium—same color, odor,
taste, and habitat. It is better known as Armillaria zelleri, but because of the obvious
affinity with T. aurantium, it is now placed in Tricholoma. The sticky yellow-orange to
orange, brown, or greenish-splashed cap, plus the attached gills, presence of a membranous
veil, and white spores are diagnostic. It might possibly be confused with Limacella glio-
derma, which has a redder cap, fibrillose veil, and fragile stem. Other species: T. robustum
and T. focale are very similar if not the same (they are said to be reddish-brown in color).

ARMILLARIA & Allies

Medium-sized to very large, fleshy mushrooms found on ground or wood. CAP convex to plane.
GILLS attached. STALK central, fleshy. VEIL typically present, well-developed, usually forming
an annulus (ring) on stalk. VOLV A absent. SPORE PRINT white or tinged yellow. Spores smooth,
amyloid (Catathelasma), to weakly amyloid or not amyloid (Armillaria and Armillariella).

GROUPED here are three small genera of fleshy, white-spored mushrooms with a cottony
or membranous veil that usually forms a distinct ring on the stem. There are no warts on
the cap nor is there a volva on the stalk as in Amanita, the gills are not free as in Lepiota
and Limacella, nor soft and waxy as in Hygrophorus, and the cap and stalk are not covered
with mealy granules as in Cystoderma. Tricholoma intergrades somewhat with Armil¬
laria, but as defined here does not usually have a veil (but see T. zelleri!).
The principal genus, Armillaria, is comprised of terrestrial forest mushrooms. The
honey mushrooms, Armillariella, somewhat resemble Armillaria but grow on wood, often
in large clusters. They were originally placed in Armillaria and some mycologists retain
them in that genus while transferring the Armillarias of this book to Tricholoma and a
separate genus, Floccularia. The third genus, Catathelasma, is terrestrial like Armillaria,
but has a double-layered veil, decurrent gills, amyloid spores, and a hard, often massive
fruiting body. It seems to be restricted to coniferous forests and is rather rare.
Several mushrooms in this group are prized edibles. The matsutake of Japan (Armillaria
matsutake) and its magnificent North American counterpart (A. ponderosa) are highly
esteemed by Japanese- and Korean-Americans. The honey mushroom (Armillariella
mellea) is well known and popular among fungophiles, while being well known and
singularly unpopular among gardeners and farmers. It grows wherever there are trees and
shrubs (even grape vines) and is almost as common in towns as in the woods. It has been
called the most serious plant disease in California gardens, because once it has infected a
bush or tree, there is no cure. One can only hope that it will co-exist with— rather than kill—
its host, and make the best of a sad situation by harvesting the mushroom bounty when it
appears! Six species of Armillaria, Armillariella, and Catathelasma are depicted here.
Key to Armillaria & Allies
1. Growing on wood (may be buried!) or at the bases of trees, sometimes in large clusters .... 2
1. Growing widely scattered to gregarious on ground (not normally in large clusters) .5
2. Cap with rusty-brown scales; stalk sheathed with similarly colored fibrils below the veil; gills
typically not decurrent; found on hardwoods in eastern North America; rare .. A. decorosa
2. Not as above; common . 3
 190 TRICHOLOMATACEAE

3. Cap some shade of yellow, tan, or brown; taste usually bitter (but not detectable by everyone);
stalk fibrous, with a stringy white pith inside, often long . 18
3. Not as above; cap paler in color or the stalk solid and hard .4
4. Cap often scaly; nearly always on or near conifers .(seeLentinus& Lentinellus, p. 141)
4. Not as above; usually found on hardwoods .(see P leurotus& Allies, p. 132)
5. Gills typically decurrent; fruiting body hard and thick-fleshed, often large (cap 7 -40 cm broad!);
odor variable but not spicy-fragrant; spores amyloid; found with northern conifers .6
5. Gills not decurrent, or if decurrent then not as above (not hard and thick-fleshed, etc.) .... 7
6. Cap dull white to grayish . Catathelasma ventricosa(see C. imperialis, p. 195)
6. Cap dingy yellowish to olive-brown to dark brown . Catathelasma imperialis & others, p. 195
7. Odor pleasingly spicy-fragrant (somewhat like cinnamon); fruiting body whitish when young,
but often developing cinnamon-brown or yellowish stains in age .A. ponderosa, p. 191
7. Not as above; not spicy-fragrant, or if so then fruiting body darker when young .8
8. Lower portion of stalk shaggy or conspicuously scaly and fruiting body showing at least some
yellow .9
8. Not with above combination of characteristics . 10
9. Cap smooth or with flattened fibrils .A. albolanaripes& others, p. 194
9. Fresh cap with yellow scales, at least near margin A. straminea (see A. albolanaripes, p. 194)
10. Cap and stalk covered with cinnamon-brown to chestnut-brown or vinaceous-brown threads
(fibrils) which may break up into scales; veil membranous, usually forming a ring (annulus)
on stalk; odor sometimes spicy-fragrant; typically found in summer or fall A. caligata, p. 192
10. Not as above; differently colored or veil not membranous or found in spring and early summer
(shortly after snow melts); odor not spicy-fragrant . 11
11. Odor typically farinaceous, cucumberlike, fishy, or like raw peanuts, or unpleasantly pungent
(crush flesh in the cap if unsure) . 12
11. Odor typically mild, not distinctive, or merely fungal. 15
12. Found under mountain conifers shortly after the snow melts; cap becoming ochre-buff to
grayish, brownish, violet-gray, etc. (occasionally whitish) .A. olida, p. 193
12. Not as above (habitat or season usually different) . 13
13. Cap bright yellow-orange to orange-brown, reddish-brown, rusty, pinkish-brown, brick-red
or splashed with green; odor farinaceous or rancid . 14
13. Not as above .20
14. Veil fibrillose, often disappearing; stalk up to 1.5 cm thick, often fragile (seeLimacella, p. 291)
14. Veil membranous, forming a ring; stalk not fragile .(see Tricholoma zelleri, p. 188)
15. Fruiting body white and gills usually decurrent or fruiting body developing reddish to vinaceous-
red stains or streaks and veil fibrillose, evanescent; found with conifers, especially at higher
elevations . (see Hygrophoraceae, p. 103)
15. Not as above . 16
16. Stalk with cottony or shaggy scales below the veil; fruiting body grayish or paler; found under
mountain conifers, especially in Rockies . . A.fuscaSc others (see A. albolanaripes, p. 194)
16. Not as above . 17
17. Stalk tough and fibrous, usually with a stringy white pith inside; cap usually with small dark
hairs or scales, especially toward center; taste usually latently bitter (but not detectable by
everyone); gills usually decurrent (but sometimes adnate); on wood or ground; common 18
17. Not as above; on ground .20
18. Veil absent .Armillariella tabescens( see A. me Ilea, p. 196)
18. Veil present, at least when young. 19
19. Stalk usually bulbous or thicker at base; veil cottony, not typically forming a prominent ring;
often on ground, not in large clusters . . Armillariella bulbosa (see A. mellea group, p. 196)
19. Not as above; often with a prominent ring, often clustered Armillariella mellea group, p. 196
20. Stalk viscid; odor alkaline; fruiting body whitish; rare . A. viscidipes
20. Not as above; stalk not viscid; fruiting body white or variously colored; common .21
21. Cap 2-4 (6) cm broad, not white; stalk 3-6 mm thick; odor usually farinaceous; usually found
with willow; spores not amyloid; not common .(see Tricholoma, p. 176)
21. Not with above features; common.(see Amanita, p. 263)
Armillariaponderosa, showing the veil that sheathes the stalk. Also see the color plate and the photo
on p. 49. (The latter photo shows a young specimen with an unbroken veil.)

Armillaria ponderosa (White Matsutake; Matsutake) Color Plate 37

CAP 5-20 (35) cm broad, convex to plane; surface dry or slightly viscid when moist, at first
white, but in age developing pale cinnamon to pinkish-brown or yellow-brown stains or
with fibrils that become these colors; margin at first inrolled and cottony. Flesh thick, very
firm, white; odor distinctly spicy-aromatic (like cinnamon). GILLS white, discoloring
or spotted rusty-brownish to cinnamon in age; crowded, adnate to adnexed or notched.
STALK 4-15 cm long, 1-5 cm thick, solid, tough, hard, equal or with a narrowed base;
white above the ring, usually scaly or fibrillose below and colored more or less like cap.
VEIL thick, membranous, sheathing the stalk, at first white; forming a prominent cottony
ring which flares outward at first, then collapses against the stalk in age. SPORE PRINT
white; spores 5-7 * 4.5-5.5 microns, broadly elliptical to nearly round, not amyloid.
HABITAT: Widely scattered to gregarious on ground in forests, thickets, and pine
barrens; found throughout northern North America, but particularly abundant in the
Pacific Northwest, where it is harvested commercially. In the mountains of Idaho,
Washington, and Oregon it is common under mixed conifers and second-growth Douglas-
fir, while on the coast it favors sandy pine forests. It also likes to lurk in thickets of
ericaceous shrubs (e.g., rhododendron, huckleberry, manzanita), which makes for very
difficult collecting. In coastal California, however, it prefers tanoak-madrone stands to
conifers, though it also fruits in sandy soil under manzanita with a pine canopy. In our
area “patches” are hard to find but fairly reliable, producing one crop each year, generally
in November or December.
EDIBILITY: Edible and highly prized by Asian-Americans. In San Francisco and San
Jose it sells fresh for as much as $25 a piece! However, its tough, chewy texture does not
appeal to everyone. Special techniques are required to render it tender while highlighting
its unique flavor. (If you bring some to me, I will give you a free demonstration!)
COMMENTS: The unique spicy odor—a provocative compromise between “red hots”
and dirty socks—is the hallmark of this magnificent mushroom. Its robust stature, whitish
color (at least when young), and prominent veil are also distinctive. It might be mistaken
for a “JAR” (Just Another Russula) or an Amanita, but the veil and odor distinguish it.
Old cinnamon-stained specimens may be rather unattractive, but the young, firm, white,
cottony buttons are undeniably gorgeous. The only other mushrooms with the same odor
are the fragrant form of A. caligata, the matsutake of Japan, plus some specimens of
Lentinusponderosus (which grows on wood), and Inocybepyriodora (a small poisonous
species with brown spores). Hygrophorus subalpinus and Catathelasma species are
somewhat similar, but lack the odor. The white matsutake has recently been transferred to
Tricholoma and given a new name, Tricholoma magnivelare.
Armiliaria caligata, mature specimens. Note the dark fibrils on the cap and stalk, and the prominent
annulus (veil).

Armillaria caligata
CAP 4-12 cm broad, broadly convex becoming plane or with uplifted margin; surface
dry, covered with flattened cinnamon-brown to chestnut or vinaceous-brown fibrils
which typically separate and cluster in age to form small scales or patches, revealing the
whitish to pinkish flesh beneath. Flesh thick, white; odor variable: distinctly spicy-fragrant
to fruity, mild, or unpleasant; taste mild to nutty, bitter, or disagreeable. GILLS close,
adnexed to adnate (rarely slightly decurrent); white, the edges developing brownish
stains. STALK 4-9 cm long, 1-3 cm thick, more or less equal, solid, firm, white or pallid
above the ring, fibrillose or scaly below and colored like cap. VEIL membranous, sheath¬
ing the stalk, forming a distinct flaring ring which collapses in age; underside colored like
cap, upper surface white. SPORE PRINT white; spores 5-8 * 4-5.5 microns, broadly
elliptical, smooth, not amyloid.
HABITAT: Solitary to scattered or in groups on ground in woods, widely distributed.
In the West, it fruits mainly under mountain conifers in the summer and fall, but isn’t com¬
mon. In eastern North America it occurs under oaks and ericaceous shrubs. The largest
fruiting I’ve seen was under spruce in the Rocky Mountains. I haven’t found it in our area.
EDIBILITY: Edible—the forms which do not have a disagreeable taste and/ or odor are
said to be as good as A. ponderosa!
COMMENTS: Also known as Tricholoma caligatum, this species is easily separated from
its cousin A. ponder osa by the cinnamon-brown to purple-brown fibrils on the cap and
stalk. It might be confused with Hygrophoruspurpurascens, but the latter has a fibrillose,
evanescent veil and a redder (not as brown) cap, plus slightly waxy gills. The above
description encompasses several varieties and forms of A. caligata. The western version
typically has dark fibrils and the spicy-cinnamon odor of A. ponderosa. Eastern material,
on the other hand, is apt to be more cinnamon-colored with a mild to fruity to pungent
or downright disgusting odor. Other species: The matsutake of Japan, A. (-Tricholoma)
matsulake, is very close to the fragrant western variety of A. caligata and may actually
be the same species. It is edible, of course, and highly prized.

192
Armillaria olida is a prominent “snowbank” mushroom of the Sierra Nevada and Cascades. These
specimens are fairly typical, except that the cap is sometimes paler. Note slight annulus formed by
the veil. The odor is also distinctive (see description).

Armillaria olida (Cucumber Armillaria)

CAP 6-15 cm broad, convex becoming plane to broadly umbonate, or in age often de¬
pressed or with an uplifted margin; surface dry to somewhat viscid, color variable: whitish
(when still under the duff) to gray to bluish-gray, purplish-gray, brown, or developing
olive, buff, or ochre tones; often overlaid with white cottony or fibrillose veil remnants
(these sometimes scattered but more often merged to form a central patch). Flesh thick,
firm, white; odor very distinctive: usually like cucumber, watermelon rind, rotting
potatoes, or freshly mowed grass, but sometimes like old fish. GILLS close, white or
tinged gray, typically adnexed or notched but sometimes adnate when very young and
free in old age. STALK 6-14 cm long, 1.8-4 cm thick, equal or swollen below (or even
with a bulb), solid, very firm, dry, white or at times pale buff, usually sheathed below the
ring by veil remnants which are often ochre- or cinnamon-tinged. VEIL cottony or fibril¬
lose, usually forming a slight median to superior ring on the stalk, but sometimes disap¬
pearing. SPORE PRINT white; spores 9-12.5 x 4.5-6.5 microns, elliptical, smooth, not
amyloid.
HABITAT: Solitary to gregarious or in small clumps of 2-3 individuals in duff under
mountain conifers, often partially buried or forming “mushrumps”; known only from the
West, fruiting during the spring or shortly after the snow melts. It is quite common in
the Sierra Nevada and is sometimes a good morel-indicator. I have seen large fruitings
in Yosemite National Park in April.
EDIBILITY: Unknown—or at least not commonly eaten. The taste is said to resemble
the odor, which is not particularly pleasing.
COMMENTS: When the veil remnants are not obvious, this robust springtime Armillaria
is likely to be mistaken for a Tricholoma. In fact, as the definition of Armillaria is nar¬
rowed and that of Tricholoma is broadened to include more veiled species, it will probably
be transferred to that genus. Its distinctive cucumber or fishy odor, grayish cap, and
growth under mountain conifers in the spring plus the frequent presence of veil remnants
on the cap and/or stalk form a distinctive combination of characteristics. The cap color
can be reminiscent of T. portentosum and T. virgatum, but those species lack a veil and
are not so strongly scented.

193
Armillaria albolanaripes, mature specimens. The golden to yellow-brown color (see color plate)
and the shaggy stalk are distinctive. A. straminea (not illustrated) is similar, but has a scaly cap.

Armillaria albolanaripes (Sheathed Armillaria) Color Plate 43

CAP 5-12 cm broad, convex or slightly umbonate to plane; surface moist or slightly viscid,
yellow to golden-yellow or more often brown at the center and yellow at the margin; with
flattened fibrils or scales which darken in age. Flesh white or tinged yellow; odor mild.
GILLS adnexed or notched, white to pale yellow, close. STALK 2-8 cm long, 1-2.5 cm
thick, equal or thicker below, dry, white above the ring, sheathed with soft cottony scales
below, the scales white at first, yellow- or brown-tipped in age. VEIL white, cottony,
fragile, leaving a ragged superior ring on stalk and/ or remnants on cap margin. SPORE
PRINT white; spores 5-8 x 3-5 microns, elliptical, smooth, weakly amyloid.
HABITAT: Solitary or in scattered groups in woods and along paths; widely distributed.
It is common under conifers in the mountains of western N orth America in spring, summer,
and early fall, but I find it only rarely in our area, usually under oak.
EDIBILITY: Edible but insipid—I have fried it. The closely related A. straminea (see
comments) is said to be a popular edible mushroom in Colorado.
COMMENTS: This handsome mushroom is easily identified by its yellow-brown cap,
creamy to pale yellow gills, and soft cottony scales on the stem. The veil can be seen in young
specimens but does not always form a distinct ring. The scaly stem might be mistaken for
a volva, but there are no warts on the cap. A manita aspera is similarly colored but has white
gills and a warted cap. In the montane aspen-conifer forests of the West, A. albolanaripes
has several close relatives, including: A. pitkinensis, with a grayer cap and stalk and only
slight yellow tints (but gills yellowish in old age); A.fusca, a grayish version with no yellow
at all; and A. straminea (-A. luteovirens), more widely distributed and colored like A.
albolanaripes, but with conspicuous bright yellow scales on the cap (or at least the cap
margin) as well as on the stalk. In some regions, whitish or“albino” forms of A. straminea
and A. albolanaripes can also be found. All of the above species have amyloid spores
and are given their own genus, Floccularia, by some taxonomists (the same ones who
retain the honey mushrooms in Armillaria and transfer the matsutakes to the genus
Tricholoma).
Cathathelasma imperialis. Note the decurrent gills, scaly or fibrillose cap, and prominent annulus
(ring) on stalk. C. ventricosa (not illustrated) is similar but has a paler cap.

Catathelasma imperialis (Imperial Mushroom)

CAP 10-40 cm broad, convex to plane; surface slightly viscid when moist but soon dry,
smooth, fibrillose-scaly, or cracked into scales or plaques(areolate); dark brown to brown,
dingy yellow-brown, or olive-brown. Flesh very thick(up to 15 cm!) and hard, white; odor
and taste sharply farinaceous. GILLS decurrent, pallid or buff to yellowish or pale grayish-
olive (in age), close, many forked. STALK 12-18 cm long, 3-8 cm thick, tapered below to a
bluntly pointed base; dry, dingy brownish to pinkish-buff below the ring; solid, hard. VEIL
membranous, double-layered, its lower surface often areolate while still covering the gills;
typically forming a double ring, the upper one thick, striate above, and often flaring; the
lower one sheathing the stalk as a thin membrane or gelatinous zone, or indistinct. SPORE
PRINT white; spores 10-15 x 4-5.5 microns, cylindrical, smooth, amyloid.
HABITAT: Solitary, scattered, or in groups on ground under conifers (mainly spruce
and fir), late summer and fall, northern North America. It is more common in the Rocky
Mountains than on the west coast. It does not occur in our area, but C. ventricosa (see
comments) is fairly common under Sitka spruce in northern California.
EDIBILITY: Edible and very tempting because of its size—but tough. Buttons are as
large as baseballs, and just as hard!
COMMENTS: This mountain of a mushroom may well qualify as the“Most Humongus
Gilled Fungus Among Us.” Its often gargantuan size, hard flesh, dingy brownish cap,
growth on the ground, double veil, and white spores set it apart. The cap may be
smooth, areolate, or fibrillose-scaly. The buttons are smaller, of course, but still distinct
by virtue of their hardness. Lentinus ponderosus is also gigantic, but grows from stumps
and lacks a veil. A slightly “smaller” sister species, C. ventricosa, also occurs under nor¬
thern conifers. Its stalk may be quite long but its non-viscid cap averages “only” 7-35 cm
across and is dingy whitish to grayish. It looks something like the white matsutake(74rm/7-
laria ponderosa) but lacks the spicy odor of that species. It is said to be a good edible in
spite of its hard, unpleasant-tasting raw flesh. Other species: C. singeri of the Rocky
Mountains looks like a Hygrophorus with its dingy yellowish viscid cap, but has amyloid
spores; C. macrospora has broad spores. All of these species grow and decay slowly.

195
196 TRICHOLOMATACEAE

Armillariella mellea group (Honey Mushroom) Color Plates 39,42

CAP 3-15 cm broad or more, convex becoming plane or sometimes broadly umbonate or
in age uplifted; surface viscid or dry, usually with scattered minute dark brown to blackish
fibrillose scales or erect hairs, especially toward the center; color variable: yellow, yellow-
brown, tawny, tan, pinkish-brown, reddish-brown, etc. Flesh thick and white when young,
sometimes discolored in age; odor mild, taste usually latently bitter. GILLS adnate to
slightly decurrent or sometimes notched; white to yellowish or sordid flesh-color, often
spotted darker in age. STALK 5-20 cm long, 0.5-3 (5) cm thick, tough and fibrous with a
stringy pith inside; usually tapered below if growing in large clusters, or enlarged below if
unclustered and on the ground; dry, whitish above the ring, soon yellowish to reddish-
brown below and often cottony-scaly when very young. VEIL cottony-membranous,
white to yellowish, forming a superior ring on stalk or occasionally disappearing. SPORE
PRINT white; spores 6-10 x 5-6 microns, elliptical, smooth, not amyloid.
HABITAT: In small or massive clusters on stumps, logs, and living trees, or scattered to
gregarious (occasionally solitary) on ground—but growing from roots or buried wood;
common on a wide variety of trees and shrubs, and practically worldwide in distribution.
In our area it occurs year-round, but is most common in the fall and early winter. I have
seen truly stupendous fruitings on oak as. well as walnut (in an orchard) and other trees.
It is a virulent parasite of timber, fruit, and garden trees, but can also be a harmless
saprophyte on dead trees or on the dead wood (heartwood) of living trees. It is called “oak
root fungus” in California because of its insatiable appetite for oaks, and “shoe string root
rot” because of the stringy black mycelial strands (rhizomorphs) by which the mycelium
spreads. These “runners” may extend up the host’s trunk or infect neighboring trees by
traversing great distances through the soil. On oak trees the mycelium can frequently be
seen as whitish fanlike growths between the bark and wood. It generally feeds on the roots
and lower trunk of its host, reducing it to a pathetic white, spongy pulp. The mycelium is
also thought to be the culprit responsible for the “aborted” fruiting bodies of Entoloma
abortivum. Actively growing mycelium may phosphoresce at night, giving the wood an
eerie luminous aura called “foxfire.” I nhabitants of subarctic regions are said to mark their
trails with bits of glowing wood infected by A. mellea.
EDIBILITY: Eminently edible. Use only firm caps and discard the tough stalks. It is an
abundant food source, crunchy in texture, and a very passable substitute for the shiitake
(Lentinus edodes) in stir-fried dishes. The bitter taste cooks out, but some forms are better
than others, and some(e.g., those that grow on buckeye or hemlock) can cause digestive
upsets. The common name, incidentally, is a reference to its color (which, like honey, is
extremely variable), not its taste (which isn’t the least bit sweet).
COMMENTS: There is very little that can or cannot be said about the honey mushroom.
Also known as A rmiliaria mellea, it is among the most variable and cosmopolitan of the
fleshy fungi, and in its innumerable guises will confound you time and time again. Espe¬
cially variable are its color, shape, viscidity, and manner of growth, but there are several
key, relatively constant features that distinguish it: (1) the presence of a veil (2) the
tough, fibrous stalk (3) frequent presence of small dark hairs on cap(4) the bitter taste when
raw (some people, however, are unable to detect it) (5) the growth on wood (though
it may be buried) (6) the white or faintly yellowish spores(in any mature cluster the lower
caps will be covered with white spore dust).
There are at least two distinct, widespread variants (one study recognized 14 different
“species” in the A. mellea complex). One has a yellow to yellow-brown cap that is viscid or
dry but becomes slimy in wet weather. It also has a yellow-tinged veil and tapered
stalk, and usually grows in clusters. The second variety, on the other hand, has a hairier
This form of the honey mushroom (Armiliariella mellea) usually grows in small tufts on the ground
rather than in large clusters on wood (as shown in color plates). Also, the stalk is usually swollen at
the base and the veil is fragile and cottony. It approaches the European form now called A. bulbosa.

pinkish-brown to reddish-brown or dingy brown cap with a white cottony veil and
frequently enlarged stem base. It grows scattered or in small tufts, often on the ground.
This form, which is close to A. bulbosa {a European species), is especially confounding
to beginners. Intermediate forms abound also. In view of the extreme variability, be¬
ginners should eat only those clearly growing in clusters on wood, and be certain that the
spores are whitish. The poisonous Galerina autumnalis grows on wood and has a ring on
the stalk, but is smaller and more fragile, with a smoother cap and brown spores. Pholiota
species also have brown spores, while Gymnopilus has rusty-orange spores. In the eastern
and southern United States you may encounter^, tabescens, a very similar, clustered,
wood-inhabiting, white-spored mushroom that lacks a veil (and annulus) and has a dry
cap. It is also edible.

SQUAMANITA
Fairly small to medium-sized terrestrial, mainly woodland mushrooms. CAP usually scaly,
fibrillose-scaly, or granulose. GILLS usually attached. STALK typically central, arising from a
conspicuous, cylindrical to bulbous, often hollow, underground"tuber.” VEIL typically present,
sometimes forming a slight annulus (ring) on stalk. VOLV A absent or present as a collar or scaly
rings above the “tuber.” SPORE PRINT white or pink. Spores smooth, thin- or thick-walled;
amyloid, dextrinoid, or neither. Hyphae in gill tissue typically parallel or nearly so.

THIS small, rare, oddball genus is distinct by virtue of the underground bulb or “tuber”
from which the stem arises. Many mycologists place it in the Agaricaceae (along with
Lepiota, Agaricus, and Cystoderma) rather than in the Tricholomataceae, but its affinities
are unclear. Since some Amanita species have a swollen, rooting stem base that could be
mistaken for a “tuber,” Squamanita has been keyed out under that genus. Amanitas,
however, differ fundamentally in their divergent rather than parallel gill tissue (a micro¬
scopic feature, see p. 19).
Squamanita is unlikely to be encountered by the average mushroom hunter. One
odoriferous species is described here and two others are keyed out.

197
198 TRICHOLOMATACEAE

Key to Squamanita
1. Cap and stalk grayish to purple-gray, lilac-gray, or darker, but covered with an ochre-brown
granulose coating, at least when fresh.S. paradoxum{see S. odorata, below)
1. Not as above; cap and stalk often scaly, but granulose layer absent .2
2. Fruiting body purple-gray to purple-brown except for the yellowish to buff tuber; odor dis¬
tinctly fruity (somewhat like grape soda) .S. odorata, below
2. Purplish tones absent; cap ochre to ochre-brown to buff, with a whitish or grayish tuber (or
clusters of tubers); found in eastern North America .N. umbonata

Squamanita odorata
CAP 1-4.5 cm broad, obtusely bell-shaped or convex, expanding somewhat in age but
usually retaining a broad umbo; surface dry, densely and coarsely scaly or fibrillose-scaly,
the scales often erect; usually more fibrillose toward margin; brownish-purple to purplish-
gray or lilac-gray, often darker in age. Flesh colored like cap; odor strongly and persistently
fruity-fragrant (like grape soda or grape juice). GILLS adnate or notched, fairly well¬
spaced, colored more or less like cap. STALK 1-3.5 cm long,(2)3-10 (15) mm thick, arising
from a swollen, sometimes hollow, juglike underground “tuber” 1-2.5 cm high and up to
2 cm thick; colored more or less like cap and covered with conspicuous scales like those on
the cap, except for the smooth, sometimes silky apex and yellowish to buff-colored
“tuber”; hollow or partially hollow in age. VEIL not forming a distinct ring on stalk.
SPORE PRINT pinkish; spores 6.5-9 * 4-6 microns, elliptical, smooth, not amyloid.
HABITAT: Usually in groups or clumps on ground in woods; widely distributed but
apparently very rare. I have examined specimens collected under conifers in Washington.
EDIBILITY: Unknown. Although too rare to be of value, the odor is certainly intriguing.
COMMENTS: Formerly known as Coolia odorata, this little mushroom is as bizarre as
it is rare. The coarsely scaly purplish cap and stalk, similarly colored gills, yellowish-buff
“tuber,” and strong grapelike odor make a most distinctive set of features. Other species:
S. paradoxum(-Dissodermaparadoxum) is gray to lilac- or purplish-tinted in age beneath
an ochre-brown granulose coating. It also occurs in the Pacific Northwest, but is rare.

CYSTODERMA
Small to medium-sized, terrestrial or wood-inhabiting mushrooms. CAP dry, with a coating of
mealy or powdery granules, at least when fresh. GILLS typically whitish or pallid, usually attached.
STALK central, lower portion sheathed with mealy granules or scales. VEIL present, oftenforming
an annulus (ring) on stalk. VOLVA absent. SPORE PRINT white. Spores smooth, sometimes
amyloid but not dextrinoid.

THE outstanding feature of this small genus is the layer of mealy granules that coats the
cap and lower stem. Rain may wash the granules off the cap, but the stem normally retains
them. A veil is always present and in several species it forms a prominent ring. Armillaria,
Armillariella, and Catathelasma have a veil and attached gills, but are larger and lack the
granulose coating. Most Cystodermas were originally placed in Lepiota and some myco¬
logists retain them in the same family. Lepiotas, however, typically have free gills, while
in Cystoderma the gills are usually attached to the stalk.
Cystodermas are common in northern coniferous forests, especially in beds of moss.
About 20 species occur in North America. Several are very attractive but little is known of
their edibility. Two species are described here; both are rare in our area.

Key to Cystoderma
1. Stalk generally 8 mm thick or more; fruiting body medium-sized . 2
1. Stalk generally less than 8 mm thick; fruiting body rather small or sometimes medium-sized 5
Cystodermafallax, mature specimen. Note the umbonate cap, prominent annulus (ring), and coating
of granules on the stalk and cap. It grows singly as well as in small groups or clusters.

2. Spore print pale yellow-brown to orange-buff; large . . . (seeRozites& Phaeolepiota, p. 411)

2. Spore print white or whitish; medium-sized . 3
3. Cap white when young (but often pale cinnamon or buff in age); rare . C.ambrosii
3. Not as above; cap not white .4
4. Cap more or less orange; veil forming a persistent, well-developed annulus (ring) on stalk;
found on rotting hardwoods, mainly in eastern North America . C. granosum
4. Cap cinnabar-red to rusty-orange, etc.; veil evanescent, not usually forming a well-developed
annulus; widely distributed .C. cinnabarinum(see C. amianthinum, p. 200)
5. Veil typically forming a distinct, well-developed annulus(ring) on stalk .6
5. Veil evanescent or merely forming a ragged zone at top of granular sheath on stalk .7
6. Cap white or tinged pinkish or lilac; rare .C. carcharias
6. Cap rusty-brown to tawny-brown; widely distributed and common .C.fallax, below
7. Growing on wood; spores amyloid .C. gruberianum (see C. amianthinum, p. 200)
7. Growing on ground or in moss; spores amyloid or not amyloid . 8
8. Cap often (but not always) radially wrinkled, tawny to ochraceous to brown or rarely white;
spores amyloid .C. amianthinum, p. 200
8. Cap dark reddish-brown to brick-colored to tawny or paler (rarely white), but not wrinkled;
spores not amyloid .C. granulosum (see C. amianthinum, p. 200)

Cystoderma fallax
CAP 2 -5 cm broad, convex to plane or frequently with an umbo; surface dry, with con¬
spicuous mealy granules which are erect at first but flattened and more powdery inage(or
often wear away completely); cinnamon-brown to rusty-orange to tawny-ochre; margin
often hung with remnants from the veil. Flesh thin, whitish or tinged cap color. GILLS
adnexed to adnate, close, white to pale pinkish-buff or tinged yellow. ST ALK 3-7 cm long,
3-5(7) mm thick, equal or enlarged below, smooth and pallid above the ring, sheathed with
cinnamon-brown to rusty-ochre granules or flaky scales below. VEIL forming a large,
delicate but persistent, often flaring ring on the stalk; ring median to superior, smooth and
pallid on upper side, colored like the cap underneath. SPORE PRINT white; spores
3.5-5.5 x 3-4 microns, broadly elliptical to nearly round, smooth, amyloid.
HABITAT: Solitary, scattered, or in small groups or tufts onground under conifers or in
mixed woods, sometimes also on rotting wood; widely distributed and common in the
199
200 TRICHOLOMATACEAE

summer and fall in the Pacific Northwest and Rocky Mountains. Fruiting in the fall and
early winter in our area, but rather rare.
EDIBILITY: Unknown.
COMMENTS: One of the most attractive and delicately adorned of our woodland fungi,
this Cystoderma is easily identified by its rusty-orange to cinnamon color, prominent ring
on the stalk, whitish gills which are attached to the stem, and granulose coating on the cap
and stem (rain may wash the granules off the cap, but not the stem). The illustration does
not do it justice, but since when is justice usually done?

Cystoderma amianthinum
CAP 2-5 cm broad, bell-shaped or somewhat conical becoming convex or umbonate to
nearly plane; surface dry, prominently wrinkled (radially) in one form; covered with mealy
or powdery granules which may wear off in age, tawny-ochre to ochre-brown, ochre-buff,
or yellowish; margin often hung with veil remnants. Flesh thin, odor mild or strongly
pungent. GILLS adnexed to adnate, crowded, white or creamy or tinged yellow-orange.
STALK 2.5 -7 cm long, 3-8 mm thick, equal or slightly enlarged below, smooth and whitish
above the veil, sheathed with granules or granulose scales below and colored like the cap.
VEIL fragile, forming a slight ring on stalk or often disappearing. SPORE PRINT white;
spores 4-7 * 3-4 microns, elliptical, smooth, amyloid. Cap cuticle staining rusty-brown
to reddish-brown in KOH (potassium hydroxide).
HABITAT: Solitary, scattered, or in groups under or near conifers, especially in moss;
widely distributed in northern regions and probably the most common member of the
genus. I have seen it in late summer, fall, and early winter in northern California and the
Pacific Northwest, but it does not seem to occur south of San Francisco.
EDIBILITY: Not recommended. Some sources list it as edible, but it doesn’t have much
substance and can be confused with poisonous species (e.g., Lepiota castanea).
COMMENTS: This petite mushroom is quite attractive when growing amongst colorful
lichens or in beds of bright green moss. It is best recognized by its granulose cap and stalk,
ochre color, and fragile veil which disappears or forms only a slight ring on the stalk (rather
than a prominent one, as in C.fallax). It is easily mistaken for a small Lepiota, but the gills
are usually attached to the stem rather than free. In one variety the cap is conspicuously
wrinkled, in another it is not. Other species: C. granulosum is similar, but has a reddish-
brown to tawny, non-wrinkled cap and non-amyloid spores; C. gruberianum is a small
species that grows on rotten wood; C. cinnabarinum is a larger, farflung species with a
rusty-orange to beautiful cinnabar-red or Vermillion cap and stalk. Whitish-capped
forms of C. amianthinum and C. granulosum also occur, but are rare.

ASTEROPHORA
Small mushrooms parasitic on other mushrooms. CAP often powdery. GILLS thick and well¬
spaced or poorly formed to practically absent. STALK present. VEILand VOLV A absent. SPORE
PRINT white to brownish when obtainable. Spores mostly produced asexually, smooth or spiny.

THIS small genus contains a staggering total of two species. Both are outlandish oddballs
that grow exclusively on other agarics, particularly Russula and Lactarius species. They
differ from Collybia tuberosa and other mushroom-inhabiting mushrooms in having
thick and well-spaced or poorly formed gills. They are also unique in that they produce
very few spores on basidia. Instead the hyphae block off to form asexual spores called
chlamydospores. Asterophora is listed in some books as Nyctalis.
ASTEROPHORA 201

Key to Asterophora
1. Cap more or less round and puffball-like, white becoming brownish and powdery as spores
mature; gills often malformed or practically absent .A. lycoperdoides, below
1. Not as above; cap not powdery; gills thick, well-spaced, usually decurrent, eventually disinte¬
grating into powdery spores .A. parasitica (see A. lycoperdoides, below)

Asterophora lycoperdoides
CAP 0.5 -2 cm broad, nearly round; surface dry, whitish becoming brown and powdery
from spores. Flesh thin, odor farinaceous. GILLS often malformed or barely present;
well-spaced, thick, whitish. STALK 1-3 cm long, 3-8 mm thick, more or less equal, white
becoming brownish. SPORE PRINT white when obtainable; spores 5-6 * 3.5^1 microns,
elliptical, smooth. Chlamydospores 12-18 microns, round, bumpy or spiny, thick-walled,
brownish.
HABITAT: In colonies on old mushrooms, particularly species in the Russula densifolia
group. Widely distributed but not common; very rare in our area.
EDIBILTY: Unknown.
COMMENTS: This oddball might be mistaken for a puffball because of its poorly formed
gills and powdered round cap. However, no puffballs are known to be parasitic on gilled
mushrooms! A. parasitica is also widely distributed, but even rarer than A. lycoperdoides.
It has thick, well-spaced, decurrent gills and a white to grayish, brownish, or lilac-tinged
cap, plus smooth and elliptical chlamydospores.

MARASMIUS, COLLYBIA,& Allies

Minute to medium-sized mushrooms, some of which shrivel up in dry weather and then revive when
moistened, others of which do not. CAP usually convex to plane, but sometimes bell-shaped;
not viscid in most cases; margin usually incurved when young. GILLS usually free, adnexed, or
notched, but sometimes adnate(or in Marasmius, even decurrent). STALK usually thin and pliant,
tough, cartilaginous, or wiry; usually central. VEIL and VOLVA absent. SPORE PRINT white to
buff or rarely tinged pinkish. Spores smooth, usually not amyloid. Cells in the upper layer of the
cap cuticle usually forming a palisade (Marasmius), or not forming a palisade (Collybia).

THESE minute to medium-sized mushrooms typically have adnexed to free gills and a
cartilaginous or wiry stem. The cap is typically convex to plane or if conical then with an
incurved margin when young (rather than straight as in Mycena). Two large genera
{Marasmius and Collybia) plus several smaller ones are treated together here because
they are difficult to separate in the field. The traditional trademark of Marasmius is its
astonishing reviving ability. If dried-up specimens are placed in a bowl of water they will
quickly swell up, magically reassuming their original shape and dimensions. In the wild,
species of Marasmius often seem to spring up in droves right after or during a rain, when
in fact they were already there, shrivelled up and inconspicuous. I n addition, they can often
be told by their tough texture and wiry or hairlike stem. {Xeromphalina is somewhat
similar, but usually has decurrent and/ or more brightly colored gills.)
Collybia has traditionally been separated from Marasmius on the basis of its slightly
fleshier, non-reviving fruiting body. However, some species have been shuttled back and
forth between Marasmius and Collybia because they revive somewhat when moistened.
Recognizing the arbitrary nature of this character, taxonomists now differentiate
Collybia from Marasmius primarily on microscopic features such as the structure of the
cap cuticle. As a result, Collybia, as currently defined, includes a few species which do
revive, while Marasmius includes some that do not. The gills in Collybia are usually ad¬
nexed or even free; in some cases they are more broadly attached, leading to confusion
with Clitocybe. The stalk is usually thin and pliant; if thick, it has a cartilaginous outer
202 TRICHOLOMATACEAE

rind that helps distinguish it from Tricholoma.

Both Collybia and Marasmius are “troubled” taxonomically. They have been fertile
fodder for the “splitters” (see p. 10), who have recently erected a number of “satellite”
genera. Some of these are easily distinguished in the field (e.g., Caulorhiza and Oudeman-
siella usually have a “tap root”; Flammulina has a viscid cap and velvety stem; Crinipellis
has dextrinoid hairs on the cap; Strobilurus usually grows on cones; Callistosporium is
olive-brown and yellow); others differ microscopically (e.g., Micromphale and Maras-
miellus); still others are not recognized here (e.g., Rhodocollybia and Microcollybia).
Marasmius is a very large genus centered in the tropics. As might be expected, it is more
diverse in the humid deciduous forests of eastern North America than in the West. Most
species are saprophytic on sticks and leaves and many are exquisitely constructed. Colly-
bias, on the other hand, are by and large a listless lot. They are also saprophytic on humus
and wood and are among our most common woodland agarics; a few may be mycorrhizal.
Most of the genera treated here are difficult from a taxonomic standpoint and have
little to offer the mushroom-eater. Two exceptional exceptions are the fairy ring mush¬
room or “Scotch Bonnet,” Marasmius oreades (forgive my promiscuous use of super¬
latives, but it is an exceptionally flavorful fungus!), and the garlic mushrooms (M. cope-
landi and allies). A representative sampling of “marasmioid” and “collybioid” fungi
is presented here and several additional species are keyed out but not described.

Key to Marasmius, Collybia, & Allies

1. Typically growing on fallen cones (sometimes buried!) or magnolia pods; rarely found on rotten
wood, and if so then stalk thin, more or less rooting, and hairy over lower portion .2
1. Typically growing on ground, wood, or other mushrooms . 5
2. Cap conical or bell-shaped when young, often reddish- or vinaceous-tinged; gills frequently
with reddish to dark purple edges .(see Mycena, p. 224)
2. Not as above . 3
3. Found on fallen magnolia “cones” .Strobilurus conigenoides (see S. trullisatus, p. 211)
3. Found on cones of conifers . . 4
4. Gills very crowded; spores amyloid . . Baeospora myosura (see Strobilurus trullisatus, p. 211)
4. Not as above . .Strobilurus trullisatus & others, p. 211
5. Stalk with numerous side-branches, at least on lower portion .C. racemosa, p. 213
5. Stalk lacking side-branches . 6
6. Cap minute (typically 1 cm broad or less) and pale; stalk whitish, very thin; usually (but not
always) colonizing the blackened remains of other mushrooms C. tuberosa & others, p. 212
6. Not growing on other mushrooms; if small, then not as above . 7
7. Fruiting body minute (cap less than 1 cm broad), stalk short, gills adnate to decurrent; growing
on bases of madrones Micromphale arbuticola (see Marasmius androsaceus group, p. 208)
7. Not as above . 8
8. Gills violet or lilac when fresh (but they may fade!). 9
8. Gills not violet or lilac . 10
9. Gills crowded; stalk not white; odor mild; spores amyloid; usually found on rotten wood;
widely distributed in northern latitudes .Baeospora myriadophylla
9. Gills well-spaced; stalk whitish to pale gray; odor usually unpleasant; spores not amyloid;
usually terrestrial; restricted to eastern North America (?) ... C. iocephala (-M. iocephalus)
10. Odor distinctly garlic- or onionlike, at least when flesh is crushed; cap small or minute (usually
less than 2.5 cm broad); often gregarious but not normally clustered . 11
10. Not as above; odor may be fetid or otherwise distinctive, but if garliclike then cap typically 2 cm
broad or more or fruiting bodies often clustered . 12
11. Stalk smooth (hairless); habitat variable .M. scorodonius (see M. copelandi, p. 207)
11. Stalk minutely hairy (use hand lens); found on leaves.M. copelandi & others, p. 207
12. Cap small or minute, with coarse tawny to brown hairs; stalk thin (less than 2 mm thick), also
hairy (minutely so), wiry-tough; hairs on cap dextrinoid . Crinipellispiceae & others, p. 210
12. Not as above; cap usually without hairs . 13
MARASMIUS, COLLYBIA, & ALLIES 203

13. Odor fetid; stalk velvety; gills yellowish to brown or tinged reddish; cap and stalk brown to red-
brown; found on sticks or bark in eastern U.S. Micromphalefoetidum (-Marasmiusfoetidus)
13. Not as above . 14
14. Odor sweet and heavy (like benzaldehyde); cap brown to reddish-brown, vinaceous-brown, or
dark brown, at least toward the center; stalk usually at least 5 mm thick . 15
14. Not as above . 16
15. Stalk white (but may develop vinaceous or brownish stains below) ... C. oregonensis, p. 218
15. Stalk brown to dark brown or vinaceous-brown C. subsulcatipes(see C. oregonensis, p. 218)
16. Growing in grass, often in arcs or rings; stalk tough; gills fairly well-spaced (not crowded or
close); cap white to tan, buff, or brownish but not gray or vinaceous, usually less than 6 cm
broad; spore print white; very common and widespread .M. oreades, p. 208
16. Not as above . 17
17. Stalk with a tapered underground “tap root”(dig up carefully!); spore print white; fruiting body
without reddish or rusty stains; cap opaque (not normally translucent-striate when moist) 18
17. “T ap root” lacking or not well-developed, or if present then cap translucent-striate when moist or
fruiting body often reddish-stained and spore print pinkish-buff . 19
18. Cap blackish, dark brown, grayish, whitish, or yellowish-brown (but if the latter then usually
viscid when moist); found from the Rockies eastward Oudemansiella radicata& others, p.219
18. Not as above; cap chestnut-brown to warm tan or yellow-brown, not viscid .
.Caulorhiza umbonata & others, p. 218
19. Spore print pinkish; growing in grass, straw, or manure .(see Clitocybe tarda, p. 152)
19. Not with above combination of features.20
20. Gills yellow; cap and stalk olive to olive-brown or yellowish (but may develop dark reddish-
brown tones as it dries); usually on rotten wood . . Callistosporium luteo-olivaceum, p. 211
20. Not as above .21
21. Stalk dark, stiff, bristle-like, lessthan 1 mm thick; cap typically less than 1 cm broad (rarely 2 cm);
cap not whitish when fresh (but may fade!); substrate (twigs, needles, leaves) usually with black
horsehair-like rhizomorphs (mycelial threads) .... M. androsaceus group & others, p. 208
21. Not with above features (but may have some of them) . 22
22. Gills adnate to decurrent and fairly well-spaced; cap 2-4 cm broad and predominantly whitish
(may be slightly darker at center), usually wrinkled; stalk becoming brownish from the base
upward; terrestrial in the forests of the Pacific Northwest .M. umbilicatus
22. Not as above .23
23. Gills adnate to decurrent, white, very widely spaced; cap white or tinged gray to olive-gray or
even slightly yellowish, translucent-striate when moist; stalk whitish; found under pine in
coastal California, often in large numbers .M. sp. (unidentified), p. 206
23. Not as above .24
24. F ruiting body small or minute (cap usually2 cm broad or less); cap white or whitish or tinged pale
yellowish (the center may be tinged brown), but may develop reddish or pinkish stains in age;
stalk less than 3 mm thick. 25
24. Not as above; either differently colored or larger . 29
25. Gills free or nearly free; margin of cap usually with veil remnants andl or the cap and stalk
minutely powdered; stalk whitish; growing on ground . (szeLepiota seminuda, p. 307)
25. Not as above . 26
26. Stalk black beneath a coating of minute white hairs; cap 1 -2 cm broad; found on leaves or twigs
in eastern North America .Marasmiellus nigripes{see M. candidus, p. 206)
26. Not as above ... 27
27. Growing on fallen leaves .M. delectans & others (see Marasmiellus candidus, p. 206)
27. Growing on sticks, berry canes, wood, etc.28
28. Gills very widely spaced; stalk relatively shortness than3 cm long); abundant on the west coast,
infrequent elsewhere .Marasmiellus candidus, p. 206
28. Not as above; abundant in eastern North America, rare or absent elsewhere on continent . . .
.M. rotula{see Marasmiellus candidus, p. 206)
29. Growing in compact bundles on rotting conifers (the wood sometimes buried or very decom¬
posed—see Color Plate 49); margin of cap incurved when young .30
29. Not as above (but may grow tufted on rotting conifers or in dense clusters on ground) ... 31
204 TRICHOLOMATACEAE

30. Stalk white to grayish .... Clitocybulafamilia& C. abundans(see Collybia acervata, p. 215)
30. Stalk vinaceous-brown to reddish-brown (or somewhat paler when dry), at least at apex . 55
31. Stalk solid or firmly stuffed (not hollow), straight and equal except for very base (which may be
slightly swollen), sometimes scurfy or dandruffy at apex or throughout, or longitudinally lined
(but without hairs); gills crowded, white (except for one large species with tan to pinkish-
cinnamon gills); cap typically rather flat (broadly convex to plane, sometimes with a blunt
umbo); surface of cap usually smooth and dark brown to grayish, sometimes ochre-brown,
yellowish, or whitish (but not reddish-brown or vinaceous-brown); spores amyloid, usually
roughened; found in many habitats, but especially in grass or landscaped ground or under
mountain conifers soon after the snow melts .(see Melanoleuca, p. 169)
31. Not as above; spores typically neither amyloid nor roughened; stalk sometimes hollow, some¬
times clothed with minute hairs but not often scurfy; gills crowded to widely spaced; usually
found in woods or near trees (but not always) . 32
32. Cap with gray to black hairs or fibrillose scales .(see Tricholoma terreum group, p. 182)
32. Not as above . 33
33. Stalk very thin (less than 1.5 mm); cap flesh-colored to light brown, often wrinkled, up to 12
mm broad; stalk very minutely hairy (pubescent), not shiny; found on needles and twigs of red¬
wood, spruce, fir Micromphale sequoiae & others (see Marasmius androsaceus group, p. 208)
33. Not as above . 34
34. Cap grayish to dark brown, olive-brown, or black; growing in moss, Sphagnum bogs, or on
burnt ground, or sometimes simply associated with conifers . 59
34. Not as above; cap differently colored or habitat different . 35
35. Stalk smooth (hairless) or finely powdered, or with hairs only at the base . 36
35. Stalk pubescent or velvety (covered with minute hairs) over at least the lower half by maturity
(use hand lens if unsure) . 51
36. Gills reddish-brown to dark brown or blackish-brown; cap and stalk similarly colored (but cap
may fade); flesh staining green in KOH; fairly common in eastern North America and the
Pacific Northwest .C. alkalivirens
36. Not as above; gills typically paler .37
37. Base of stalk with a litter-binding mycelial pad; cap yellowish-brown to reddish-brown; stalk
1-3 mm thick, shining; growing in groups or dense clusters on hardwoods leaves and debris
in eastern North America .M. cohaerens& others
37. Not as above . 38
38. Growing in grass; fruiting body small (cap usually less than 2.5 cm broad) and vinaceous- or
reddish-tinged; stalk not tough and polished; cap not pleated .
.Mycena sp. (unidentified) (see Marasmius oreades, p. 208)
38. Not as above . 39
39. Cap and stalk pale or whitish (cap may have grayish-brown center); gills fairly well-spaced;
known from California and South America.M. albogriseus(see M. oreades, p. 208)
39. Gills close or crowded, or if well-spaced then cap and stalk differently colored .40
40. Gills widely spaced; stalk usually polished; cap often (but not always) pleated .41
40. N ot as above; gills typically fairly close or crowded .42
41. Cap bay-brown to reddish-brown, brown, or wine-red; stalk 5-13 cm long; common on west
coast .M. plicatulus, p. 209
41. Not as above; either cap differently colored or stalk shorter or found elsewhere .
.M. siccus & others (see M. plicatulus, p. 209)
42. Cap striate when moist and often translucent; cap typically conical or bell-shaped when young
.(seeMycena, p. 224)
42. Not as above .43
43. Stalk tough, grooved or twisted, often with a rooting base, tan to brown (not white!); growing
on hardwoods; rare (not positively known from North America) .C. fusipes
43. Not as above; common . 44
44. Stalk 0.5-2.5 cm thick, often with a rooting base, white or yellowish (but may develop reddish
stains below); cap usually over 4 cm broad and whitish, but often becoming reddish, pinkish, or
vinaceous-brown at the center and sometimes entirely those colors from the beginning; found
under conifers, usually on decayed wood or lignin-rich humus .C. maculata, p. 217
44. Not as above; either differently colored, smaller, or with a different habitat . 45
MARASMIUS, COLLYBIA, & ALLIES 205

45. S talk dark red except at apex; cap tan to buff, often plane at maturity; growing in tufts or clusters
in humus and under trees; not common . C. marasmioides(see C. acervata, p. 215)
45. Not as above (if tufted or clustered then stalk differently colored, including reddish-brown) 46
46. Gill edges coarsely ragged or toothed, even when young; spore print white; spores amyloid;
widespread (but not reported from California) .(seeLentinus& Lentinellus, p. 141)
46. Gill edges entire or finely serrated (or in age sometimes coarsely serrated); spore print white or
slightly colored; spores rarely amyloid; very common in California and elsewhere.47
47. Cap yellowish to light brown, often fading to whitish; stalk whitish or tinged yellow; found under
eastern hardwoods (or mixed woods) M. strictipes& M. nigrodiscus (see M. oreades, p.208)
47. Not as above .48
48. Usually growing in grass and cap typically pinkish or growing under mountain conifers and cap
vinaceous-red to purplish-red with ochre-yellow gills .... (see Lyophyllum &, Allies, p. 173)
48. Not as above .49
49. Spore print white to pale cream; cap averaging 1-5 cm broad (occasionally larger); cap color
variable but often tawny; gills white or pale yellow, their edges often entire; spores not
dextrinoid; common under hardwoods and conifers.C. dryophila & others, p. 215
49. Spore print cream to buff or pinkish-buff; cap averaging 3-8 cm broad (sometimes larger),
various shades of brown but not tawny; gills white or with reddish stains, the edges often finely
scalloped at maturity; at least some of the spores dextrinoid; found mainly(but not exclusively)
under conifers . 50
50. Cap vinaceous- to reddish-brown, not fading appreciably; gills sometimes reddish-stained . .
.C. extuberans & others (see C. butyracea, p.216)
50. Cap reddish-brown, brown, tan, or even grayish; gills not reddish-stained C. butyracea, p.216
51. Cap velvety, more or less orange-brown; gills adnate to decurrent; growing on hardwoods in
eastern North America .(see Omphalina&Xeromphalina, p. 221)
51. Not as above; stalk may be velvety but cap not velvety and gills not decurrent .52
52. Cap usually viscid when moist (but may dry out!); lower portion of stalk rusty-brown to
blackish-brown and velvety when mature (usually smooth and pallid when young); found on
wood (sometimes buried!) .Flammulina velutipes, p. 220
52. Not as above; cap not normally viscid .53
53. Odor garlicky or taste distinctly acrid (burning) .
.C. polyphylla& C. peronata{see C. confluens, p. 213)
53. Not as above (but taste may be somewhat bitter) . 54
54. Gills usually crowded (sometimes merely close), white or tinged flesh-color; cap not prominently
wrinkled; stalk pubescent (covered with minute white hairs) at least over the lower half; usually
growing in tufts or clusters . 55
54. Not as above; gills darker or more widely spaced or hairs on stalk brown to gray or tawny or
cap distinctly wrinkled, etc. 57
55. Stalk reddish to reddish-brown or vinaceous-brown beneath the pubescence (may fade slightly
in age), 2-6 mm thick; found on ground or wood but not normally on lawns .56
55. Stalk buff or whitish (or pale brown toward base), (2) 5-10 mm thick; growing on rotten wood,
wood chips, or lawns; not common . C. luxurians(see M. oreades, p. 208)
56. Growing in compact bundles on rotting conifers, the wood often buried or decomposed (see
Color Plate 49) . C. acervata, p. 215
56. Found on ground under both hardwoods and conifers, often clustered but not in compact
bundles; base of stalk often with a litter-binding mycelial mat. C. confluens, p. 213
57. Stalk with an enlarged, spongy base; cap and stalk reddish-brown to tan; restricted to eastern
North America . C. spongiosa
57. Not as above . 58
58. Gills white or pallid .C. spp. (unidentified) (see C. confluens, p. 213)
58. Gills soon darker (but may be dusted white by spores) .C. fuse op urp urea group, p. 214
59. Gills usually adnate to decurrent; odor usually mild; on burnt ground, moss, etc., but not
normally in Sphagnum .(see Myxomphalia maura & others, p. 165)
59. Gills usually adnexed to adnate; often in Sphagnum bogs, or if not then odor often rank or
rancid .(see LyophyllumSt Allies, p. 173)
206 TRICHOLOMATACEAE

Marasmiellus candidus Color Plate 36

CAP 0.6 -2.5 cm broad, convex to plane or with slightly depressed center; surface dry,
shining white or translucent white, but often stained deep pinkish or reddish in old age;
often striate or grooved at maturity. Flesh very thin, pliant, soft, odor mild. GILLS fewand
far between, usually interspersed with smaller gills or veins; adnexed or adnate to slightly
decurrent, white like the cap but often pinkish- or reddish-stained in old age. STALK 0.5-3
cm long, 1 -2 mm thick, equal or slightly tapered at either end, often rather short, central or
off-center but not lateral, tough, smooth, often curved, white or with gray to pinkish-gray
base, darkening gradually to brownish-black from the base upward as it ages. SPORE
PRINT white; spores 10-15 * 3.5-6 microns, spindle-shaped to elongated tear-shaped,
smooth, not amyloid.
HABITAT: In groups or rows on dead sticks, branches, berry canes, etc.; widely distri¬
buted but most abundant along the Pacific Coast. It fruits in wet weather, mainly in the
fall and early winter in our area. It is especially abundant along creeks overgrown with
brambles, and on rotting oak, eucalyptus, bay laurel, cedar, etc.
EDIBILITY: Utterly inconsequential.
COMMENTS: Also known asMarasmius candidus, M. magnisporus, and Marasmiellus
albuscorticis, this dainty mushroom is reminiscent of a small shell. Although small, its
shining white cap stands out vividly in the forest gloom. The exceedingly well-spaced (dis¬
tant) gills are its outstanding feature. The stem, which may be off-center, is tough and dark¬
ens at maturity. The entire fruiting body may develop pinkish or sordid vinaceous tones as
it ages, leading one to falsely (but reasonably) assume that the spores are pink. In eastern
North America it is largely replaced by the equally beautiful “pinwheel Marasmius
Marasmius rotula. This little gem has a longer (1.5-8 cm), central black stem that is 1 -2 mm
thick and a white cap (1 -2 cm broad) with darker center. It grows on decaying hardwoods,
usually in large groups. Other whitish-capped eastern species include: Marasmius
delectans, growing on leaves, stalk pallid to yellowish above and dark brown below; and
Marasmiellus nigripes, stalk black beneath a coating of minute white hairs and spores
triangular or jack-shaped. Another whitish-capped easterner, Marasmius epiphyllus,
grows on sticks and leaves and has widely spaced, veined gills and a relatively long, hairlike
stem; it also occurs in the West, as does its oak-leaf-inhabiting look-alike, M. querco-
phyllus. All of the above species are too small to be of culinary value.

Marasmius sp. (unidentified) (Pine Needle Pinwheel)

CAP 1.5^1.5 cm broad, broadly convex to plane or umbilicate; surface not viscid, smooth
or wrinkled, translucent-striate when moist, pure white or with a grayish to olive-gray
tinge, sometimes becoming slightly yellowish in age; margin often wavy. Flesh very thin,
fragile, soft; odor mild. GILLS adnate to slightly decurrent, widely spaced, usually with
veins in between, white. STALK 3-7 cm long, 2-5 mm thick, equal or tapered below or
with a swollen base; often flattened, smooth, colored like cap or slightly yellower, the base
often with brownish stains; hollow, usually with hairs at the base. SPORE PRINT white;
spores 10-12 * 4-6 microns, elliptical, smooth, not amyloid. Cap cuticle cellular.
HABITAT: Scattered to densely gregarious or in troops on pine needles, often abundant
in our coastal pine forests in the late fall, winter, and spring.
EDIBILITY: Unknown, but much too miniscule to be of value.
COMMENTS: The widely spaced, adnate to decurrent gills, pale color, small size, fre¬
quently umbilicate cap, white spores, and growth under pine distinguish this pretty little
mushroom. It looks like a pint-sized Clitocybe or an Omphalina or Camarophyllus, and
is keyed out under those genera. Actually, the term “unidentified” is not quite appropriate
This dainty undescribed Marasmius is a common feature of our coastal pine forests. Note how widely
spaced the gills are.

because it has been “identified” by Dennis Desjardin, Rolf Singer, Howard Bigelow,
and other mycologists as an undescribed species. It is not as tough as most Marasmius
species and does not revive when moistened, but is assigned here to Marasmius for lack
of a better alternative. According to Desjardin, it may be the prototype for a “new” genus.

Marasmius copelandi (Garlic Mushroom)

CAPO.5-2 (2.5) cm broad, convex to plane or centrally depressed; surface smooth or often
wrinkled or striate, light brown to buff or flesh-colored, sometimes fading to whitish as it
dries; not viscid. Flesh thin, whitish; odor distinctly garlic- or onionlike; taste garlicky to
slightly acrid. GILLS pallid to flesh-colored or colored like the cap, attached (usually
adnate or notched, sometimes seceding), often somewhat crisped. STALK 2-7 cm long,
1-3 mm thick, equal or slightly thicker at either end, tough, hollow, minutely hairy; dark
purple-brown to reddish-brown with a paler apex, the base often blackish-brown (but
the hairs may appear whitish when dry). SPORE PRINT white; spores 12.5-16 *
microns, more or less narrowly pip-shaped, smooth, not amyloid.
HABIT AT: Scattered to gregarious on fallen leaves; common along the west coast. In our
area it is often abundant on tanoak leaves and chinquapin burrs in the fall and winter. A
large-spored variant occurs farther north on the leaves of salal and other shrubs.
EDIBILITY: Edible. It can be used as a seasoning or garlic substitute, but should be
cooked only slightly if at all. It makes up for its small size by fruiting in large numbers.
COMMENTS: True to its name, this species and its close relatives smell and taste
like garlic. In fact, they are often smelled before they are seen. Aside from the odor,

Marasmius copelandi is one of several small brownish species with a strong garlic- or onionlike odor.
It has passed under several names, but marasmiologist D. Desjardin says that M. copelandi is correct.

s -
208 TRICHOLOMATACEAE

there is little else to separate them from other“LBM,s.” The mycelium must also smell like
garlic, because in wet weather our tanoak humus will often have a distinct garlic odor—
even when no fruiting bodies are present! There are several very similar “garlic mush¬
rooms,” including: M. olidus, found on oak leaves in eastern North America, with slightly
shorter spores; M. prasiosmus, a European species with whitish gills and even smaller
spores (of uncertain occurrence in North America); M. scorodonius, widespread, with a
reddish-brown to pallid cap and smooth (hairless) stem, found on needles, twigs, grass
stems, etc.; M. alliaceus, a European species with a long black, minutely hairy stalk; and
M. thujinus, with a minute (1-3 mm broad) cap and garlic odor if crushed, found under
northern conifers. Several of these names have been applied to our local garlic mushroom,
but marasmiologist Dennis Desjardin says that M. copelandi is the correct name.

Marasmius androsaceus (Horsehair Fungus)

CAP 2-10 (20) mm broad, convex to plane, the center often depressed; surface dry, soon
radially wrinkled or striate, reddish-brown to pale brown or flesh-colored, fading in age.
Flesh very thin, pliant, reviving when moistened; odor mild. GILLS narrow, well-spaced,
pallid becoming flesh-colored or brownish, usually adnexed to adnate. STALK 2-7 cm
long, less than 1 mm thick, equal, hairlike, tough, stiff, entirely black or black with a brown
to reddish-brown apex; black horsehair-like rhizomorphs usually emanating from base or
visible in surroundings. SPORE PRINT white; spores 6-9 x 2.5-4.5 microns, elliptical or
pip-shaped, smooth, not amyloid. Cystidia present on gill edges.
HABITAT: Scattered or in troops on needles, twigs, or leaves; widespread. In our area
it is fairly common, along with similar species (see below) in wet weather.
EDIBILITY: Unknown—hardly worth the trouble to find out.
COMMENTS: Several dainty marasmioid fungi will more or less fit the above description.
They are barely visible when shrivelled up, but rain revives them. They differ from the more
numerous Mycenas in their dark, hairlike stem and tougher texture. M. androsaceus and
M. pallidocephalus (very similar, but partial to conifer needles and lacking cystidia on the
gill edges) usually possess black rhizomorphs, and sometimes form stems with no caps.
Micromphale sequoiae is similar but grows only on redwood needles, has a brown stalk,
and lacks thick rhizomorphs; likewise Micromphale perforans, which grows on spruce and
fir needles. Micromphale arbuticola, found in swarms at the bases of madrones, is a minute
brownish bark-inhabitor with a slight garlic-onion odor; Marasmius capillar is, partial to
dead leaves in eastern North America, has a black stalk and minute brownish cap with a
white center. See also M. epiphyllus and M. quercophyllus under Marasmiellus candidus.

Marasmius oreades (Fairy Ring Mushroom) Color Plates 38, 47

CAP 1 -5 (6) cm broad but usually 2-4 cm; at first bell-shaped or umbonate with an incurved
margin, then convex or plane but often retainingan obtuse umbo, the margin often uplifted
in old age; surface smooth, dry, color variable: reddish-tan to light brown, tan, buff, or even
white; margin faintly striate when moist. Flesh tough, pliant, pallid, reviving when
moistened; odor agreeable. GILLS adnate, adnexed, or free, fairly well-spaced, broad,
white to pale tan, sometimes discoloring brownish in old age. STALK 2-8 cm long, 1.5-6
mm thick, equal or tapering downward, tough and pliant, smooth, colored like the cap or
paler (whitish). SPORE PRINT white; spores 7-10 x 4-6 microns, elliptical to somewhat
irregular, apiculate, smooth, not amyloid.

HABITAT: Gregarious in grass, usually in arcs or rings; widely distributed and very
common in lawns, parks, cemeteries, etc.; also common in pastures in the Pacific North¬
west, but rarely straying far from suburbia in our area. Found year-round except during
MARASMIUS 209

cold spells, but most abundant in California in late spring, summer (on watered lawns),
and fall. Several crops are produced each year, but its presence can be detected even when it
isn’t fruiting—just look for “fairy rings” (patches of brown grass rimmed by a lush zone of
darker green grass). The living mycelium on the periphery of the ring stimulates the grass to
grow, while the dried-up mycelial matter within the circle inhibits growth.
EDIBILITY: Delectably delicious—one of the few “LBM’s” worth learning. What it
lacks in substance it makes up for in abundance. Discard the tough stems and use the caps
whole. They’re superb is just about anything—omelets, soups, sauces, stir-fried dishes,
even cookies. Or simply saute in butter and serve on toast! What’s more, they dry easily,
don’t decay quickly, and are usually free of maggots. Don’t pass up shrivelled, sun-dried
specimens—they are easily resurrected or can be stored in an airtight jar for later use.
COMMENTS: At first glance this seems like yet another Boring Ubiquitous Mushroom
(“BUM”). However, a closer look reveals that it is really quite attractive, with a lean, clean,
subtle symmetry all its own. Many “BUM’s” and “LBM’s” grow on lawns, but the fairy
ring mushroom can be distinguished by the following features: (1) cap obtusely umbonate
in many specimens (2) white spores (the grass beneath mature caps is often dusted with
white spore powder) (3) broad (deep) white to buff gills which are fairly well-spaced and
not decurrent (4) thin, tough stem (5) growth in grass (6) the ability of dried specimens to
revive dramatically when moistened. Be especially careful not to confuse it with the
poisonous Clitocybe dealbata, which is white-spored and grows in grass (often with M.
oreades), but has thin, crowded, adnate to decurrent gills and a convex to plane(not umbo¬
nate) cap. The growth in rings is not a good means of distinguishing it, as many mushrooms
are capable of growing in circles, including C. dealbata. Other species: M. albogriseus is
fairly common in central and southern California under trees, shrubs, and chaparral or
even in grass. It tends to have a grayer or browner or yellower cap than M. oreades, at least
at the center, and a hollow stem (the stalk of M. oreades is usually stuffed with a white pith),
but is otherwise quite similar. M. (-Collybia) strictipes of eastern North America is also
somewhat similar but grows in the woods and has closer gills and a yellowish cap. Another
woodland easterner, M. nigrodiscus, is larger (cap up to 11 cm broad) and often has a
striate stalk, but is similar in color to M. albogriseus. Collybia luxurians sometimes grows
in grass but has close gills and a reddish-brown cap. Finally, there is an unidentified Mycena
that often grows in grass. However, it is smaller than M. oreades, its stalk is not as tough,
it is usually reddish- or vinaceous-tinged, and its cap is convex to plane (not umbonate).

Marasmius plicatulus (Pleated Marasmius) Color Plate 45

CAP 1-4(5) cm broad, obtusely conical to bell-shaped, often expanding to convex or plane
or with uplifted margin in age; surface dry, with a velvety or frosted appearance when fresh,
furrowed or wrinkled in age or upon drying; bay-brown to reddish-brown, brown, wine-
red, or maroon. Flesh thin, pliant; odor mild. GILLS adnate to nearly free, well-spaced,
broad, white to buff to pinkish or tinged cap color. STALK 5-13 cm long, 1.5-3 mm
thick, equal, tough but brittle, usually long and thin, smooth, polished, reddish-black to
deep chestnut below, often paler (pinkish or sometimes pallid) above; base often with
whitish mycelium. SPOREPRINT white; spores 11-15 x 5-6.5 microns,elliptical,smooth,
not amyloid.
HABITAT: Widely scattered to gregarious in humus under trees and shrubs; apparently
endemic to the west coast. Common in our area in the late fall and winter(at least one month
after the rainy season begins) under eucalyptus, oak, conifers, in brambles, etc.
EDIBILITY: Like myself, too tough and thin to be edible.
COMMENTS: One of the most exquisite of all mushrooms—the frosted wine-red to
brown cap, widely spaced (distant) pallid gills, and long, shining reddish-black stalk are
Left: Marasmius plicatulus is one of our most beautiful mushrooms. See color plate for close-up of
gills. Right: Marasmius haematocephalus, a gorgeous tropical species (see comments below). M.
siccus of eastern North America (not illustrated) closely resembles it. (Michael Fogden)

distinctive. The stalk is so brittle that the mushroom must be dug up (rather than plucked)
to keep it intact. The conical cap may lead to confusion with Mycena, but the tough,
polished stem is characteristic of Marasmius. There are several similar and equally
exquisite species with distant gills, including: M. bellipes, with smaller spores and a small
cap (up to 15 mm broad) and short stalk; M. borealis, whose cap is not pleated or striate;
M. siccus and M.fulvoferrugineus of eastern North America, with even more distant gills,
a shorter (2-7 cm) stalk, an orange-brown to ochre-tawny to rusty-brown, deeply ribbed
or pleated cap (like a miniature umbrella) and spores 15-21 microns long (I have seen the
oranger of the two, M. siccus, preserved nicely in plastic cubes); and M. haematocephalus
(see photo above), a gorgeous tropical leaf-inhabitor with a dark red cap.

Crinipellis piceae
CAP 3-7 (10) mm broad, convex to broadly convex or nearly plane, the center sometimes
slightly depressed; surface whitish to buff or tinged tawny except for the dark (tawny-
brown to brown or blackish) center, which is often surrounded by a dark circle; covered
with coarse tawny to brownish hairs and sometimes minute scales, not viscid; margin often
ciliate (fringed with projecting hairs). Flesh very thin, white. GILLS white, close, adnexed
or free. ST ALK 2-6 cm or more long, up to 1 mm thick, more or less equal, very thin and
tough, brown to blackish-brown beneath a coating of minute hairs. SPORE PRINT white;
spores 7-10 x 3-4.5 microns, cylindrical, smooth, not amyloid. Hairs on cap dextrinoid.
HABITAT: Solitary, widely scattered, or in groups on twigs, needles, and debris of
conifers, especially spruce; known only from the west coast (and Asia). It fruits practically
year-round in damp weather and is, according to Dennis Desjardin, the most numerous
“marasmioid” fungus of the coastal forests of northern California. It does not seem to
occur in our area, but neither does spruce.
EDIBILITY: Much too miniscule to merit attention.
COMMENTS: This minute Marasmius-like mushroom is easily told by the coarse dex¬
trinoid hairs on the cap, frequently ciliate cap margin, and thin, dark stalk. Other species:
C. campanella is a slightly larger northern species with a rusty-orange to chestnut-brown
hairy cap and stem and a tendency to grow on conifer twigs (especially cedar) that are
still on the tree. C. zonata of eastern North America is a “large” (cap 1-2.5 cm) species
with coarse tawny hairs; it grows on dead wood. C. stipitaria has a minute central nipple
on the cap. None of these are worth eating.
CALLISTOSPORIUM 211

Callistosporium luteo-olivaceum
CAP 1.5-6.5 cm broad, convex or slightly umbonate becoming plane or shallowly
depressed; surface not viscid, often minutely scurfy at first but becoming smooth; dark
olive to olive-brown or olive-yellow, often becoming yellower (yellow-brown to honey-
colored) in age and developing dark reddish-brown tones when dried. Flesh thin, pallid
or yellow or tinged cap color; odor mild to pungent or slightly fruity; taste mild or slightly
bitter. GILLS yellow to golden-yellow, tending to redden when dried; close, notched or
adnexed or at times adnate. STALK 2.5-7 cm long, 0.3-1 cm thick, equal or slightly thicker
at either end, often flattened, smooth to fibrillose or scurfy (especially over the lower
portion), sometimes streaked in age; colored like cap or slightly darker, tending to turn
deep reddish-brown from the base upward as it dries. SPORE PRINT white; spores
4.5-6.5 x 3-4.5 microns, elliptical to nearly round, apiculate, smooth, not amyloid but
many of them staining vinaceous in KOH (potassium hydroxide).
HABITAT: Solitary, scattered, or in small groups or tufts on rotten wood (often buried!)
under conifers; widely distributed but not common. I have found it in our coastal pine
forests in the winter. It is said to occur on hardwoods also, particularly in the tropics.
EDIBILITY: Unknown.
COMMENTS: This distinctive mushroom has the stature of a Collybia and was originally
placed in that genus. However, the olive and yellow coloration plus the tendency to grow
on rotten wood (sometimes very decomposed!) distinguish it. C. graminicolor is a similar
but smaller (cap up to 2 cm broad) northwestern species with larger spores.

Strobilurus trullisatus
CAPO. 5-1.5 cm broad, convex to plane or slightly depressed; surface dry, often striate or
wrinkled, minutely granular, white to pinkish-buff or brownish. Flesh very thin. GILLS
typically adnate to adnexed, close, white or tinged pinkish-buff. STALK 2-5 cm long,
1-1.5 mm thick, equal, dry, minutely granular; apex white, lower portion yellowish to
brownish or tawny; base with yellow to tawny-orange hairs and mycelial threads. SPORE
PRINT white; spores 3-6 x 1.5-3 microns, elliptical, smooth, not amyloid.
HABITAT: In colonies (usually 4-10) on old Douglas-fir cones or rarely cones of other
conifers; common throughout the range of Douglas-fir. In our area it usually fruits after
the first fall rains.
EDIBILITY: Who knows? Who cares?
COMMENTS: Also known as S. kemplonae and Colly bia trullisata, this little mushroom
is one of several species that grow only on rotting cones. (Some wood-inhabiting mush¬
rooms, such as Mycenapurpureofusca, may grow on cones but are not restricted to them.)
Other Strobilurus species include: S. conigenoides, a whitish species that grows only on

Strobilurus trullisatus is a nondescript mushroom that grows exclusively on cones. Baeospora

myosura (not illustrated) is rather similar but has very crowded gills.
212 TRICHOLOMATACEAE

the fallen seed pods (“cones”) of magnolias in eastern North America; S. occidentalis,
which occurs on the cones of Sitka spruce; S. albipilatus (-Collybia albipilata), with a
pinkish-buff to brown or dark brown cap, commonintheSierraNevadaandotherwestern
mountains on pine cones and other coniferous debris; and S. lignitilis, which tends grow
on buried decaying wood and has a grayish-brown cap. Finally, there is Baeospora
myosura (-Collybia conigena), which is slightly larger than S', trullisatus and has a smooth,
buff or tan to pinkish-brown cap, very crowded gills, a coarsely hairy stem base, and
amyloid spores. It grows on the cones of various conifers but favors Douglas-fir, at least
in California.

Collybia tuberosa
CAP 3-10 mm broad, convex to plane or centrally depressed; surface smooth, dry, whitish
to buff, sometimes with a darker (yellowish to brownish or pinkish-buff) center. Flesh very
thin, white. GILLS white or rarely tinged pinkish, adnate to adnexed, close or crowded.
STALK 1 -3 cm long, up to 1 mm thick, equal, dry, minutely downy, white or tinged brown,
often arising from a small orange-brown to reddish-brown to blackish, appleseed-like
body or “tuber” (sclerotium). SPORE PRINT white; spores 3-6 * 2-3 microns, elliptical,
smooth, not amyloid.
HABITAT: In colonies on the blackened remains of old mushrooms, particularly larger
Russula and Lactarius species (e.g., R. albonigra), occasionally in humus; widely distri¬
buted. I have found it only once in our area, in December, but the very similar C. cookei
and C. cirrhata (see comments) are fairly common.
EDIBILITY: Unequivocally inconsequential.
COMMENTS: This dainty little Collybia is one of four widespread species that colonize
decayed mushrooms. The host may be so deteriorated, however, that it is not recognizable
as a mushroom. The other three species are: C. cookei, practically identical but with
rounder, more prominent, tan to yellow or yellow-orange sclerotia (and occasionally
found on rotten wood or in humus); C. cirrhata, also very similar but with white mycelial
threads instead of sclerotia, found in humus as well as on old mushrooms (and particularly
common in the Sierra Nevada); and C. racemosa, which has stubby lateral branches on the
stem (see description). All of these are placed in Microcollybia by some taxonomists.

Left: Collybia tuberosa and its close relatives grow in colonies on decaying mushrooms and other
debris. Sclerotia (beadlike bodies from which the mushrooms arise) are not visible in this picture.
Right Collybia racemosa is easily told by its small size and unique branching stalk.
COLLYBIA 213

Collybia racemosa (Branched Collybia)

CAP 3-10 mm broad, bluntly conical to convex becoming umbonate to nearly plane;
surface smooth, not viscid, dark gray to gray or brownish-gray, the margin often paler.
Flesh very thin. GILLS typically adnexed, close, gray or brownish-gray. STALK 3-8 cm
long, 0.5-3 mm thick, with numerous short lateral side-branches, especially over the lower
half or two-thirds; gray to brownish-gray, often entirely buried and sometimes originating
from a small blackish beadlike body (sclerotium). SPORE PRINT white; spores 4-5.5 x
2-3 microns, oblong to elliptical, smooth, not amyloid. Asexual spores (conidia) often
produced on the swollen tips of the side-branches.
HABITAT: In small groups or colonies on old decayed or blackened mushrooms (e.g.,
Russula albonigra) or occasionally in coniferous duff; widely distributed but seldom
encountered, perhaps because it is so easily overlooked. I have found it only once in our
area, in December, but it is said to be fairly common in the Sierra Nevada.
EDIBILITY: Unknown, but much too puny and rare to be of value.
COMMENTS: This curious Collybia is the only one with side-branches on the stem (see
photo on p. 212). The stalk may meander somewhat through its substrate, branching
along the way, and the sclerotium from which it originates is not always evident. Stems
without caps are sometimes found, implying that the formation of asexual spores on the
side-branches can be enough to “satisfy” the mycelium’s reproductive urges.

Collybia confluens (Tufted Collybia)

CAP 2-5 cm broad, convex to plane or slightly umbonate, the margin sometimes uplifted
or wavy in age but incurved at first; surface smooth, hygrophanous: reddish-brown to
pinkish-cinnamon or flesh-colored when moist (often darker at center and pallid at mar¬
gin), fading to pinkish-buff, grayish-pink, or whitish as it dries. Flesh thin, white. GILLS
crowded, narrow, adnate soon becoming adnexed or even free, whitish to flesh-colored.
STALK 3-10 cm long, 2-5 mm thick, equal, hollow, pliant, sometimes flattened or grooved,
tough, usually darker than cap (reddish-brown), but covered with a minute white pubes¬
cence (downy hairs); base often with white mycelial mat attached. SPORE PRINT white
or tinged yellow; spores 7-9 x 3-4 microns, narrowly elliptical, smooth, not amyloid.
HABITAT: Gregarious, often in tufts or clusters, on ground in woods; widely distributed.
In the Pacific Northwest it is quite common under conifers, but in eastern North America it
favors hardwoods. I have not seen it in our area, but similar species (see comments) occur.
EDIBILITY: Edible with caution; it is tough and similar species have not been tested.
COMMENTS: The fine white pubescence on the stalk (use hand lens!) and crowded gills
help to separate this species from C. dryophila and C. butyracea. It shrivels up in dry
weather and revives somewhat when moistened, and as a result was originally placed in
Marasmius. A similar, unidentified species whose gills are adnate to adnexed and not so
crowded and whose stalk lacks the litter-binding mycelial mat is quite common in our area
in the fall and winter; it sometimes rivals C. dryophila for abundance but appears later in
the season and, like C. confluens, it has a minutely downy stem. Another local, downy-
stemmed species appears to be undescribed. It resembles the C.fuscopurpurea group, but
has whitish gills and a pallid to brownish-tan cap. It differs from C. confluens in having
attached gills which are fairly well-spaced, and it usually grows in tufts or clusters on
decaying wood, wood chips, or in lignin-rich humus. A third species, C. polyphylla, has a
garlicky to slightly unpleasant odor and taste, while a fourth, C. peronata (-Marasmius
urens) has an acrid (burning) taste. The latter two species are widely distributed but do
not seem to occur in California.
Collybia fuscopurpurea group. These ubiquitous “LBM’s” have a tough texture and usually grow in
groups or clusters.

Collybia fuscopurpurea group

CAP 1-4 cm broad or slightly larger, convex becoming plane, slightly depressed, or with
the margin uplifted in age; surface dark reddish-brown to brown, purple-brown, or
chocolate-brown when fresh, usually paler (near tan) when dry, usually radially wrinkled
or finely striate. Flesh whitish or colored like cap, thin, reviving somewhat when moistened;
odor mild. GILLS attached (usually notched or adnexed), fairly well-spaced, pallid or pale
pinkish-tan becoming more or less cap color, then dusted whitish by spores. STALK
2-10 cm long, 1-4 mm thick, equal or tapered downward, dry, rather tough and pliant,
brown with the apex usually paler; clothed with minute grayish to brownish hairs (unless
very wet) throughout or over the lower two-thirds; often curved near the litter-binding
base. SPORE PRINT white; spores 6-8 x 3-4 microns, elliptical, smooth, not amyloid.
HABITAT: Densely gregarious (often in clusters) among leaf litter, humus, and woody
debris in woods, under trees (especially oak), or in wood chips or landscaped areas; widely
distributed. Common year-round in our area but particularly abundant in the early fall.
EDIBILITY: Unknown.
COMMENTS: Formerly known as Marasmius fuscopurpureus, this common species
“complex” often forms dense swarms in leaf litter and woody debris, especially where the
ground has been recently disturbed. Like most “LBM’s,” it blends in well with its sur¬
roundings. It can be recognized by the reddish-brown to dark brown fruiting body, convex
to plane cap (never bell-shaped as in Mycenal), white spores, pubescent (finely hairy)
stem, and gregarious disposition. However, separating the numerous species within the
“complex” is a job for specialists. One such specialist, Dennis Desjardin, says that
several of California’s representatives in this “complex” (including the most common
one) appear to be undescribed. There are a number of other difficult-to-identify Collybia
and Marasmius species with a pubescent stalk and litter-binding mycelium. Several of
these (e.g., M. cohaerens) are described in the key to Marasmius and Collybia; others
are discussed under C. confluens.

214
COLLYBIA 215

Collybia acervata (Clustered Collybia) Color Plate 49

CAP 1-4 (5) cm broad, convex with an incurved margin, becoming broadly convex in age;
surface smooth, hygrophanous, not viscid; dark reddish-brown when fresh and moist,
fading to pale reddish-brown, pinkish-buff, or paler (sometimes with darker and lighter
zones) as it dries. Flesh thin, pallid. GILLS close or crowded, narrow, typically adnexed or
notched or free, white to dingy pinkish or vinaceous-buff. STALK 4-12 cm long, 2-6 mm
thick, more or less equal, dry, hollow, pliant but brittle, smooth above, with fine whitish
hairs over lower half or at base; reddish-brown to vinaceous-brown or sometimes paler in
age. SPORE PRINT white; spores 5-7 * 2-3 microns, elliptical, smooth, not amyloid.
HABITAT: In compact bundles or clusters on rotting conifers (but often appearing
terrestrial); widely distributed. It is common in the summer and fall in the Sierra Nevada
and Rocky Mountains and in the Pacific Northwest, but apparently absent in coastal
California south of San Francisco.
EDIBILITY: Inedible. It is said to have a bitter taste when cooked and is apparently
slightly poisonous to some people.
COMMENTS: The bundled growth habit (see color plate!), white spores, and reddish-
brown stem make this an easy mushroom to recognize. It always grows on wood, but may
appear terrestrial if its host is buried or in a very advanced stage of decay. The cap varies
in color according to the amount of moisture present, but is never translucent-striate as
in Mycena. Other species: C. marasmioides (-C. erythropus, C. bresadolae) tends to grow
in tufts or clusters on the ground, but has a pale tan to creamy-buff cap and beautiful
dark red (paler at apex) stem. A mushroom meeting this description occurs in our area
but is rare. Clitocybula (-Collybia) familia has the aspect of C. acervata (densely clustered
growth habit on rotting conifers—see photo on p. 898), but it has a watery white to smoky-
gray to somewhat brownish or tan (never reddish-brown) cap, white to grayish stem,
and round, amyloid spores. It is edible and widely distributed, but in California it
seems to be restricted to the Sierra Nevada. Clitocybula (-Collybia) abundans resembles
C. familia, but has a smaller fibrillose cap and elliptical, amyloid spores.

Collybia dryophila (Common Collybia; Oak-Loving Collybia)

CAP 1-5 (7) cm broad, broadly convex with an incurved margin, becoming plane or with
with an uplifted, often wavy margin in age, sometimes also slightly umbonate; surface
smooth, hygrophanous: chestnut-brown to reddish-brown, yellow-brown, tawny, or
ochre when young and moist, but fading to tan, pinkish-tan, yellowish-tan, or buff as it
dries. Flesh thin, white. GILLS crowded, usually notched or adnexed, white to pale yellow.
STALK 2-8 cm long, 2-6 mm thick, equal or with a swollen base, slender, smooth, hollow,
rather tough and cartilaginous; pale cream or colored like cap (but often paler); white
mycelium often visible at base or in surrounding humus. SPORE PRINT white or pale
cream; spores 5-7 * 2-3.5 microns, elliptical, smooth, neither amyloid nor dextrinoid.
HABITAT: Scattered to gregarious or in small tufts in woods or near trees, often forming
arcs or rings; widely distributed. It is abundant in our area shortly after the first fall rains,
but is less common thereafter. As its name implies {dryophila-oak-loving), it is fond of
oak, but is also common under pine and other conifers, as well as around the edges of old
sawdust piles. At higher elevations it often fruits in the spring as well as in the summer
and fall.
EDIBILITY: Edible, but some people are apparently sensitive to it. Only the caps are
tender enough to eat. It is a proficient concentrator of mercury, but occasional con¬
sumption should pose no threat as mercury is a cumulative poison.
Collybia dryophila is a cosmopolitan “LBM” with white or yellowish gills that are usually notched,
adnexed, or even free. Note the relatively slender equal stalk and small size.

COMMENTS: The appearance of this species in large fairy rings under oaks is a sure sign
that the coastal California mushroom season is under way. The hygrophanous reddish-
brown to tawny, rusty, or tan cap, more or less adnexed gills, growth on the ground, and
smooth (hairless) stalk distinguish it from all but C. butyracea, which has a greasier
(buttery) cap, buff-colored spores, and ragged gill edges. Both species are reminiscent of the
fairy ring mushroom (Marasmius oreades), but grow under trees and have crowded gills;
neither grows in the tight bundles characteristic of C. acervata, and neither has downy hairs
on the stem like C. confluens. In some regions C. dryophila is frequently covered by lumps
or masses of somewhat jelly- or tumorlike tissue, caused by the fungus Christiansenia
mycetophila. However, I have not observed this phenomenon in California. The yellow-
gilled form of C. dryophila is sometimes listed as a separate species, C. sub sulphur ea.

Collybia butyracea (Buttery Collybia)

CAP 3-8 (12) cm broad, convex becoming plane or uplifted, often with a broad umbo;
surface smooth, greasy or slippery when moist but not truly viscid; dark reddish-brown to
chesnut-brown to dull brown (or grayish-brown in one form), fading as it dries or ages to
tan, grayish-tan, reddish-tan, or ochre-buff; margin at first incurved. Flesh thin, soft,
whitish or watery. GILLS close or crowded, free or adnexed, white, the edges usually
uneven, eroded, or finely scalloped at maturity. STALK 2-10 cm long, 4-10 mm thick,
equal or more often thicker below and/ or pinched at the base; often longitudinally striate
or twisted; rather tough and cartilaginous, hollow at least in age, smooth; colored like cap
in age but often buff when young; base usually with white mycelial down. SPORE PRINT
creamy to yellowish, buff, or pinkish-buff; spores 6-8 * 3-3.5 microns, elliptical, smooth,
many of them dextrinoid.
HABITAT: Scattered to gregarious or tufted in humus under conifers (especially pine)
or occasionally hardwoods; widely distributed. It is sometimes common in our coastal
pine forests in the late fall, winter, and early spring.
EDIBILITY: Edible; about like C. dryophila. Care must be taken to identify it correctly!
COMMENTS: Also known as Rhodocollybia butyracea, this handsome and widespread
species is often confused with C. dryophila. However, its cap is apt to be greasier when
moist, the gills are more apt to be finely scalloped or eroded, and the spores are slightly
more colored in deposit and at least somewhat dextrinoid. A grayish-brown version of the
species {form asema) also occurs in North America, especially on the west coast. Other
species: C. badiialba and C. disiorta are two closely related species with round or nearly
round spores, a reddish-brown to vinaceous-brown cap, and white gills that may or may

216
Collybia butyracea is often confused with C. dryophila, but is slightly different in color and slightly
larger, and often has a club-shaped stalk (thicker at base).

not develop reddish stains in age. Both favor conifers and grow in humus or on rotten
wood. The former occurs in coastal California and the Pacific Northwest while the latter
is more common in eastern North America. Another species with a vinaceous-brown cap,
C. extuberans, has elliptical spores and is widely distributed.

Collybia maculala (Spotted Collybia)

CAP (2.5) 4-12 cm broad, convex with an incurved margin at first, becoming broadly
umbonate to plane or undulating; surface dry or lubricous but not truly viscid, smooth,
typically white to buff, pale tan, or pinkish-tinged, often darker or redder in age, especially
at the center and often developing rusty or reddish spots and stains. Flesh thick, white; odor
mild or somewhat fragrant; taste usually bitter (but mild in var. occidentalis). GILLS
white to pale pinkish-buff, often developing rusty or reddish stains (like the cap) in age;
adnate to adnexed or notched to nearly free, crowded. STALK 4-15 cm long, 0.5-2 (3) cm
thick, often tapered below to form a more or less rooting base; solid (at least when young),
firm, smooth or fibrillose, often striate or grooved and easily splitting longitudinally;
white to pale buff, usually developing rusty or reddish stains below. SPORE PRINT
creamy to pale peach or pinkish-buff; spores 5-11 x 4-6 microns, round or nearly round
in the typical variety, elliptical in var. occidentalis; smooth, at least a few dextrinoid.
HABITAT: Solitary, scattered, or in groups or tufts on decaying wood or lignin-rich
humus under conifers; widely distributed. I have not seen it in our area, but it is quite
common in the Pacific Northwest and Sierra Nevada in the late spring, summer, and fall.
EDIBILITY: Inedible because of the frequently bitter taste.
COMMENTS: Also known a.s R hodocollybia maculata, this rather large species is closely
related to C. oregonensis and has the same stature, but typically has a paler cap and mild
odor. The pinkish to rusty-reddish spots and stains and tendency to grow on rotten wood
are good fieldmarks. It could conceivably be mistaken for a Tricholoma, but the stalk is
longer than in most Tricholomas and it splits more easily. It is a variable species, and a
number of varieties have been designated based on differences in color, taste, and spore
shape. The above description embraces several such varieties. Others include: var.
scorzonerea, a large version with yellowish gills and a frequently yellowish-tinged stalk
and/or cap; and two small varieties from the Pacific Northwest with a vinaceous-brown
to blackish cap.

217
Left: Collybia oregonensis is an amazingly fragrant mushroom. Right: Oudemansiella radicata
(p. 219), a common mushroom in eastern North America, is rather variable in color but always has
broad gills (as shown here) and a deeply rooting stalk (not visible).

Collybia oregonensis (Fragrant Collybia)

CAP 4-10 cm broad, broadly convex to plane; surface smooth, slightly viscid when moist
but soon dry; deep chestnut-brown to vinaceous-brown or reddish-brown at the center,
often paler or pinker or redder toward the margin, and gradually fading overall in age.
Flesh thin, white or reddish-stained; odor heavy and sweet (like benzaldehyde), taste
somewhat bitter. GILLS crowded, creamy or pale yellow to buff, often reddish-stained in
age; adnexed, notched, or seceding; edges sometimes overlapping each other and eroded in
age. STALK 6-20 (30) cm long, (0.5) 0.8-2 cm thick, usually quite long and deeply rooted;
equal above or swollen slightly near ground level, tapered gradually below to a point; dry,
whitish, but usually developing reddish to reddish-brown stains especially over the lower
portion; solid or hollow in age. SPORE PRINT whitish to buff; spores 6-8 x 3.5-5 microns,
elliptical, smooth, at least some of them dextrinoid.
HABITAT: Solitary or in small groups around old stumps and in lignin-rich humus;
apparently endemic to the Pacific Coast, not common. I’ve seen it in the Sierra Nevada,
and in one local spot (in mixed woods) where it fruits every year, usually in the fall.

EDIBILITY: Unknown. Despite its sweet odor it has a somewhat bitter taste.
COMMENTS: The strong, heavy benzaldehyde or almond extract odor and vinaceous-
brown cap are distinctive. The fragrance is reminiscent of Russula fragrantissima, but
without the fetid component of that species. The stalk often roots deeply but does not form
a true pseudorhiza (“tap root”) as in Caulorhiza umbonata. The gills are not waxy as in the
almond-smelling Hygrophorus species (e.g., H. bakerensis) and the odor is usually thicker
and more pronounced. C. subsulcatipes, known only from Washington, has a similar but
weaker odor, a brown to dark vinaceous-brown stalk with a “tap root,” a brown to vina-
ceous-buff cap, and dull vinaceous gills in age. Both of these species are placed in a separate
genus, Rhodocollybia, by some collybiologists.

Caulorhiza umbonata (Redwood Rooter)

CAP 3-15 cm broad, at first conical or umbonate, then expanding but usually retaining
the umbo; surface smooth, dry, chestnut-brown to warm tan or yellow-brown; margin at
first incurved. Flesh thin, pallid; odor mild. GILLS notched or adnexed to nearly free,
close, broad, white to yellowish or tinged cap color. STALK 6-50 cm long or more, 0.4-1.5

218
Caulorhiza (- Collybia) umbonata is a flagrant fungal feature of our redwood forests. Note the conical
or umbonate cap and long “tap root” which may extend as much as three feet into the humus.

(2) cm thick, most of it underground in the form of a long, tapered “tap root”; smooth,
twisted-striate, buff to tan or colored like cap but paler; rather tough and cartilaginous.
SPORE PRINT white; spores 5-8 * 3-5 microns, elliptical, smooth, amyloid.
HABITAT: Solitary, scattered, or in groups near or under redwood (rarely elsewhere);
restricted to the range of coastal redwood (Sequoia sempervirens). It is very common in
our area in the fall and winter.
EDIBILITY: Like myself, not firmly established. H owever, if it were poisonous we would
probably know by now. I’ve sampled small quantities without ill effects.
COMMENTS: Anchored by its long “tap root,” this distinctive species, like a dandelion,
resists being uprooted. The “tap root,” pallid gills, umbonate cap, and white spore print,
plus the association with redwood distinguish it. The “tap root” is brittle, however, and
will break off and stay behind in the ground unless it is dug up very carefully. Collybia
umbonata is a synonym. Collybia subsulcatipes (see comments under C. oregonensis) is
somewhat similar but has a sweet odor. Caulorhiza hygrophoroides is a closely related
hardwood-lover with a smaller (up to 5 cm) cap. None of these have the grayish-brown
colors and exceptionally broad gills of Oudemansiella radicata and its relatives.

Oudemansiella radicata (Beech Rooter)

CAP 2.5-12 cm broad, bell-shaped becoming broadly convex to plane or with a broad
umbo, the margin sometimes uplifted in old age; surface smooth or radially wrinkled, often
viscid or tacky when moist, usually dark brown to grayish-brown, but in some variants
grayish and in others whitish, blackish, or olive- to yellowish-brown; margin at first in¬
curved. Flesh thin, whitish; taste mild. GILLS broad, fairly well-spaced, thickish, usually
adnexed or notched; white. STALK 5-25 cm long,0.3-1 (1,5)cm thick, usually thickest at
or near ground level, with a tapered underground portion or “tap root”; whitish at apex
or in upper portion, colored more or less like the cap below, dry, rather stiff and brittle,
longitudinally lined or twisted-striate; smooth or in one form scurfy. SPORE PRINT
white; spores 12-18 * 9-12 microns, broadly elliptical, smooth, not amyloid.
HABITAT: Solitary, scattered, or in small groups on or near hardwood stumps and roots
(often appearing terrestrial) in woods, grassy clearings, etc.; fairly common in eastern
North America, spring through fall (especially on beech), apparently absent in the West.
EDIBILITY: Edible but not choice—or so I am told. Only the caps are worth eating.

219
220 TRICHOLOMATACEAE

COMMENTS: Formerly known as Collybia radicata, this common eastern mushroom

can be told by its broad white, well-separated gills (see photo on p. 218) and rooting stalk.
The latter snaps easily, however, so that if it is not dug up carefully the telltale “tap root”
will remain behind in the ground. The color of the cap is quite variable, but is usually in
the dark brown to grayish-brown range. The viscidity of the cap is not always evident.
Tricholomopsisplatyphylla is somewhat similar but does not normally have a “tap root,”
while Melanoleuca species are terrestrial and have narrower, crowded gills. Other species:
O. longipes is a similarly colored but slightly smaller species with a dry cap and much
smaller spores; it occurs in the Rocky Mountains on hardwoods such as aspen.

Flammulina velutipes (Velvet Foot; Velvet Stem) Color Plate 26

CAP 1-5 (7) cm broad, convex to plane or broadly umbonate; surface smooth, slimy or
viscid when moist, reddish-brown to yellow-brown, yellow-orange, orange-brown, or
tawny, the margin often paler (yellower); fading in dry weather; margin at first incurved.
Flesh thin, white or yellowish. GILLS adnate to adnexed or notched, white to pale yellow,
close. STALK 2-11 cm long, 3-5(12) mm thick, equal or thicker below, slender, tough, of¬
ten curved, sometimes slightly off-center; smooth and pallid to yellowish to orange-brown
when young, but developing a rusty-brown to blackish-brown velvety pubescence (tiny
hairs) from the base upward as it matures. SPORE PRINT white; spores 6.5-9 * 3-5
microns, elliptical to pip-shaped, smooth, not amyloid. Gills with cystidia.
HABITAT: In tufts or clusters on or near stumps, logs, and roots of hardwoods (but some¬
times appearing terrestrial); very widely distributed. It is fairly common in our area from
fall through spring on poplar and willow, and in coastal sand dunes on bush lupine. In the
Rocky Mountains it is abundant in the late summer on aspen; in eastern North America it
fruits from the late fall through early spring on elm and other hardwoods.
EDIBILITY: Edible, but the sticky skin should be removed before cooking. In colder
regions it is an important edible and is called the ‘winter mushroom” because it fruits very
late in the season (even during winter thaws) when other fungi are not available. In our
balmy climate, however, its season coincides with that of other, more flavorful mushrooms,
so it is not often gathered. A cultivated form of it called the “snow-puff mushroom” or
enokitake can be bought in many markets. It looks something like a pure white bean sprout
with its long, smooth (not velvety!) stem and negligible cap.
COMMENTS: The smooth, sticky, yellow-orange tobrownishcapandstalk whichisdark
brown and velvety (at least below) at maturity plus the white spores, absence of a veil,
and clustered growth habit on dead hardwoods typify this hardy mushroom. It was origi¬
nally placed in Collybia, but is now given a genus of its own due to its viscid cap and the
prominent sterile cells (cystidia) on the gills. In arid climates, however, the cap may appear
dry rather than viscid. The fruiting bodies can apparently survive freezing, and after
thawing out will continue to produce spores!

Flammulina velutipes is easily told by velvety brown stalk, sticky cap, cheerful color (see color plate),
and growth on wood.
 221

OMPHALINA& XEROMPHALINA
Small to minute, often brightly colored, saprophytic mushrooms. CAP usually plane to depressed
or umbilicate at maturity, not conical; margin often incurved when young. GILLS often yellow,
orange, or pinkish, typically adnate to decurrent. ST ALK usually central, thin, cartilaginous, often
pliant or rather tough, hollow. VEIL and VOLVA absent. SPORE PRINT white to pale yellow.
Spores smooth, amyloid (Xeromphalina) or not amyloid (Omphalina).

THESE are dainty, brightly colored mushrooms with decurrent gills and a cartilaginous
stem. They were originally grouped together in the obsolete genus Omphalia, but
Xeromphalina was created for the species with a dark stem, tougher texture, ability to
revive somewhat when moistened, and amyloid spores.
Omphalina is interesting from a botanical standpoint because several species grow only
with lichens. It is now thought that the mushrooms are actually the fruiting bodies of the
fungal component of the lichen (a lichen is a symbiotic relationship between an alga and a
fungus). Other Omphalinas grow in grass, moss, or on soggy logs. They are easily confused
with Mycenas (which usually have a conical or bell-shaped cap), while even experts cannot
agree on the distinction between Omphalina and some of the small, slender Clitocybes.
Xeromphalinas occur on logs and in humus and are more common than Omphalinas,
at least in California. Their tough, usually dark stem and ability to revive after being dried
are reminiscent of Marasmius, but their yellow to orange gills and amyloid spores are
distinctive. In addition to Xeromphalina and Omphalina, the genus Gerronema is recog¬
nized by some mycologists; it embraces several Omphalinas not specifically associated with
lichens. None of these mushrooms are large enough to eat. If your mushroom does not key
out convincingly below, check Mycena, Marasmius, and the Hygrophoraceae.

Key to Omphalina & Xeromphalina

1. Stalk tough, wiry, or horny, at least the lower portion dark brown to orange-brown or reddish-
brown when mature . 2
1. Not as above . 5
2. Typically growing on wood (sometimes buried), often in clusters or dense groups .3
2. Typically growing on ground or humus, usually scattered to gregarious but not clustered . 4
3. Cap less than 2.5 cm broad; growing on conifers .X. campanella, p. 222
3. Cap typically at least 2 cm broad; cap velvety, more or less orange-brown; growing on hard¬
woods in eastern North America and the tropics .X. tenuipes
4. Taste bitter; gills adnate .X.fulvipes{see X. cauticinalis, p. 222)
4. Taste more or less mild; gills adnate to decurrent .X. cauticinalis, p. 222
5. Cap and stalk slimy-viscid when moist, more or less yellow . . {setMycena lilacifolia, p. 236)
5. Not as above . 6
6. Fruiting body orange (or at times pinkish-orange or fading to yellow or orange-buff); gills well¬
spaced and waxy-looking; found in groups or clusters on rotting conifers, especially in the
Pacific Northwest and California .O. luteicolor{see O. ericetorum, p. 223)
6. Not as above . 7
7. Cap and/ or gills greenish-yellow to green, olive, olive-brown, or dark green; growing on rotting
conifers or on ground, moss, or lichens . O. wynniae& others (see Clitocybe odor a, p. 161)
7. Not as above; green shades absent. 8
8. Cap 0.5^4 cm broad, ochre to grayish-ochre; gills yellow to orange-yellow; growing on rotting
conifers; spore print often yellow- or salmon-tinged .O. chrysophylla
8. Not as above; cap usually less than 2.5 cm broad .9
9. Cap minute, up to 1.5 cm broad, yellow-orange to orange or orange-red; stalk orange; gills pallid
or yellowish; growing in moss .O. (-Rickenella ox Mycena) fibula
9. Not as above; differently colored or larger or growing elsewhere . 10
10. Cap grayish to gray-brown; gills often yellowish (especially in age); found on rotten wood in
eastern North America .O. strombodes
10. Not as above ..O. ericetorum & others, p. 223
Xeromphalina fulvipes (see comments under X. cauticinalis) forms carpets of miniature fruiting
bodies under redwood and other conifers. Gills are typically adnate and taste is bitter.

Xeromphalina cauticinalis
CAP 0.5 -2.5 cm broad, convex becoming plane or with a small central depression; surface
smooth, not viscid, reddish-brown to tawny or ochraceous-tawny, fading to yellowish.
Flesh very thin, pliant; taste mild. GILLS adnate to decurrent, yellow, with veins between.
STALK 2-8 cm long, 1-2.5 mm thick, equal or more often with a small bulb at the base;
pliant, tough, tawny or yellowish above, dark brown below; base with tawny mycelium.
SPORE PRINT white; spores 5.5-7 x 3-4 microns, elliptical, smooth, amyloid.
HABITAT: Scattered to densely gregarious on conifer needles and debris; common in
western North America. In our area it appears (along with X. fulvipes—see comments) in
the late fall and winter under redwood and other conifers.
EDIBILITY: Unknown.
COMMENTS: This is one of several terrestrial Xeromphalinas. The decurrent yellow gills
distinguish it from Mycena, and the stalk is much tougher than in Hygrocybe. Other
species: X. fulvipes is similar but has a bitter taste, adnate gills, and minute orange hairs on
the stem. I have found it under redwood several times, but it is not as common as X.
cauticinalis;X. picta is a minute terrestrial species with a greatly swollen stalk apex.

Xeromphalina campanella Color Plate 44

CAP 0.3 -2.5 cm broad, convex becoming broadly convex with a depressed center; surface
smooth, not viscid, yellow-brown to tawny to orange-brown or cinnamon-brown; margin
striate when moist. Flesh thin, pliant, yellowish. GILLS yellowish to dull orange, fairly
well-spaced, decurrent with veins in between. STALK 1-5 cmlong,0.5-3 mm thick, equal
or enlarged at base, pliant, tough, smooth, horny, often polished; yellowish above, brown
to reddish-brown below; usually curved; base with bright tawny hairs. SPORE PRINT
white to pale buff; spores 5-8 x 3-4 microns, elliptical, smooth, amyloid.
HABITAT: In groups or dense clusters on rotting conifers; widely distributed. It is
common throughout the West, fruiting in our area in the fall, winter, and early spring.
EDIBILTY: A miniscule morsel that is hardly worth eating.

222
OMPHALINA & XEROMPHALINA 223

COMMENTS: The yellow to orange, decurrent gills plus the thin polished stem and small
size typify this dainty mushroom. Its occurrence on conifers is so dependable that its pre¬
sence can be taken as “proof’ that its host is some type of conifer. It might be mistaken
for a Mycena, but the shape of the mature cap is quite different. Galerina autumnalis
is also somewhat similar, but has a veil and brown spores. The cluster pictured in the color
plate is darker (browner) than normal, but shows the shape and growth habit quite well.
Other species: X. kauffmanii is very similar but occurs with hardwoods in eastern North
America; X. orickiana has a dark reddish-brown cap, grayish to brownish gills, and grows
grows on redwood.

Omphalina ericetorum (Lichen Agaric)

CAP 0.5-2.5 (3.5) cm broad, at first plane with an incurved margin, becoming deeply
depressed or funnel-shaped in age; surface smooth, not viscid, dull cinnamon to brownish,
fading to yellowish or straw color or paler as it ages; margin striate, often wavy. Flesh very
thin, pliant. GILLS decurrent, well-spaced, sometimes veined, pale yellowish. STALK
1-3 cm long, 1-3 mm thick, equal or enlarged at base, often curved, smooth, pliant, pale
reddish-brown above, yellow-brown to pale brown below; often pale yellowish in age.
SPORE PRINT white to yellow; spores 7-9 ><4-6 microns, elliptical, smooth, not amyloid.
HABITAT: Usually in groups on old lichen-laden conifer logs or scum-covered soil,
always associated with the lichen Botrydina vulgaris; widely distributed in the cool tem¬
perate zone, but rare in our area. It is one of the commonest mushrooms of the Arctic.
EDIBILITY: Much too puny to be of value.
COMMENTS: The small size, depressed cap, thin stem, decurrent gills, and distinctive
habitat are the principal fieldmarks. The gills may appear somewhat waxy as in Hygrocybe
and Camarophyllus, but the latter don’t normally grow with lichens. O. umbellifera is a
synonym. There are several similar species, including: O. hudsoniana, similarly colored
but associated with the lichen Coriscium viride;0. postii, with a pinkish-red to orange cap,
found in moss; O. pyxidata(scc photo below), fairly common in our area in grassy or open
places, with pinkish- or vinaceous-tinged gills that fade to yellowish or creamy in age; and
O. luteicolor (COLOR PLATE 24), a beautiful waxy-gilled orange to salmon-colored
species that fades with age and is common in the Pacific Northwest, northern California,
and the Sierra Nevada on rotting conifers, usually in groups or clusters.

Two small Omphalinas with different habitats (both are discussed above). Left: Omphalina erice¬
torum is associated with lichens (which, however, are often very inconspicuous). Right: Omphalina
pyxidata grows in moss, grass, and soil. Note how both species have decurrent gills. Similar greenish
species are discussed under Clitocybe odora (p. 162).
224 TRICHOLOMATACEAE

MYCENA
Small to minute saprophytic mushrooms that do not revive when moistened (after being dried).
CAP typically conical to bell-shaped when young, but often expanding in age; often translucent-
striate; margin usually straight when young, rarely incurved. GILLS usually attached. STALK
central, thin, usually hollow;fragile or cartilaginous. VEIL and VOLVA absent. SPORE PRINT
white. Spores smooth, amyloid or not amyloid. Cystidia usually present on gills.

THIS is a very large group of very small mushrooms with a thin, fragile or cartilaginous
stem and bell-shaped to conical cap (at least when young). The gills are not waxy as in the
Hygrophoraceae; the dried fruiting body does not revive when moistened as in Marasmius,
and is not particularly tough; Collybia and Omphalina are similar but usually have a
convex to plane or umbilicate cap. Other small mushrooms with conical to bell-shaped
caps (Coprinus, Conocybe, Nolanea, etc J do not have white spores.
In sheer numbers Mycenas are more abundant than any other mushrooms, but because
of their diminutive dimensions, most people are oblivious to their presence. In the wake
of heavy rains they fruit in untold quantity in the woods, especially on needle beds under
conifers, where they form thick carpets of delicate domes. They are strictly saprophytic—
on logs, stumps, sticks, leaves, soil, and humus. They turn up occasionally in lawns,
gardens, and flower pots, but do not grow in dung like Coprinus and Bolbitius.
The most common forms are gray or brown, but a few, such as M. acicula, are brightly
colored. Alexander Smith’s monumental monograph lists 218 North American species,
but dozens more occur. The overwhelming majority cannot be identified positively without
a microscope (some are so small they can barely be seen with the naked eye!). The size and
shape of the sterile cells (cystidia) on the gills plays a particularly important role in their
identification. However, Mycena can be divided into manageable groups based on gross
features such as size, color, odor, habitat, and viscidity. In addition, a few species exude a
juice or latex when the base of the stem is broken and squeezed. Lactarius species also
possess a latex, but are much larger and fleshier with thicker stems.
In every mixed bag of individuals I take mushroom hunting, there’s always one or two
with a keen eye for detail. I n the normal course of events, I begin with a brief spiel about the
marvels of the mushroom world before dispatching the group to rush about madly in
search of the elusive 25-lb. Boletus edulis (trampling myriad Mycenas with every step),
while I make a sly beeline for my secret Boletus patch. These one or two keen-eyed
individuals, however, are content to remain where they are, meticulously examining every
leaf, twig, and cone, and uncovering in the process not only a multitude of marvelously
minute Mycenas, but an astonishing assortment of other clandestine creatures—slugs,
centipedes, spiders, snakes, salamanders, etc.
As I am rather small myself, I harbor a profound respect for these exceptional indivi¬
duals. In a society where we are taught from birth to think big, it is encouraging to find
some who are still able to make the distinction between quality and quantity, who appre¬
ciate the fact that size alone is not a measure of intrinsic worth.
My lust satiated and my basket laden with Boletus, I return later to find these patient
and perceptive souls sprawled out on their bellies in the exact spot where I left them—for
they consider the day well spent if they discover fifty Mycenas whose combined mass is no
bigger than their thumb! Finding myself in good company, I bring out the cheese and bread
—and if I’m lucky they bring out the wine—and we proceed to have an impromptu picnic,
sharing our discoveries, then savoring the silence around us while awaiting the riotous
return of the rest of the group.
For the benefit of these discerning individuals I have included nineteen species of
Mycena in this chapter, which is nineteen more species than the average individual cares
to know. Of course, they constitute only a fraction of the “YAM’s” (“Yet Another My-
MYCENA 225

cena”) that can be found. They are much too small to eat, and some may actually be
poisonous, but they deserve to be better known. They are exquisite in their daintiness and
are among the most attractive of the fleshy (fleshless?) fungi. If your “Mycena” does not
key out convincingly, check Marasmius & Collybia, Omphalina & Xeromphalina, and
the waxy caps (Hygrophoraceae).

Key to Mycena
1. Base of stalk exuding a red to blackish-red juice when cut or squeezed .2
1. Not as above .4
2. Growing on wood ...M. haematopus, p. 231
2. Growing on ground or humus .3
3. Flesh in cap exuding a reddish juice when cut and squeezed .M. sanguinolenta, p. 232
3. Flesh in cap exuding a watery orange-yellow juice when cut and squeezed .
.M. subsanguinolenta (see M. sanguinolenta, p. 232)
4. Fruiting body bright orange to yellow; gills yellow with orange margins; found on hardwoods
in eastern North America (usually clustered) .M. leaiana{ see M. lilacifolia, p. 236)
4. Not as above . 5
5. Stalk (and often the cap) viscid or slimy when fresh and moist .6
5. Stalk not viscid or slimy; cap not normally viscid either . 10
6. Stalk yellow, orange, greenish-gray, or lilac-tinged when fresh, at least at the apex .7
6. Stalk white, gray, or brown (not brightly colored) .9
7. Gills usually decurrent; found on rotting conifers ...M. lilacifolia, p. 236
7. Not as above .8
8. Cap olive-brown to blackish; stalk discoloring brown from the base upward;
common under mountain conifers when or shortly after the snow melts .
.M. griseoviridis(see M. epipterygia, p. 237)
8. Not as above .M. epipterygia & others, p. 237
9. Cap dry (not viscid) .M. rorida, p. 237
9. Cap viscid or slimy, at least when moist . M. vulgaris & others (see M. rorida, p. 237)
10. Fruiting body minute (cap typically less than 1 cm broad and stalk less than 1 (2) mm thick), or if
slightly larger then growing on bark (often mossy) of living trees . 11
10. Fruiting body small but not minute (stalk gene rally at least 1 mm thick; cap at least 5 mm broad
and often larger); growing in a variety of habitats but not usually on bark of living trees . 14
11. Cap brightly colored (red, pink, orange, or yellow).M. acicula & others, p. 228
11. Not as above (cap white, gray, brown, vinaceous, etc.) . 12
12. Typically growing on the bark of trees . 13
12. Typically growing on leaves, twigs, humus, stems, or occasionally on bark ..
.M. capillaris & others, p. 227
13. Cap brownish to vinaceous-brown or vinaceous-buff.
.M. corticola& M. madronicola (see M. clavularis group, p 227)
13. Cap white to grayish or grayish-brown .M. clavularis group & others, p. 227
14. Gills marginate (the edges of at least the larger ones significantly darker and differently colored
than the faces—use hand lens if unsure!) . 15
14. Gills not marginate (edges same color or paler than the faces) .22
15. Gill edges scarlet to bright orange . 16
15. Gill edges pink, dark reddish, purple, yellowish, etc., but not scarlet to bright orange .... 17
16. Gill edges scarlet; hairs at base of stalk orange . M. strobilinoides, p. 228
16. Gill edges orange; hairs at base of stalk yellow-orange .
.M. aurantiomarginata(see M. strobilinoides, p. 228)
17. Gill edges greenish-yellow to yellow to yellow-brown .
.M. citrinomarginata & others (see M. capillaripes, p. 229)
17. Gill edges pink to reddish, purplish-red, reddish-brown, etc. 18
18. Edges of at least the larger gills pink to rosy; growing on ground .... M. capillaripes, p. 229
18. Gill edges darker than above (but faces may be pink or rosy); growing on ground or wood 19
226 TRICHOLOMATACEAE

19. Gills pink to rosy with sordid reddish edges; growing on ground under conifers .
.M. rosella(see M. capillaripes, p. 229)
19. Not as above .20
20. Odor radishlike; typically growing on ground . M. pelianthina & others (see M. pura, p. 230)
20. Odor not radishlike; growing on wood, cones, wood chips, or in lignin-rich humus .21
21. Gill edges very dark purple or grayish-purple .M. purpureofusca, p. 229
21. Gill edges rosy to vinaceous-brown . . M. elegantula & others (see M. purpureofusca, p. 229)
22. Lilac, purplish, blue, or bluish-green tints usually present somewhere on fruiting body ... 23
22. Not as above .24
23. Lilac or bluish tints often present; odor radishlike .M. pura, p. 230
23. Blue or bluish-green tints usually present when fresh; odor more or less mild .
.M. amicta& M. subcaerulea(see M. pura, p. 230)
24. Cap brightly colored (bright coral-pink to orange or yellow) when fresh (may fade in age) 25
24. Cap typically some shade of gray, white, brown, dull vinaceous, etc. (not brightly colored) 26
25. Growing on walnut or hickory nut shells .... M. luteopallens (see M. strobilinoides, p. 228)
25. Growing on ground or in humus . . . M. amabilissima & others (see M. strobilinoides, p. 228)
26. Typically growing on ground, needles, twigs, leaves, etc.27
26. Typically growing on logs, stumps, branches, etc.35
27. Found in grass or disturbed soil; cap brick-red to vinaceous-tinged, soon convex or plane, less
than 2 cm broad.Mycena sp. (unidentified) (see Marasmius oreades, p. 208)
27. Not as above .28
28. Found in grass; stalk tough; cap white to brown, never grayish (seeMarasmius oreades, p. 208)
28. Not as above . 29
29. Extreme base of stalk exuding a droplet of milky fluid when squeezed (fresh specimens only!)
. M. galopus, p. 232
29. Not as above . 30
30. Odor of crushed flesh distinctly alkaline, bleachlike, antiseptic, or at least sharp .
.M. leptocephala & others (see M. alcalina, p. 234)
30. Odor mild or farinaceous, but not as above . 31
31. Stalk with whitish fibrils or particles; cap olive-brown when fresh .M. scabripes, p. 233
31. Not as above . 32
32. Cap margin not often incurved at first; cap often translucent-striate when moist, up to4 (5) cm
broad, grayish to buffy-brown; stalk usually less than4 (6) mm thick; spores often amyloid 33
32. Not as above; margin of cap usually incurved when young and I or cap opaque or differently
colored; spores typically not amyloid .(seeMarasmius, Collybia, & Allies, p. 201)
33. Stalk often with a tapered underground rooting portion; cap sordid tan to buffy-brown; usually
growing near hardwood stumps . M. galericulata& others, p. 235
33. Not as above . 34
34. Stalk base typically exuding a droplet of clear liquid when squeezed; gills often reddish-stained
in age .M. atroalboides(see M. galopus, p. 232)
34. Not as above . . . M. murina group & sundry assorted YAMS (see M. murina group, p. 234)
35. Gills (and often cap) developing sordid pink to reddish spots or stains in age .
. M. maculata & others, p. 235
35. Not as above (but gills may be tinged evenly pinkish in age) .36
36. Growing solitary or in rows, pairs, or groups, but not normally clustered M. subcana, p. 233
36. Not as above . 37
37. Usually growing on hardwood stumps; stalk often rooting in substrate M. galericulata, p. 235
37. Not as above; usually on conifers . 38
38. Fruiting body relatively large (cap 2-5 cm broad), grayish; usually growing near melting snow
(or shortly after snow melts) in mountains .M. overholtsii(see M. subcana, p. 233)
38. Not as above . 39
39. Cap translucent-striate at maturity (when moist), gray to brownish-gray or paler; flesh very
thin .M. occidentalis & many others (see M. subcana, p. 233)
39. Not as above .(see Marasmius, Collybia, & Allies, p. 201)
MYCENA 227

Mycena capillaris (Miniscule Mycena)

CAP 2-6 (8) mm broad, bluntly conical or bell-shaped to convex, sometimes plane in age;
surface smooth, gray, soon fading to white, translucent-striate when moist. Flesh very thin,
odor mild. GILLS adnate to nearly free, well-spaced, grayish then white. STALK 3-7 cm
long, about 1 mm thick, threadlike, equal, very fragile, smooth, dark gray when young
becoming whitish in age. SPORE PRINT white; spores 8-10 * 4-5 microns, elliptical,
smooth, not amyloid.
HABIT AT: S olitary or in small groups on fallen oak(and beech) leaves; widely distributed.
Fairly common in our area in wet weather, but difficult to distinguish from similar species.
EDIBILITY: Unequivocally inconsequential.
COMMENTS: The mycelium of this miniscule whitish Mycena is usually confined to a
single leaf, so that you can pick up the leaf and carry it home, mushroom and mycelium
intact. There are multitudes of other minute white to grayish or brownish Mycenas. They
are especially abundant after long periods of wet weather, but are extremely difficult to
separate (let alone see!) without a microscope. Their ranks include: M. tenerrima andM.
osmundicola, even smaller, with well-spaced, nearly free gills, growing on bark, twigs, and
woody debris (the former with a very short, usually curved stem);M. delicatella, withclose
gills, gregarious on needles and twigs under conifers; M. albidula, with well-spaced,
decurrent gills and a convex cap, on leaves and bark of hardwoods; M. stylobates, with a
flat circular disc at base of stalk; M. paucilamellata, cap only 1 -2 mm broad and gills very
few and foldlike to practically absent; M. ignobilis, growing in mud; and M. juncicola,
vinaceous-tinged, growing on the culms (stems) of sedges. These Mycenas are among the
tiniest of all gilled mushrooms. Marasmius species can also be very tiny but tend to have a
convex to plane cap and dark, wiry or hairlike stem. See also the M. clavularis group.

Mycena clavularis group (Bark Mycena)

CAP 4 -7 mm broad, rounded to convex or somewhat bell-shaped; surface smooth, striate
and often grooved or fluted toward margin; white or translucent, the center and striations
frequently grayish or grayish-brown. Flesh very thin, pallid. GILLS few and widely
spaced, white or tinged gray, attached to the stalk or separating to form a collar around
it. STALK 0.5-2.5 cm long, 0.5-2 mm thick, equal except for a small basal bulb or disc;
white or translucent, smooth, fragile; base often with long white hairs, inserted in bark.
SPORE PRINT white; spores 8-10.5 microns, round or nearly round to broadly ellip¬
tical, smooth, amyloid.
HABITAT: Scattered to gregarious on mossy bark of trees (both living and dead); widely
distributed but not often noticed. I n our area it is common in wet weather, especially on the
lower trunks of living oak and madrone trees.
EDIBILITY: Academic—a tremendous number would be needed for a meal!
COMMENTS: The above description will apply to a number of small Mycenas that
typically grow in groups or troops (but not clusters) on the bark of trees(see photograph on
p. 229). Whether the common form in our area is the “true” M. clavularis or a very similar
species is for the Mycena-experts (of which there are few) to decide. There are several
species with a similar growth habit, including: M. tenerrima, a minute (cap up to 4 mm
broad) whitish species; M. madronicola, a brownish to vinaceous-buff or grayish species
with adnate to slightly decurrent gills; and M. corticola, which has a dark purplish to
vinaceous-brown cap that finally fades to pale grayish-brown, and grows abundantly on
the bark of both hardwoods and conifers in eastern N orth America, often by the hundreds.
228 TRICHOLOMATACEAE

Mycena acicula
CAP 3-7 (10) mm broad, convex or bell-shaped, but sometimes expanding in age; surface
not viscid, coral-red when young, soon fading( often from margin inward) to bright orange-
yellow or yellow. Flesh very thin, yellow, odor mild. GILLS attached (usually adnate), pale
orange to yellow or whitish. ST ALK 1 -7 cm long, less than 1 mm thick, threadlike, equal,
brittle, orange-yellow to yellow, smooth except for hairy base. SPORE PRINT white;
spores 9-11 * 3.5^U5 microns, elongated-elliptical, smooth, not amyloid.
HABITAT: Solitary, scattered, or in small groups on leaves and debris in woods, especially
along streams and in other wet places; widely distributed. Common throughout the
mushroom season in our area, but easily overlooked. I have seen it in large numbers in a
thicket of oak saplings and blackberries along a stream.
EDIBILITY: Unknown, but a great many would be needed for a mouthful!
COMMENTS: This minute Mycena is a delight to behold, its coral-red to yellow cap
contrasting vividly with the dim backdrop of decaying leaves and humus. One usually has
to get down on hands and knees to find it! M. oregonensis is a similar miniature species
whose cap is yellow to yellow-orange from infancy; it grows on needle carpets as well as oak
leaves. For larger brightly colored Mycenas, see M. strobilinoides.

Mycena strobilinoides (Flame M ycena)

CAP 1 -2 cm broad, conical to bell-shaped; surface smooth, not viscid, striate when moist;
red, soon fading to scarlet, then slowly to orange and finally yellow or even whitish; margin
often scalloped. Flesh thin, yellowish; odor mild. GILLS adnate to slightly decurrent, well¬
spaced, yellow to pale pinkish-orange, the edges scarlet (at least when young). STALK
3-6 cm long, 1-2 mm thick, equal, thin, fragile, orange to yellow; base with orange hairs.
SPORE PRINT white; spores 7-9 * 4-5 microns, elliptical, smooth, amyloid.
HABIT AT: Scattered to densely gregarious in needle beds under conifers(especially pine);
widely distributed, but most abundant in the mountains of the Pacific Northwest and
northern California. It fruits in mild, moist weather. I have not seen it in our area.
EDIBILITY: Unknown.
COMMENTS: This little mushroom is likely to attract attention because of its beautiful
color. The scarlet to yellow cap and stalk, scarlet-edged gills, and orange hairs at the stem
base make a most distinctive combination of characters such that even beginners should
have no trouble identifying it. It is one of the more prominent Mycenas in the Pacific
Northwest, sometimes forming colorful “carpets” so thick that it is difficult to examine
them closely without stepping on some. M. aurantiomarginata is a closely related species
with an olive-brown to orange-tinted cap, brilliant orange gill edges, and yellow-orange
hairs at the stalk base. It is commoner at lower elevations in the Pacific Northwest and I
find it fairly frequently in our area as well. Brilliantly colored Mycenas without differently
colored gills edges include: M. monticola, cap coral-red to pink, stalk coral-pink when
fresh; M. amabilissima, cap bright pinkish-red fading to whitish and stalk soon fading to
whitish; and M. adonis, with a smaller (5-12 mm broad), scarlet cap that becomes orange
or yellow-orange and then fades to orange-buff, plus a pale yellow to whitish stalk. All
three of these occur mainly on needle carpets under conifers. M. luteopallens, on the other
hand, grows on old walnuts and hickory nuts in eastern North America. It is bright orange
to yellow before fading. All of the above have stalks which are typically at least 1 mm thick.
For minute, brightly colored species with thinner or threadlike stalks, see M. acicula.
Left: Mycena clavularis group (p. 227) typically grows on bark of standing trees. Right: Mycena
capillaripes is one of many species that form dense carpets of delicate domes on duff under conifers.

Mycena capillaripes
CAP 0.5-2.5 cm broad, oval becoming bell-shaped or conical, often with a blunt umbo,
sometimes nearly plane in age; surface smooth, not viscid, usually some shade of vina-
ceous-gray or gray, the center often browner; translucent-striate when moist, usually
wrinkled or furrowed when dry. Flesh thin; odor typically nitrous or bleachlike(especially
when crushed), but sometimes radishlike. GILLS attached (usually adnate), well-spaced,
typically pallid or vinaceous-gray, at least the longer ones with pale pink or rosy edges(use
hand lens!). STALK 3-10 cm long, 1-3 mm thick, equal or slightly thicker at either end,
hollow, smooth, fragile, colored more or less like cap or paler above, sometimes slightly
yellower in age; base with coarse white hairs. SPORE PRINT white; spores 7-12 * 4-6
microns, elliptical, smooth, amyloid.
HABITAT: Scattered to densely gregarious on conifer duff (but occasionally also under
hardwoods); widely distributed. In our area it often occurs in droves under Monterey pine
in the fall and winter.
EDIBILITY: Inconsequential.
COMMENTS: This is one of several Mycenas that carpet our pine forests with a miniature
“fungal jungle” of petite parasols. The faint reddish or vinaceous tints on the cap plus
the pink-margined gills are distinctive. M. rosella is a widely distributed pink to grayish-
rose species with pale to bright rosy gills that have darker (pale to dark reddish) edges. It
is uncommon in our area but often abundant under pine and other conifers farther north.
Other terrestrial species with colored gill margins include: M. citrinomarginata, with pale
yellow to yellow-brown gill edges and spores 8-11 microns long;M. olivaceobrunnea, with
smaller spores, a smaller cap, and yellowish gill edges; andM. elegans, with pale greenish-
yellow gill edges. In these species the cap is typically some shade or mixture of olive-brown,
gray, and/ or yellow.

Mycena purpureofusca
CAP 0.5-2.5 cm broad or occasionally larger, obtusely conical with a slightly incurved
margin, becoming broadly conical or bell-shaped or sometimes expanding to nearly plane
in old age; surface smooth, not viscid, dark purplish with a paler (lilac) margin when young,
fading to purplish-gray or vinaceous-lavender; translucent-striate when moist and mature.
Flesh thin, pliant, odor mild. GILLS attached (usually adnate), fairly close, pallid to
Mycenapurpureofusca commonly grows on pine cones and other woody debris.

grayish with dark grayish-purple edges. STALK 3-1 Ocm long, 1-3 mm thick, equal, hollow,
cartilaginous, colored more or less like cap or paler above; base with white hairs and some¬
times rooting. SPORE PRINT white; spores 8-14 * 6-8.5 microns, broadly elliptical,
smooth, not amyloid.
HABITAT: Solitary or in small groups or tufts on conifer wood and debris; widely
distributed and frequent in our area in the fall and winter. I find it often on old pine cones.
EDIBILITY: Unknown, and like myself, likely to remain so.
COMMENTS: The purplish conical cap, dark purplish gill edges, and growth on wood
or cones typify this species. The stalk does not exude a red latex like M. haematopus. The
color and stature are reminiscent of Marasmiusplicatulus, which has a very tough, brittle
stalk and pallid gill edges and grows on the ground. Collybiafuscopurpurea has a similar
name but has a convex to plane cap and is also terrestrial. Other species: Mycena elegantula
is closely related, but has a vinaceous-brown to pinkish cap and pale pink to vinaceous gill
edges; it also grows on coniferous wood and debris, including cones. M. rubromarginata
grows on spruce and fir and has a browner cap and reddish-brown gill edges.

Mycena pura (Lilac Mycena)

CAP 2-5 (6.5) cm broad, obtusely umbonateorconvex becoming broadly convex or plane,
sometimes with an uplifted margin; surface smooth, hygrophanous, color variable: various
shades or mixtures of lilac, pink, gray, and blue-gray, but varying also to blue-green with a
yellowish center, or sometimes even whitish tinged purple or blue at the center; translucent-
striate when moist, otherwise opaque. Flesh thin, soft; odor and taste radishlike. GILLS
adnate or adnexed, usually tinged cap color (lilac, etc.), but sometimes grayish or white;
edges pallid. STALK 3-7 (10) cm long, 2-7 mm thick, equal or thicker below, hollow,
smooth, pallid or colored like the cap or paler. SPORE PRINT white; spores 5-9 * 3-4
microns, elliptical to cylindrical, smooth, amyloid.
HABITAT: Solitary or in groups or small tufts on ground in woods, widely distributed.
Common in our area from fall through early spring, but rarely in large numbers. I find it
most often with oak, Douglas-fir, and pine.
EDIBILITY: Edible according to some sources, but one study revealed traces of the toxin
muscarine. Definitely not recommended—it is too small to be of value anyway.

230
Mycena pur a is a terrestrial species that sometimes resembles a miniature blewit. Note how the cap
becomes convex or plane in age.

COMMENTS: This is one of the larger Mycenas as well as one of the more beautiful. The
color is exceedingly variable, but there is usually a trace of lilac somewhere on the fruiting
body, especially the stem. The convex rather than conical cap is atypical for a Mycena,
and for this reason it is keyed out under Collybia. The radishlike odor is another important
fieldmark. The gill edges are not purple or pink as in M. capillaripes and M. purpureofusca,
and it is strictly terrestrial. It never grows in huge clusters and does not normally fruit in
dense troops either. I have seen fairly robust specimens which could have been mistaken for
blewits (Clitocybe nuda). Inocybe lilacina is similarly colored, but has brown spores.
M. pelianthina and M. rutilantiformis are two very similar species with a radishlike odor.
They differ in having dark reddish to purple gill edges, and the latter typically shows yellow
at the apex of the stalk. They are not as highly colored as M. pura and both grow on the
ground under hardwoods. M. amicta is a western species whose cap and stalk are blue to
blue-green or tinged those colors. It grows on coniferous debris, while M. subcaerulea is
its eastern, deciduous- woodland counterpart.

Mycena haematopus (Bleeding Mycena) Color Plate 46

CAP 1-3.5 (5) cm broad, oval to bell-shaped when young, with the margin oftenextending
beyond the gills; in age sometimes convex or plane with an umbo and uplifted margin;
surface smooth, not viscid, reddish to vinaceous-brown to reddish-brown or pinkish-
brown, margin often paler (vinaceous-gray); striate at maturity when moist; margin often
scalloped in age. Flesh thin, exuding a dark red juice when cut. GILLS attached (usually
adnate or adnexed), fairly close, pallid, often developing reddish stains; edges white or in
one form reddish. STALK 3-8 (14) cm long, 1-3 mm thick, equal, dull reddish or reddish-
brown, or sometimes pallid; smooth, fragile; base with coarse hairs, exuding a dark red
juice when cut or squeezed. SPORE PRINT white; spores 7-11 * 5-7 microns, elliptical,
smooth, amyloid.
HABITAT: Solitary or more commonly tufted or in groups on decaying logs and stumps
(mostly hardwoods). Very widely distributed and common, fruiting in our area from fall
through spring.
EDIBILITY: Edible according to some sources, but too small to be of value.
COMMENTS: One of the most distinctive and common Mycenas, this species resembles
several others in color, but if the base of the stalk is squeezed a dark red blood like fluid will
emerge. Its growth on wood separates it from other “bleeding” Mycenas(see M. sanguino-
lenta). Also distinctive is the frequent presence of a sterile band of tissue around the cap
margin of young specimens. It is suggestive of a veil but breaks up in age or forms a
scalloped rim around the cap.

231
Left: Mycena sanguinolenta is a common terrestrial species that “bleeds” when broken. Right:
Mycena galopus looks like numerous other grayish or brownish Mycenas, but its base exudes a milky
droplet when squeezed.

Mycena sanguinolenta (Terrestrial Bleeding Mycena)

CAP 0.3-1.5 (2.5) cm broad, conical to bell-shaped or sometimes convex in age; surface
smooth, not viscid, color variable: some shade of pale reddish-brown to bright reddish-
brown to orange-brown, etc; margin often vinaceous and furrowed at maturity. Flesh thin,
reddish, exuding a dark red juice when cut; odor mild. GILLS attached (typically adnate),
well-spaced, pallid or tinged flesh-color or reddish, edges dark reddish-brown. ST ALK 2-
7.5 cm long, 1 -2 mm thick, equal, hollow, fragile, colored more or less like cap, smooth; base
with white hairs and exuding a dark red juice when broken or squeezed. SPORE PRINT
white; spores 8-11 x 4-6 microns, elliptical, smooth, weakly amyloid. Cystidia present on
gill faces and edges.
HABITAT: Solitary or widely scattered to gregarious or tufted on leaf mold and needles
in woods or at their edges; widely distributed. Common in our area in fall and winter,
usually scattered under oak. I n northern California and the Pacific North west it sometimes
fruits in large numbers under conifers, especially spruce.
EDIBILITY: Unknown, and like most of us, destined to remain so.
COMMENTS: True to their names, this species and its close cousin, M. haematopus,
“bleed” when broken. As such they are the easiest Mycenas to recognize and among the
most common as well. M. sanguinolenta is easily distinguished from M. haematopus by its
growth on the ground, absence of a sterile band of marginal tissue on the cap, and some¬
what different color. The striate or furrowed cap is reminiscent of Marasmiusplicatulus,
but that species has a tough, polished stem and does not“bleed.” Other species: M. subsan-
guinolenta is closely related but has dark red to orange juice, is slightly yellower overall,
and lacks cystidia on the gill faces. It often fruits in large numbers under conifers in
northern California.

Mycena galopus (Milky Mycena)

CAP 0.5-2.5 cm broad, conical to bell-shaped orumbonate, the margin sometimes curling
up in age; surface smooth, not viscid, at first grayish-black (at least at center), but soon
fading to gray or grayish-brown, then pale gray or even whitish; translucent-striate when
moist. Flesh thin, soft, odor mild. GILLS well-spaced, attached (usually adnate), white
to grayish. STALK 4-8 (12) cm long, 1-2 mm thick, equal, fragile, hollow, smooth, dark
grayish-brown to gray below, paler above; base with white hairs, exuding a droplet of milky
juice when squeezed. SPORE PRINT white; spores 9-13 x 5-6.5 microns, elliptical,
smooth, not amyloid or sometimes weakly amyloid.

232
MYCENA 233

HABITAT: Scattered to gregarious on ground and humus in woods, widely distributed.

Common in our area during wet weather, especially under redwood and pine, but tending
to fruit widely scattered over a large area rather than in dense troops or“carpets” like some
of our other pine-loving species (e.g., M. murina, M. capillaripes).
EDIBILITY: Who knows?
COMMENTS: This ubiquitous Mycena will be encountered by anyone who takes their
“LBM’s” seriously. At first glance there is little to distinguish it from the myriad of other
anonymous brown or grayish Mycenas. However, if you break the very base of the stalk
and squeeze gently, a drop of milky or cloudy fluid will emerge. This is best observed in
the field, while the fruiting body is fresh. A similar species, M. atroalboides, exudes liquid
that is clear rather than milky, and often has reddish spots on the gills in old age. It is not
common in our area but is often abundant farther north under conifers.

Mycena scabripes (Rough-Stemmed Mycena)

CAP 1.5-3 (5) cm broad, obtusely conical becoming convex, umbonate, or plane; surface
not viscid, blackish-brown or dark brown at the center, olive-brown toward the margin,
but fading somewhat in age to olive-gray or gray; not distinctly striate. Flesh thin,
brownish-gray, odor mild. GILLS adnate to adnexed or free in age, pallid or tinged cap
color, sometimes spotted reddish-brown in age. STALK 3-9 cm long, 2-4 mm thick, equal,
hollow, grayish or colored like cap, with a thin but distinct coating of silky white to grayish
fibrils and particles. SPORE PRINT white; spores 7-9 * 4-5 microns, elliptical, smooth,
amyloid.
HABITAT: Scattered to gregarious or in small tufts in humus and debris in woods; widely
distributed but not common, or at least seldom noticed. I have found it in our area in the
fall and winter under redwood and oak.
EDIBILITY: Who knows? Who cares?
COMMENTS: The distinguishing feature of this undistinguished Mycena is the presence
of silky white fibrils and particles on the stem. The olive-brown, more or less umbonate
cap is also distinctive.

Mycena subcana (Neutral Gray Mycena)

CAP 0.5-2.5 (3) cm broad, oval or obtusely conical to bell-shaped, becoming convex in
age; surface smooth or with a hoary sheen, grayish to pale gray, usually darker at the center,

Left: Mycena scabripes is a nondescript species with a minutely scaly or dandruffy stalk. Right:
Mycena subcana typically grows singly or in pairs or groups (but not clusters) on sticks and branches.
234 TRICHOLOMATACEAE

sometimes fading to whitish; not viscid, translucent-striate nearly to center when moist.
Flesh thin, grayish, odor mild. GILLS attached (usually adnate), well-spaced, whitish to
pale gray. STALK 1.5-6 cm long, 1.5-3 mm thick, equal or enlarged at base, smooth, same
color as cap or paler, with downy white hairs at base. SPORE PRINT white; spores
8-10 * 5-6 microns, elliptical, smooth, amyloid.
HABITAT: Solitary or in small groups (not large clusters!) on dead sticks and branches,
sometimes on living trees; fairly common on the west coast. In our area it fruits in the fall
and winter on conifers (especially redwood); I have also found it on tanoak.
EDIBILITY: Who knows? Who cares? I don’t.
COMMENTS: This grayish species is representative of a multitude of indifferent grayish,
wood-inhabiting “YAM’s.” The salient macroscopic features of M. subcana are its
color, its translucent-striate cap, and its preference for dead sticks and branches
rather than stumps or logs (a completely different niche!). Unlike many of its kin, it does
not grow in clusters, but instead fruits alone or in attractive rows of solitary or sometimes
paired specimens. Typifying those that grow in clusters on stumps or logs is M. occiden-
talis, a grayish species found throughout the West on conifers. Also clustered on conifers
is M. laevigata, a whitish species with grayish-black to bluish-gray tints on cap and stalk
when very young and yellowish discolorations in age; and M. overholtsii, a large grayish
species found in clumps near melting snow in the spring. Another species,M. parabolica,
has a growth habit in between that of M. subcana and M. occidentalis—it grows in groups
or small clusters on rotting wood and has a sooty black cap that fades to gray in age.

Mycena alcalina (Alkaline Mycena)

CAP 1^4 cm broad, conical to bell-shaped or convex-umbonate, expanding somewhat in
age; surface smooth, dark brownish-black to grayish-black when very young, soon fading
to gray, grayish-brown, or yellow-brown; striate when moist. Flesh thin, fragile, odor
faintly to strongly alkaline (like bleach). GILLS rather close, adnate to slightly decurrent,
whitish to grayish, sometimes stained reddish-brown in age. STALK 3-10 cm long, 1.5-3
mm thick, equal, pallid to grayish or often sordid yellowish-brown in age; fragile, hollow.
SPORE PRINT white; spores 8-11 * 5-7 microns, elliptical, smooth, amyloid.
HABITAT: In groups or tufts on decaying conifer logs, or sometimes densely gregarious
on needles under conifers; common and widespread throughout North America. In some
regions it is most abundant in the spring, but in our area it fruits in the fall and winter.
EDIBILITY: Who knows? Who cares? I don’t. Do you?
COMMENTS: The sharp, bleachlike odor is usually quite distinct in this species, and
helps distinguish it from the throngs of other grayish Mycenas that grow on wood. The
odor is best detected by crushing the cap. Another common species with an alkaline odor,
M. leptocephala, is terrestrial, growing on sticks, needles, or even in grass. It is only rarely
umbonate, and slightly smaller than M. alcalina. Other species: M. metata is terrestrial,
with a slightly browner cap and faint but sharp odor; M. iodiolens has an antiseptic odor.
For similar species without a distinctive odor, see M. subcana and the M. murina group.

Mycena murina group (Yet Another Mycena) Color Plate 48

CAP 0.7 -3 cm broad, conical to bell-shaped, striate nearly to the center when moist; surface
smooth, not viscid, dark gray to gray when moist, paler (gray to whitish) when faded,
darker when young. Flesh thin, odor mild. GILLS grayish with pallid edges, attached,
fairly well-spaced. ST ALK 3-8 cm long, 1 -4 mm thick, equal, fragile, smooth, pale to dark
gray with pallid apex; base with white hairs. SPORE PRINT white; spores 8-11 * 5-7
microns, elliptical, smooth, amyloid.
MYCENA 235

HABIT AT: Scattered to densely gregarious by the hundreds on needle duff under conifers,
widely distributed. This species and its look-alikes are often abundant in our coastal pine
forests in the late fall and early winter. They also appear under hardwoods, in grass, even
in planter boxes and flower pots—i.e., wherever there is organic matter to nourish them.
EDIBILITY: Who knows? Who cares? I don’t. Do you? Do you care if I do?
COMMENTS: This species, which has passed under the name M. stannea, is one of a slew
of nondescript, odorless, gray to brownish, terrestrial Mycenas that are very prevalent
under conifers. They are differentiated from each other largely on the basis of microscopic
characteristics such as the size and shape of spores and sterile cells (cystidia). The one pic¬
tured in the color photograph is not quite as gray as typical M. murina and may well be a
distinct species, but to the average mushroom hunter it is a “YAM” (“Yet Another My-
cena”), and thus hardly worth the time and trouble to distinguish. Other “Y AM’s” include
M. latifolia, M.fHopes, M. abramsii,M. pseudotenax, M. atroalboides, ad infinitum. For
those species with a sharp or bleachlike odor, see comments under M. alcalina, and for
those with colored gill margins, see M. capillaripes.

Mycena maculata (Reddish-Spotted Mycena)

CAP 1.5-5 cm broad, conical to bell-shaped expanding to broadly convex or plane with a
broad umbo, the margin sometimes uplifted in age; surface smooth, blackish-brown soon
fading to brown or brownish-gray, but usually spotted with reddish-brown and sometimes
entirely watery gray; striate when moist. Flesh thin, firm, grayish, slowly bruising reddish-
brown. GILLS adnate to adnexed, whitish to pale gray, soon stained with reddish spots,
sometimes entirely sordid reddish in old age; edges pallid. STALK 4-8 (12) cm long, 2-5
mm thick, nearly equal, smooth, hollow, cartilaginous, pallid above, colored like cap or
paler below; base with dense white hairs and soon stained reddish-brown, sometimes
rooting. SPORE PRINT white; spores 7-9 x 4-5 microns, elliptical, smooth, amyloid.
HABIT AT: In small tufts or large clusters on logs and stumps, especially of conifers; widely
distributed. It is common in our area in cool, wet weather on redwood, Douglas-fir, etc.,
occasionally also on hardwoods. “The most abundant Mycena on conifer wood in the
Pacific Northwest,” says Alexander Smith, author of a monograph on the genus.
EDIBILITY: Who knows? Who cares? I don’t. Do you? Do you care if I do? I won’t if
you don’t.
COMMENTS: This rather unattractive species can be recognized by the sordid reddish
stains that develop in age and its growth in clusters on rotten wood. The stalk sometimes
has a “tap root” as in M. galericulata. It doesn’t “bleed” like M. haematopus, and the gill
edges aren’t purple as in M. purpureofusca. M. occidentalism see comments under M. sub-
cana) is also common in clusters on conifers, but does not develop reddish stains. Another
reddish-staining species, M. rugulosiceps, has larger spores and grows on hardwoods
(alder, rhododendron, etc.) as well as on conifers. M. inclinata is a closely related species
that is especially common in eastern North America. It often develops reddish or pinkish
tones as it ages but favors hardwoods and has fibrillose flecks on the lower half of the stalk.
See top of next page for a photograph of M. maculata.

Mycena galericulata
CAP 2-5 (7) cm broad, conical when young becoming broadly bell-shaped to umbonate
to plane or with an uplifted margin in age; surface moist but not viscid, often radially
wrinkled nearly to center; buffy-bro wn to grayish, dingy tan, cinnamon-brown, or grayish-
brown. Flesh watery gray; odor faintly to distinctly farinaceous or radishlike. GILLS
adnate to adnexed, often with cross-veins between, white to grayish or in age flushed pale
Left: Mycena maculata (see p. 235) is one of many Mycenas that grow in clumps on rotten wood.
Center: Mycena galericulata, young specimens with broadly conical caps; note how stalk roots deeply.
Right: Mycena galericulata, mature specimens which have opened up so as to resemble Collybias.

pinkish, but not spotted or stained. STALK 5-10 (14) cm long, 2-4 (6) mm thick, equal,
cartilaginous, hollow, smooth, often rooting; grayish-white or colored more or less like
cap, often darker below. SPORE PRINT white; spores 8-11 * 5.5-7 microns, elliptical,
smooth, amyloid.
HABITAT: Solitary to scattered, gregarious, or in small clusters on decaying hardwood
stumps, logs, and debris; widely distributed and not uncommon in our area in the fall and
winter. One form—possibly a distinct species—grows on manzanita burls.
EDIBILITY: Edible, but not recommended—it is one of the few Mycenas large enough
to eat, but many of its difficult-to-distinguish relatives have not been tested.
COMMENTS: This widespread species can be mistaken for a Collybia because the cap
is convex to plane in age. The telltale traits are its pale color, frequently long, rooting stem,
and penchant for growing scattered or in small, loose clusters. The gills are often tinged
pinkish in age, but do not become reddish-spotted as in M. maculata. Other species:
M. vitilis has a long stem that often has a rooting base that is covered with white hairs
below the ground. It grows in humus under hardwoods, especially alder.

Mycena lilacifolia
CAP 0.8 -2.5 cm broad, convex to helmet-shaped, the center slightly depressed in age;
surface viscid to slimy and translucent-striate when moist, smooth, lilac when very young,
but soon bright to pale yellow. Flesh thin; odor mild. GILLS well-spaced, usually
decurrent, pale lilac becoming whitish to pale yellow or pinkish. STALK 1 -4 cm long, 1 -2
mm thick, equal or enlarged at the base, smooth, slimy-viscid when moist, same color as
gills but soon yellow; base often with lilac mycelium. SPORE PRINT white; spores
6-7 x 3-3.5 microns, elliptical, smooth, not amyloid.
HABITAT: Solitary, scattered, or in groups on rotting conifers, widely distributed.
It usually fruits in cool weather and is fairly common in our coastal pine forests.
EDIBILITY: Unknown, but too slippery to be of value.
COMMENTS: This species is best distinguished from other viscid Mycenas by its yellow
color and growth on conifers. The gills will sometimes retain their lilac tint longer than
the cap and stalk, but I have found specimens which showed absolutely no trace of their
original lilac color. It might be looked for in Omphalina because of the decurrent gills
(see photo on p. 237), but the slimy stem and cap distinguish it. Viscid, brightly colored
wood-inhabiting species with non-decurrent gills include: M. leaiana, bright orange to
yellow, growing in clusters on hardwoods in eastern N orth America, and certain varieties
of M. epipterygia(see description on next page), which grow on conifers.

236
Mycena lilacifolia (p. 236) is one of several Mycenas with a slimy cap and stem. Note decurrent gills.

Mycena epipterygia (Yellow-Stemmed Mycena)

CAP 0.8-2 (2.5) cm broad, oval becoming umbonate, bell-shaped, or convex; surface
smooth, slimy or viscid when moist, color variable: mostly yellow with olive or grayish
tones, sometimes fading in age. Flesh thin, yellowish; odor mild or faintly fragrant.
GILLS attached, whitish or tinged yellow, fairly well-spaced. STALK 5-9 cm long,
1 -2 mm thick, equal, fragile, hollow, smooth, slimy or viscid when moist, yellow, but some¬
times* fading to whitish in old age. SPORE PRINT white; spores 8-10 x 5-6 microns,
elliptical, smooth, amyloid.
HABITAT: Scattered to gregarious in needle duff under conifers, widely distributed. It
is partial to cold weather, fruiting in northern regions in the late fall until frosts set in, and in
the winter in our coastal pine forests. One variety with a rank odor occurs on decaying
conifers. The related M. griseoviridis (see comments) frequently fruits nearly melting
snow in the spring and summer.
EDIBILITY: Unknown.
COMMENTS: This is one of several attractive Mycenas with a viscid cap and viscid,
yellow to greenish-gray stem. The viscid layers may dry out in age, but the colors are dis¬
tinctive. Similar terrestrial species include: M. viscosa, with a strong rancid odor and taste;
M. epipterygioides, with a dark olive-gray cap that does not fade to white; andM. griseo¬
viridis, with a deep olive to olive-brown to blackish cap, a yellow stem that becomes brown
or deep vinaceous-brown from the base upward, a cucumberlike odor, and thorny cystidia
on the gill edges. The latter species is often common under mountain conifers soon after
the snow melts, but is not restricted to that habitat. Any of these species can be mistaken
for a waxy cap (e.g., Hygrocybepsittacina), but their gills are not noticeably waxy.

Mycena rorida (Slippery Mycena)

CAP (2) 5-10 (15) mm broad, at first rounded or bell-shaped, then expanding to plane or
nearly so; surface dry, pale brown or brownish-gray to tan, fading to white or yellowish-
white. Flesh thin, odor mild. GILLS adnate to decurrent, well-spaced, white. STALK 2-3
(5) cm long, about 1 mm thick, equal, whitish, covered with a sheath of slime when fresh and
moist. SPORE PRINT.white; spores 8-12 x 4-6 microns, elliptical, smooth, amyloid.
HABITAT: Solitary or scattered to gregarious on ground and debris under conifers and in
mixed woods; widely distributed, but not common in our area. I have found it under
redwood in November.
EDIBILITY: Too small and slippery to be of value.
COMMENTS: This little mushroom is easily identified by its pale color, dry cap, and
sheath of slime on the stem. Plucking it can be a difficult proposition if you try to grasp it by
the stalk. Other Mycenas with a pallid to grayish, viscid stem include: M. tenax, with a
gray-brown to pale gray cap and strong disagreeable odor;M. vulgaris, similar but with an
extremely viscid cap and slight odor; and M. clavicularis, with a dry grayish cap and mild
odor. All of these occur in groups or troops under conifers.

237
 spores
ENTOLOMATACEAE
THIS is the largest family of pinkish-spored mushrooms and also the most difficult. It
is unusual among agarics in having spores which are distinctly angular (several-sided) in
side and/ or end view. The gills are typically attached to the stalk (in contrast to the other
major pinkish-spored family, the Pluteaceae), and the stalk is not cleanly separable from
the cap. The spore color, though characterized as “pinkish,” actually ranges from deep
flesh-color to salmon- or rosy-pink to dull reddish or sordid pinkish-cinnamon, whereas
the pinkish-spored members of the Tricholomataceae (e.g., Clitocybe) have pale or dull
pinkish, non-angular spores.
The Entolomataceae have a very confusing nomenclatural history. Four principal
genera have traditionally been recognized—Clitopilus, Entoloma, Nolanea, and Leptonia
—but the presence of intermediate forms has led some mycologists to consolidate all but
Clitopilus in a single giant genus, Entoloma, while recognizing a few smaller genera (e.g.,
Rhodocybe) that differ microscopically. To muddle matters more, the name Rhodophyllus
is often used instead of Entoloma, and Rhodophyllaceae instead of Entolomataceae.
The result is that each species has a surfeit of superfluous synonyms—for instance,
Leptonia sericella, Alboleptonia sericella, Entoloma sericellum, and Rhodophyllus
sericellus are all names for the same fungus! (Human beings are certainly a confusing—and
confused—lot!)
The mushrooms do little to ameliorate matters, for they are exasperatingly difficult to
demystify. Some Entoloma-experts have resorted to such abstruse activities as measuring
the urea concentration of the fruiting body or determining the type of symmetry exhibited
by the spores in their efforts to delineate the various genera and species!
In North America the Entolomataceae are largely woodland fungi, but some species
grow in open or grassy areas. The vast majority are terrestrial (in contrast to the wood-
inhabiting genus Pluteus), but a few occur on rotting wood. Several species are poisonous,
and in view of the difficulty in identifying them, beginners should avoid the entire group.
Particularly dangerous are some of the large fleshy Entolomas—a fact which clearly
invalidates the old adage that “any mushroom with pink gills is edible” (a reference, no
doubt, to Agaricus).
From two to nine genera are recognized, depending on whether a “lumper” or “splitter”
is doing the recognizing. Several of the genera are small and rare (e.g., Claudopus and
Pouzarella)\ these are included in the following key but not treated beyond it.

Key to the Entolomataceae

1. Growing on dung or on other mushrooms; widely distributed but rare .21
1. Growing on wood, humus, or ground . 2
2. Stalk lateral to absent; fruiting body small, usually on wood (rarely on ground) . 3
2. Stalk present, usually central or slightly off-center; on ground or sometimes on wood .... 7
3. Gills pale orange, orange, or yellow or gill edges split lengthwise (see Tricholomataceae, p. 129)
3. Not as above .4
4. Odor strongly unpleasant (skunklike) . Claudopus grayeolens
4. Not as above . 5
5. Cap felty or hairy, white, 4-20 mm broad; spores not angular .(see Crepidotus, p. 405)
5. Not as above; spores angular under the microscope .6
6. Cap brownish-orange to grayish-brown beneath a dense layer of whitish fibrils; odor usually
farinaceous; stalk silky-hairy, base often with white mycelial threads . Claudopus byssisedus
6. Cap whitish to gray or pinkish-gray, not as hairy as above, 3-7 mm broad; odor mild .
.Claudopus depluens
7. Gills purple or purple-tinged; spores not angular .(see Clitocybe & Allies, p. 148)
7. Not with above features; gills rarely purple, and if so then spores angular.8
ENTOLOMATACEAE 239

8. Gills often decurrent(but sometimes adnate or even notched); cap and stalk never bluish, violet,
or black; spores warted or longitudinally lined in side view, angular only in end view .
. Clitopilus& Rhodocybe, below
8. Not as above; spores usually angular in side view or not angular at all .9
9. Cap and/ or stalk blue, blue-black, blue-gray, violet, steel-gray, or black, at least when young
and fresh. 10
9. Not as above . 12
10. Stalk usually 5 mm thick or more; usually growing on ground . 11
10. Stalk slender, usually hollow and fragile or cartilaginous, typically 1.5-5 mm thick at apex, or if
thicker then growing on wood .Leptonia & Allies, 248
11. Stalk fibrillose-scaly, often with a metallic luster or iridescence; cap usually fibrillose-scaly
also .LeptoniaSi Allies, p. 248
11. Stalk smooth to finely fibrillose, or if fibrillose-scaly then cap smooth.Entoloma, p. 242
12. Fruiting body small and white (or in age tinged yellowish or pinkish) LeptoniaSi Allies, p. 248
12. Not as above . 13
13. Cap hairy and/ or scaly and base of stalk with coarse, stiff hairs; not common .. . Pouzarella
13. Not as above . 14
14. Stalk fleshy, 4 mm thick or more . 15
14. Stalk slender, usually hollow and fragile or cartilaginous, usually less than 5 mm thick ... 17
15. Spore print pale to dull pinkish; spores not angular . (see Tricholomataceae, p. 129)
15. Spore print deep flesh-color to salmon-pink to pinkish-cinnamon or darker; spores angular
under the microscope . 16
16. Cap conical to bell-shaped or with a pointed umbo (but may expand with age) Nolanea, p. 245
16. Not as above (but cap may be bluntly conical when young) .Entoloma, p. 242
17. Growing on or near wood; spores smooth or at least not angular . 18
17. Usually terrestrial (but occasionally on wood); spores angular under microscope . 19
18. Gills usually free (or nearly free) and close together . (seePluteus, p. 254)
18. Not as above . (see Tricholomataceae, p. 129)
19. Stalk yellow-green or gills pale with pink edges.. LeptoniaSi Allies, p. 248
19. Not as above .20
20. Cap conical to bell-shaped or umbonate, or if convex to plane then smooth or silky; cap not
umbilicate .Nolanea, p. 245
20. Cap convex to plane or umbilicate and usually finely fibrillose, scurfy, scaly, or fibrillose-scaly,
at least at the center (and particularly in age) .Leptonia & Allies, p. 248
21. Parasitic on other mushrooms (mostly chanterelles and polypores); fruiting body minute,
grayish to white; small stalk usually present. Claudopusparasiticus
21. Not as above; growing on dung . Clitopilus& Rhodocybe, p. 239

CLITOPILUS& RHODOCYBE
Small to medium-sized, mostly terrestrial mushrooms. CAP convex to plane or depressed. GILLS
usually adnate to decurrent. STALK thick or slender, fleshy or rather fragile, central or off-
center. VEIL and VOLVA absent. SPORE PRINT pinkish to salmon, flesh-colored, or pinkish-
cinnamon. Spores angular in end view only; in side view longitudinally ridged (Clitopilus) or
bumpy to warty (Rhodocybe).

THESE two small genera of pinkish-spored mushrooms typically have adnate to decurrent
gills and spores which are angular only in end view. They are a rather lackluster lot, most
likely to be confused with species of Entoloma, which usually have notched gills, and
Clitocybe, which have paler, non-angular spores. Several small species of Leptonia and
Eccilia have decurrent gills, but their spores are angular in both side and end view. There
are also several Rhodocybes with notched gills; these are likely to be confused with Colly-
bia or Entoloma unless their spores are examined under the microscope.
Both Clitopilus and Rhodocybe are small genera, but some of their species are quite
common. Two are described here (one from each genus) and several others are keyed out.
240 ENTOLOMATACEAE

Key to Clitopilus& Rhodocybe

1. Odor like a candy store or bubble gum (break open cap if unsure) .(see TVolanea, p. 245)
1. Not as above . 2
2. Cap reddish-brown to orange-brown, cinnamon, pinkish, yellowish-ochre, or tan .3
2. Cap white or grayish to grayish-brown, olive-brown, etc.4
3. Cap 2-12 cm broad, opaque; stalk usually at least 5 mm thick; fairly common in California,
but more widely distributed .R. nuciolens & others, p. 241
3. Cap usually less than 5 cm broad and translucent-striate when moist; stalk 1-7 mm thick;
widespread .R. nitellina{see R. nuciolens, p. 241)
4. Found on dung or compost; cap small, stalk often poorly developed .C. passeckerianus
4. Not as above . 5
5. Cap gray to grayish-brown; “aborted” fruiting bodies often found that are somewhat puffball¬
like: whitish to pinkish-tan and bumpy on the outside, marbled within; common in eastern
North America; spores angular in side view .(see Entoloma, p. 242)
5. Not as above . 6
6. Gills decurrent; cap white to grayish and typically 3 cm or more broad . 7
6. Not as above; cap smaller and/ or dark brown to olive-brown, etc.9
7. Taste bitter; cap often cracked concentrically . R. mundula (see R. nuciolens, p. 241)
7. Not as above . 8
8. Stalk often somewhat off-center and fairly slender (4-12 mm thick); odor pleasant; typically
not growing in clusters; spores longitudinally lined under microscope . . C. prunulus, below
8. Stalk often quite thick; sometimes clustered; spores not lined (see ClitocybeSc Allies, p. 148)
9. Spores finely warted in side view, angular only in end view R. caelata(see R. nuciolens, p. 241)
9. Spores angular in side and end views .(seeLeptonia&c Allies, p. 248)

Clitopilusprunulus (Sweetbread Mushroom)

CAP 3-10 cm broad, convex becoming plane or centrally depressed; surface dry and
slightly felty or in one form slightly viscid when moist, smooth, white to gray; margin often
lobed or wavy. Flesh rather thin, white; odor strongly but pleasantly farinaceous (like
sweetbread). GILLS decurrent or occasionally adnate, fairly close, narrow, whitish to
gray, then dusted pinkish by spores. STALK 2-8 cm long, 0.4-1.2 cm thick, central or off-
center, solid, equal or tapering downward, white or grayish. SPORE PRINT flesh-colored
or salmon; spores 9-12 * 5-7 microns, elliptical, longitudinally ridged, angular only in end
view.

Clitopilus prunulus looks like a Clitocybe with its decurrent gills and white to gray color, but its
spores are pinkish and show ridges under the microscope. The tree frog shown here is very tiny!
CLITOPILUS 241

HABITAT: Solitary to scattered or in small groups on ground in open woods and grassy
places near trees; widely distributed. It is fairly common in our area in the fall and winter,
usually under oak or pine.
EDIBILITY: Edible and choice, but not recommended. It is rated highly in Europe but is
easily confused with poisonous mushrooms such as Clitocybe dealbata.
COMMENTS: The pinkish spores, decurrent gills, white to grayish cap, and strong odor
of meal are the fallible fieldmarks of this unremarkable fungus. The poisonous Clitocybe
dealbata closely mimics it but is somewhat smaller and has white spores. Clitocybe sub-
connexa and its close relatives have pinkish spores, but grow in clusters and are more
robust. The name Clitcopilus orcellus has been given to forms of C. prunulus with a white,
slightly sticky cap, but it is no longer regarded as a distinct species by most mycologists.

Rhodocybe nuciolens
CAP 2-7 (11) cm broad, broadly convex becoming plane or slightly depressed or some¬
times slightly umbonate; surface smooth, not viscid, dull pinkish to pinkish-brown,
pinkish-tan, or sometimes cinnamon-brown to brick-colored, fading as it dries (especially
toward margin), sometimes with watery spots or bands. Flesh white or pale ochre; odor
usually rather spicy. GILLS whitish becoming pinkish as spores mature; close, adnate or
notched or sometimes slightly decurrent. STALK 2-8 (15) cm long, 0.4-1.5 (4) cm thick,
equal or occasionally thicker at base, whitish or colored like cap (but usually paler).
SPORE PRINT rosy-buff; spores 5.5-7.5 * 3.5-4.5 microns, elliptical and warty in side
view, angular in end view.
HABITAT: Solitary, scattered, or in groups or tufts in humus or on rotten wood; common
in central and southern California in the fall and winter, but more widely distributed. It
occurs under both hardwoods and conifers; I find it most often with redwood.

EDIBILITY: Uncertain. Greg Wright of Los Angeles says it has an excellent flavor, but
that one collection caused sweating, chills, and other symptoms of muscarine poisoning.
COMMENTS: The dull pinkish to brick-colored cap and pinkish spores are the distin¬
guishing features of this eminently undistinguished fungus. The gill attachment is quite
variable, leading to confusion with Clitocybe inversa (which has white or buff spores),
candy caps (Lactarius fragilis and L. rufulus), and various Collybias, but the pinkish
spores and pinkish gills at maturity are distinctive. There are several other Rhodocybes
that are difficult to key to genus without examining the spores. These include: R. nitellina,
cap up to 5 cm broad and orange-brown to reddish-brown to pinkish-cinnamon or ochre
with a translucent-striate margin when moist; R. roseiavellanea, one of several uncommon
wood-inhabiting species with a more or less tan cap; R. mundula, cap 3-7 cm broad,
grayish, and bitter-tasting; R. caelata, with a small(l-3 cm) grayish cap and decurrent gills;
and/?, aureicystidiata, also small, but with a darker cap that bruises dark or dingy reddish.

Rhodocybe nuciolens is variable in size, shape, and color, but always has pinkish spores. These are
rather small specimens.
242 ENTOLOMATACEAE

ENTOLOMA
Medium-sized to fairly large terrestrial mushrooms. CAP typically convex to plane or uplifted,
usually not sca/y.GILLS typically attached (usually notched), fairly close to well-spaced, usually
pinkish or flesh-colored in old age. STALK fleshy, smooth or slightly scaly, central. VEIL and
VOLYA absent. SPORE PRINT pinkish to flesh-colored or cinnamon-pinkish. Spores angular.

ENTOLOMA is easily the most conspicuous genus in the Entolomataceae. As defined

here it is comprised of fleshy, pinkish-spored, terrestrial mushrooms that correspond in
shape and stature to the white-spored genus Tricholoma. The cap and stalk are smooth to
only slightly scaly; the gills are usually notched but vary in their attachment (however, they
are not truly free as in Pluteus, and only rarely decurrent as in Clitopilus). In contrast to
Nolanea and Leptonia, the stalk is typically fleshy, but there are numerous nondescript
forms with a “somewhat fleshy” stalk, giving the “lumpers” (see p. 10) a good excuse to
broaden the definition of Entoloma to include both Nolanea and Leptonia.
Because of their larger size Entolomas are much easier to see than Nolaneas and
Leptonias—but they are just as difficult to differentiate, if not more so. Most species are a
forgettable shade of brown, gray, dingy olive, or dull yellow, but a few feature the blue and
violet hues so typical of Leptonia.
Entolomas are almost exclusively terrestrial—a good means of distinguishing them from
deer mushrooms (Pluteus)—and grow mostly in the woods. Some are probably
mycorrhizal (in Europe several are associated exclusively with trees and shrubs of the rose
family—an unusual habitat for mushrooms). In our area they are particularly common
under hardwoods (oak, tanoak, madrone) in the late fall and winter, and are often most
abundant when other mushrooms are rather scarce.
Entolomas should not be eaten. Several species are quite poisonous—e.g., E. lividum
and E. rhodopolium—and the few that are definitely known to be edible are not easy to
recognize, with the possible exception of E. abortivum, an eastern species that frequently
produces aborted or misshapen fruiting bodies.
Entoloma is a fairly large genus even in its strictest sense. Three representative species or
species “complexes” are described here, but according to Entoloma-expert David Largent,
several of our local species are endemic and undescribed.
Key to Entoloma
1. Cap red-brown to orange-brown, cinnamon, or pinkish (see Clitopilus& Rhodocybe, p. 239)
1. Not as above; cap blue, gray, black, grayish-brown, grayish-yellow, olive-brown, whitish, etc.
(but may be dusted pinkish by spores) .2
2. Cap gray to grayish-brown and gills usually decurrent; “aborted” fruiting bodies often found
(infected by mycelium of Armillariella mellea) that are whitish to pinkish-tan and bumpy on
the outside, marbled within; common in eastern North America .E. abortivum
2. Not as above; gills typically not decurrent; “aborted” fruiting bodies absent .3
3. Cap and/or stalk bluish-gray to steel-gray to dark blue, violet-tinged, indigo-blue, or black
when fresh .4
3. Not as above . 6
4. Only the stalk bluish or violet; cap brown to dark brown and smooth; stalk rather slender; found
on wood or ground E. trachysporum var. purpureoviolaceum(see Leptonia carnea, p. 250)
4. Not as above . 5
5. Cap at least slightly viscid when moist; stalk (0.5) 1-3 cm thick; cap blue-gray to steel-gray to
violet-gray to black; common .E. madidum, p. 243
5. Cap and/ or stalk deep indigo-blue; cap not viscid, stalk usually more slender than above; not
common . E. nitidum(see Leptonia carnea, p. 250)
6. Odor nitrous or bleachlike when fresh (crush flesh if unsure) . . . E. nidorosum group, p. 244
6. Odor mild or at least not as above . 7
ENTOLOMA 243

7. Cap white or whitish when fresh . 8

7. Cap not white ..9
8. Spores not angular; rare .(see H ebeloma, p. 463)
8. Spores angular under microscope; not uncommon .
.E. prunuloides& E. speculum (see E. rhodopolium group, below)
9. Gills tinged yellow when young; stalk 1-3 cm thick; mainly found in eastern North America
.E. lividum(see E. rhodopolium group, below)
9. Not as above; widely distributed .E. rhodopolium group &. others, below

Entoloma madidum (Midnight Blue Entoloma) Color Plate 51

CAP (3.5) 5-15 (20) cm broad, obtuse or convex becoming plane or remaining broadly
umbonate, the margin sometimes uplifted in age; surface viscid to slightly lubricous when
moist, smooth or wrinkled, often with a fibrillose or streaked appearance; blue-gray to
nearly black to violet-gray or steel-gray, sometimes fading in age to fuscous or occasionally
with ochraceous stains or paler areas. Flesh thick, firm, white; odor usually farinaceous.
GILLS notched or adnexed or at times appearing free, close, white or tinged cap color,
becoming pinkish or flesh-colored as the spores ripen. STALK 4-13 cm long, (0.5) 1-3 cm
thick, solid, firm, fleshy, fibrillose, equal or with a tapered base or often thicker in middle;
upper portion blue-gray or colored like cap, base whitish to slightly yellowish (but occa¬
sionally white throughout). SPORE PRINT flesh-color to pinkish-cinnamon; spores 6.5-
8.5 (10) x 6-8 microns, nearly round to broadly elliptical but angular (5-6 sided).
HABITAT: Solitary, scattered, or gregarious on ground in woods; widely distributed but
especially common along the west coast in the fall and winter. In our area two slightly
different forms occur—one with conifers (especially redwood), the other with hardwoods
such as tanoak and madrone. The European version is said to grow in grass!
EDIBILITY: Edible, with meaty flesh and a good flavor. However, many fleshy
Entolomas are toxic, so be certain of your identification!
COMMENTS: One of the few Entolomas that can be recognized easily in the field—
the black to bluish-gray or violet-tinged cap, pinkish spores, notched or adnexed gills that
are whitish when young, and firm, fleshy stem render it distinct. It has the aspect of a
Russula, but is not as brittle and does not have white or yellow spores. It is much larger and
fleshier than the small bluish-black Leptonias (L. parva, etc.) and is not nearly as purple
as the blewit (Clitocybe nuda). The viscid or lubricous cap (when moist) separates it from
E. nitidum and Leptonia carnea, which are also more vividly colored. Tricholomaporlen-
tosum is quite similar, but has white spores and a white stem.

Entoloma rhodopolium group

CAP 3-12 (15) cm broad, convex or broadly umbonate becoming plane or slightly de¬
pressed; surface smooth, not viscid or only slightly so, gray to grayish-brown to yellowish-
gray, usually paler as it dries; margin often wavy or splitting in age. Flesh firm, white; odor
mild to slightly farinaceous, but not bleachlike. GILLS adnate or notched, whitish or
tinged cap color, becoming pinkish as the spores ripen; fairly close to rather well-spaced.
STALK 4-13 cm long, 0.5-1.5 (2) cm thick, fleshy, equal or tapered below, dry, white or
tinged gray. SPORE PRINT pinkish-salmon to deep flesh-color; spores 8-11 x 7-9
microns, nearly round but angular.
HABITAT: Solitary, scattered, or in groups or clusters onground in woods, mainly under
hardwoods; widely distributed. In California it is quite common in the fall and winter
under alder, but has many look-alikes that occur with oak, tanoak, and madrone.
244 ENTOLOMATACEAE

EDIBILITY: Poisonous—causing vomiting, diarrhea, and abdominal cramps that may

require hospitalization! All fleshy Entolomas should be strictly avoided, although some
are said to be edible—they are difficult to differentiate and not well known.
COMMENTS: The above description will fit a number of medium-sized to large, showy
woodland Entolomas with a smooth, grayish to yellowish or tan cap, fleshy stem, attached
(usually notched) gills, and pinkish spores. They are the most common and conspicuous
of our Entolomas (along with the E. nidorosum group) as well as the most dangerous. As
a group they can be recognized by their color and Tricholoma-like stature (i.e., they are
never as slender as the similarly-colored Nolaneas). Distinguishing between the various
Entolomas within the group (as loosely defined here) is strictly a matter for specialists.
The largest and most infamous of the lot, E. lividum, (also called E. sinuatum), has a tan
to grayish or pallid cap that is 6-20 cm broad or more, pale yellowish gills when young, and
a thick (1 -3 cm) stalk; it is common in eastern North America, but has not yet been reported
from California. Other species include: E. grayanum, a common fleshy eastern species;
E. clypeatum, whose cap is grayish-brown to olive-buff and streaked with darker fibrils;
E. prunuloides, with a large white viscid cap; and E. speculum, slender, with a dry white
or whitish, often umbonate cap. The latter species occurs in California, but the others
may not. (According to Entoloma-expert David Largent, many of California’s large,
fleshy Entolomas are undescribed and probably endemic.)

Entoloma nidorosum group (Nitrous Entoloma)

CAP 3-12 cm broad, convex becoming plane or depressed or occasionally broadly umbo¬
nate; surface smooth, not viscid or slightly so when moist; olive-brown to brownish to
grayish-brown or pale yellowish-brown when moist (but see comments), paler (grayish to
grayish-yellow or even whitish) as it dries; margin often wavy in age. Flesh pallid; odor
distinctly nitrous (bleachlike) when fresh, but fading after sitting awhile; taste rather
rancid-oily. GILLS white or pallid, becoming pinkish as the spores mature; usually
adnexed or notched, sometimes adnate; fairly well-spaced to fairly close. STALK 4-12
cm long, 0.3-1.5 cm thick, more or less equal or narrowed at base, whitish orfaintly yellow,
fibrillose-striate, firm, fleshy, solid. SPORE PRINT pinkish-salmon to deep flesh-color;
spores 7-9 * 6-8 microns, nearly round but angular.
Entoloma nidorosum group. The larger Entolomas are notoriously difficult to identify, but this
species “complex” has a distinctive bleachlike odor. Some are stouter than the specimens shown
here, others are more slender. The gills are usually adnexed or notched.
ENTOLOMA 245

HABITAT: Solitary to scattered or gregarious (sometimes tufted) on ground in woods,

widely distributed. Common in our area under hardwoods in the fall and winter.
EDIBILITY: To be avoided. The very similar E.ferruginans{see comments) isapparently
edible, as it has been mistaken inexplicably (but harmlessly) for the blewit (Clitocybe
nuda), in spite of its unpleasant taste and unpurple color. Several similar Entolomas are
quite poisonous, however (see the E. rhodopolium group).
COMMENTS: The bleachlike odor of fresh specimens is usually quite pronounced, and
serves to separate this species complex from other gray to brown, fleshy Entolomas. The
stature of E. nidorosum is typically rather erect and slender, but the stalk is distinctly
fleshy, in contrast to species of Nolanea. Other Entolomas with a bleachlike odor include:
E. pernitrosum, with a persistent odor (i.e., one that doesn’t fade); and E. ferruginans,
which is one of the commonest gilled mushrooms in the live oak woodlands of southern
California, and the most common Entoloma there (it also occurs in our area and probably
throughout the state). It differs from E. nidorosum in its more variable cap color (hazel
to buff, pale yellow, olive-brown, dark brown, or nearly black), thicker stalk (up to 2.5 or
even 3 cm thick) which is often tinged with the cap color, and slightly larger spores. Whether
it is indeed a distinct species endemic to California, or merely a robust version of E. nido¬
rosum, is for the Entoloma-experts to decide.

NOLANEA
Smallish, mostly saprophytic, terrestrial mushrooms. CAP usually smooth, often silky when dry,
typically conical, bell-shaped, or distinctly umbonate, at least when young, but sometimes convex.
GILLS typically attached (adnate to adnexed or sometimes decurrent), but sometimes appearing
free. STALK typically slender and fragile or cartilaginous; usually hollow and easily splitting;
central. VEIL and VOLVA absent. SPORE PRINT pinkish to flesh-colored to salmon- or
cinnamon-pinkish. Spores angular (sometimes shaped like jacks or ice cubes!), not amyloid.

THESE are small, drab, gray to brownish mushrooms with a thin, cartilaginous or fragile
stem and a conical to bell-shaped or distinctly umbonate (“nippled”) cap. The shape and
stature of the fruiting body are reminiscent of the white-spored genus Mycena, but the
spore color, of course, is pinkish. The cap is convex to plane in a few species, but never
(or only rarely) umbilicate as in Leptonia. Entoloma differs by its fleshy stem, but this is a
somewhat ambiguous character, and the two genera intergrade.
Nolanea species are differentiated largely on microscopic characteristics such as the
presence or absence of cystidia (sterile cells) on the edges of the gills. They are especially
common under conifers, but also occur with hardwoods and in lawns and pastures.
Most are of unknown edibility but some, such as N. verna, are said to be poisonous.
Three widely distributed species are described here. They are similar to one another in
appearance but differ in habitat and season. Two colorful and distinctive eastern species
are also included in the key, plus the infrequently encountered genus Eccilia.
Key to Nolanea
1. Odor sweet, like a candy store or bubble gum (break open the cap if unsure) .2
1. Odor mild or distinctive, but not as above . 3
2. Stalk (and usually the cap) yellowish . N. icterina(see N. verna group, p.247)
2. Cap and stalk more or less grayish-brown .... N. fructufragrans(see N. verna group, p. 247)
3. Cap pink to yellow or orange and conical or bell-shaped (or if expanding in age then retaining
a distinct umbo); found in eastern North America .4
3. Not as above; cap not usually brightly colored . 5
4. Cap yellow to yellow-orange .N. murraii
4. Cap more or less salmon-colored .N. salmonea
246 ENTOLOMATACEAE

5. Stalk minutely velvety; cap reddish-brown to blackish (at least at center); odor usually strong,
like cucumber or fish. (see Tricholomataceae, p. 129)
5. Not as above . 6
6. Fruiting mainly in the spring, especially in areas with snowfall .TV. verna group, p. 247
6. Not as above . 7
7. Usually growing in grass or open areas; odor of crushed flesh farinaceous (like raw meal or
cucumber) .TV. sericea & others, below
7. Usually growing in woods or under trees; odor mild or various, sometimes farinaceous ... 8
8. Cap conical to bell-shaped, or if expanding somewhat then usually retaining a distinct umbo
.TV. staurospora & others (see TV. sericea, below & TV. verna group, p. 247)
8. Cap broadly conical to convex or plane . 9
9. Gills decurrent . Eccilia
9. Gills not decurrent . TV. stricta & others, below

Nolanea stricta (Strict Nolanea)

CAP 2.5 -7 cm broad, conical or bell-shaped expanding to broadly convex or even plane,
but usually with a distinct, sometimes pointed umbo; surface smooth, hygrophanous,
watery cinnamon to grayish-brown when moist, paler and often somewhat streaked when
dry. Flesh thin, fragile, odor mild. GILLS broad, adnate to deeply notched, adnexed, or
free; pallid becoming flesh-colored in age, fairly close. STALK5-15 cmlong,2-5 (10) mm
thick, typically long and straight (strict), hollow, fragile, splitting easily, equal or tapered
slightly upward, longitudinally striate; whitish or colored like cap (but usually paler); base
whitish and often downy. SPORE PRINT salmon-pink or pinkish-cinnamon; spores
9-13 * 6-9 microns, elliptical but angular. Cystidia absent on gill edges.
HABITAT: Solitary, scattered, or in small groups on ground or very rotten wood, under
both hardwoods and conifers; widely distributed. I have found it—or a very similar
species—in our area in the late fall, winter, and spring.
EDIBILITY: Unknown—to be avoided.
COMMENTS: Also known as Rhodophyllus strictior and Entoloma strictius, this is one
of a number of attractive butdifficult-to-identify, pinkish-spored, woodland mushrooms
with a long, straight (“strict”), slender stalk that splits or breaks easily, and a gray to brown
cap. Similar species include: TV. hirtipes, with a grayish-brown cap, farinaceous odor, and
cystidia on the gill edges; and Leptonia jubata, with a convex to plane or uplifted, minutely
hairy or velvety cap. For smaller species with a conical or “nippled” cap, see comments
under TV. verna.

Nolanea sericea (Silky Nolanea)

CAP 2-4 (6.5) cm broad, convex becoming plane or with a slight umbo; surface smooth
but not viscid, hygrophanous: dark brown (umber) to dark olive-brown to grayish-brown
or hazel-brown when moist, paler or grayer with a silky, often streaked appearance when
dry. Flesh thin, watery; odor usually farinaceous, at least when crushed. GILLS adnate to
adnexed, notched, or nearly free, pallid to grayish or tinged cap color, then dusted pinkish
with spores. STALK 2.5-5 (10) cm long, (2) 3-8 mm thick, equal or with a slightly enlarged
base, often short, fragile, longitudinally fibrillose-striate and silky when dry; grayish-
brown or colored like cap, the base whitish. SPORE PRINT deep salmon-pinkish or
cinnamon-pinkish; spores 8-13 x 6-9 microns, elliptical but angular-nodulose. Cystidia
absent on gill edges.
HABITAT: Scattered to gregarious in pastures, on lawns or hillsides, in waste places,
sometimes also in woods or under willows; widely distributed and common in our area
from the fall through early spring. Unlike many mushrooms, it tends to fruit during rainy
spells rather than following them.
Nolanea sericea is common in pastures and other grassy places and occasionally turns up in the
woods as well. The cap usually has a silky sheen in dry weather.

EDIBILITY: Unknown.
COMMENTS: Only when this species is found in grassy places can the beginner hope to
distinguish it from the throngs of other brown to grayish Nolaneas, Entolomas, and
Leptonias. Be sure to look for the characteristic silky sheen that develops in dry weather or
as the cap loses moisture. The cap is not normally conical or bell-shaped as in many Nola¬
neas (see N. verna), but may have a slight umbo (the very similar TV. hirtipes has a more
prominent umbo or “nipple” and features cystidia on the gill edges). The pinkish spores and
gray-brown to brown color help separate N. sericea from our other common grass-
inhabiting mushrooms. Other species: N. edulis is a very similar but smaller species that is
common in open places in southern California; N. staurospora grows in grassy as well as
shaded or wooded areas and has a conical to bell-shaped cap that may become convex or
umbonate as it matures, and ranges in color from dark brown to date-brown,
grayish-cinnamon, or grayish-brown, but fades as it dries to tan or paler. Furthermore, it
has “cruciate-nodulose” spores (shaped like stars or jacks). I mention this feature because it
is impossible to appreciate the relief they (the spores) provide unless you’ve spent many
long, tedious hours scrutinizing the round to elliptical spores sported by the overwhelming
majority of fleshy fungi. Apart from the spores, however, it is just another “LBM” that is
easily confused with dozens of other “LBM’s.” Enough said.

Nolanea verna group (S pringtime Nolanea)

CAP 2-5 cm broad, conical or bell-shaped, often with a pointed umbo or “nipple,” or
expanding sometimes to convex or plane (but often retaining the umbo); surface dry,
usually smooth, finely f|brillose and often silky or glossy; color variable: dark to light
brown, yellow-brown, olive-brown, tan, grayish, or yellowish. Flesh thin, odor mild.
GILLS fairly close, adnate to adnexed but often seceding, pale grayish or buff or colored
like cap, becoming salmon or reddish as spores mature. STALK 2.5-10 cm long, 3-7 (10)
mm thick, equal, straight, often flattened or twisted-striate, hollow, fragile and easily
splitting; pallid or more often colored like the cap (or paler). SPORE PRINT rosy-salmon
to flesh-colored; spores 8-11 * 7-8 microns, elliptical but angular.
HABITAT: Widely scattered or in groups on ground in woods, mainly under conifers;
widely distributed and very common in the spring and early summer shortly after the snow
melts (about the same time as morels!). I have seen large fruitings in the Cascades and
Sierra Nevada, but have not yet found it in our area.
EDIBILITY: Said to be poisonous, causing moderate to severe gastrointestinal distress.

247
Nolanea verna group is common during the spring, especially under mountain conifers (often when
the morels are out). Note the conical cap when young.

COMMENTS: Also known as Entoloma vernum, this species or species “complex” has
many look-alikes in the genus Nolanea, but fruits almost exclusively in the spring.
The pinkish spores distinguish it from Mycena, and the slender, cartilaginous stem
separates it from Entoloma. The yellowish form that is common in the Sierra Nevada may
actually be TV. cuneata (according to David Largent). Similar Nolaneas include: N. holo-
coniota, also common in western mountains, with a brown to yellow-brown cap and pale
yellow to pale orangish stalk that is minutely powdered above; theN. mammosa-N. papil-
lata group, with a small (1-4 cm) nippled, bell-shaped to conical cap and long thin stalk,
usually fruiting in the late summer, fall, or winter; andiV. staurospora, with star-shaped
spores (see comments under N. sericea). Also worth mentioning are two small species that
smell like a candy store or “tooty-fruity gum” (Largent): N. fructufragrans, with a more
or less grayish-brown cap and stalk; and N. icterina, with a yellowish to olive-yellow cap
and stalk and adnexed to slightly decurrent gills. None of these are worth eating.

LEPTONIA & Allies

Small to medium-sized, saprophytic mushrooms growing on ground or sometimes on wood. CAP
typically convex to plane, depressed, or umbilicate; often fibrillose or with small scales, and often
blue, black, or violet. GILLS typically attached (adnexed to adnate or slightly decurrent), but
sometimes appearing free. STALK typically slender and fragile or cartilaginous, but sometimes
fleshy; smooth or scaly; often blue to gray, black, or violet. SPORE PRINT pinkish to rosy-pinkish,
flesh-colored, pinkish-cinnamon, etc. Spores angular, not amyloid.

I
IN stature Leptonia corresponds roughly to the white-spored genera Collybia and
Omphalina. The cap is typically convex to plane or umbilicate, rather than conical as in
Nolanea, and the stalk—with a few exceptions—is thin and fragile rather than fleshy as in
Entoloma. Leptonias can sometimes be recognized by their color alone: many species are
breathtakingly blue, vividly violet, or beautifully black, and some, such as L. carnea, have
an iridescent quality that makes them among the most striking of all mushrooms. Others,
however, are gray or brownish like umpteen Entoloma and Nolanea species.
Leptonias fruit throughout the mushroom season, but tend to be most abundant when
or where other mushrooms are scarce. Fairly few fleshy fungi, for instance, are fond of
fruiting under fern fronds in redwood duff, but that is precisely the niche in which Leptonia
thrives in California. In the Pacific Northwest, on the other hand, it is frequent under
alder and western red cedar (also rather unfruitful foraging grounds for fleshy fungi), and

248
LEPTONIA& ALLIES 249

in England and Scotland it is said to favor heaths. Most of the species are terrestrial; a few
grow on rotten wood, but can be separated from small species of Pluteus by their attached,
more widely spaced gills.
Though Leptonias are among our most beautiful mushrooms, little is known of their
edibility. David Largent recognizes over 100 species in his monograph on Leptonia, but
even the distinctively colored ones are rather difficult to identify. A mere four species
are described here, plus one species of Alboleptonia, a small genus of whitish mushrooms
often included in Leptonia.
Key to Leptonia & Allies
1. Cap and/ or stalk blue, blue-black, blue-gray, steel-gray, violet, or black, at least when young
and fresh.2
1. Not as above . 12
2. Cap smooth (not scaly!) and brown to dark brown; stalk blue or violet-blue and often iridescent,
usually at least 4-5 mm thick .(seeEntoloma, p. 242)
2. Not with above features . 3
3. Growing on wood; cap usually fibrillose-scaly; rare L. cyanea& others (see L. carnea, p. 250)
3. Growing on ground and/ or cap smooth .4
4. Stalk slender, hollow, and either fragile or cartilaginous, usually 1-5 mm thick at apex .... 5
4. Stalk usually 5 mm thick or more . 11
5. Fresh stalk fibrillose-scaly and iridescent or metallic . . L. occidentalis(see L. carnea, p. 250)
5. Not as above .6
6. Gills pallid or grayish with distinctly darker (blue-gray to black) edges .
.L. serrulata(see L. parva, p. 251)
6. Not as above (but gills may be entirely blue-gray) . 7
7. Cap and/ or stalk with distinct violet or purple tints when fresh . 8
7. Not as above .9
8. Gills pale yellow when young.L. zanthophylla(see L. nigroviolacea, p. 250)
8. Not as above . L. nigroviolacea & others, p. 250
9. Cap bluish-black to black when fresh (but may fade to brownish or gray in age) . 10
9. Cap yellow-brown to reddish-brown to grayish-brown when fresh L. gracilipes& others, p. 252
10. Gills blue-gray to blue.L. nigra (see L. parva, p. 251)
10. Gills whitish or tinged gray .L. parva & others, p. 251
11. Cap and stalk deep indigo-blue (or stalk apex sometimes with violet tints); known only from
the west coast .L: carnea, p. 250
11. Cap grayish-brown to deep violet; stalk usually paler; found mainly in eastern North America
.L. porphyrophaea (-Entoloma violaceum)
12. Fruiting body entirely white when fresh (but may develop yellowish, ochraceous, or pinkish
tones in age) .A Iboleptonia sericella & others, p. 252
12. Not as above . 13
13. Gills whitish with brown to gray-brown edges L. fuligineomarginata(see L. gracilipes, p. 252)
13. Not as above . 14
14. Edges of gills pink; cap rose-tinted when fresh; rare . L. rosea
14. Not as above . 15
15. Stalk yellow-green, usually staining blue-green when bruised; cap greenish-yellow to yellow-
brown to olive-brown; rare .L. incana
15. Not as above . 16
16. Stalk white; cap umbilicate in age .L. exalbida(see L. gracilipes, p. 252)
16. Not as above . 17
17. Gills decurrent . Eccilia
17. Gills adnexed to adnate or with a very slight decurrent tooth . 18
18. Cap striate and usually umbilicate in age .L. undulatella (see L. gracilipes, p. 252)
18. Not as above; cap not umbilicate.(see AJolanea, p. 245)
250 ENTOLOMATACEAE

Leptonia carnea (Indigo Leptonia) Color Plate 52

CAP 2.5 -7 cm broad, convex becoming broadly umbonate or plane, the margin sometimes
uplifted in old age; surface dry, densely fibrillose or fibrillose-scaly, deep indigo-blue, at
times almost iridescent, scarcely fading. Flesh yellowish or whitish, but tinged violet under
the cuticle; odor typically farinaceous. GILLS adnate to adnexed or notched, pallid or
bluish-gray, slowly becoming pinkish as spores mature. STALK 5-10 cm long, 0.5-1 (1.5)
cm thick, equal or thicker below, dry, densely fibrillose or minutely scaly, deep indigo-blue,
the apex sometimes violet-tinted; base often whitish, or in age yellow-orange within.
SPORE PRINT pinkish; spores 9-13 x 6.5-10.5 microns, elliptical but angular.
HABITAT: Solitary or in small groups in woods; known only from California, and ap¬
parently rare. I have found it several times near or under redwood in late fall and winter,
during otherwise uneventful mushroom hunts. Its deep color camouflages it well.
EDIBILITY: Undetermined, but too rare to be of value.
COMMENTS: The stunning indigo-blue color and fibrillose-scaly cap and stem render
this breathtakingly beautiful mushroom distinct (the color plate does not do it justice). Its
rareness makes it all the more pleasurable to find. Its only rival for intensity of color is
Cortinarius violaceus, which has rust-brown spores and is deep violet. The stem is thicker
than that of the numerous blue-black or violet-black Leptonias (see L. parva and L. nigro-
violacea), but thinner than that of Entoloma madidum. The gills are not purple as in the
blewit (Clitocybe nuda) and the stalk and cap are much scalier. There are several similarly
colored species, including: Entoloma nitidum, with a smoother (less scaly) cap and stalk
and smaller spores (7-9 microns long), not common; Entoloma trachysporum var.
purpureoviolaceum, with a smooth dark brownish cap and metallic blue stalk; L.
occidentalis var. metallica, smaller, with an iridescent or metallic blue, densely scaly-
fibrillose stalk (up to 5 mm thick at apex), mild to slightly fragrant odor, and white to blue-
gray flesh, also not common; and three rare wood-inhabiting species with fibrillose-scaly
stalks: L. cyaneonita, bluish-black, with an iridescent stalk; and L. cyanea and L.
violaceonigra, both violet-tinted, the former with pallid gills and the latter with bluish- or
violet-tinged gills when young.

Leptonia nigroviolacea (Violet-Black Leptonia)

CAP 1-3.5 cm broad, convex or obscurely umbonate becoming plane to slightly umbili-
cate; surface dry, fibrillose, usually with small scales at least at the center; entirely violet-
black or deep violet, or sometimes bluish-black or fading to violet-gray. Flesh thin, whitish
or tinged violet. GILLS fairly well-spaced, white or pale buff or sometimes tinged gray,
becoming pinkish in age as spores mature; attachment variable but usually adnexed to
adnate. STALK 3-8 cm long, 2-5 mm thick, equal or tapered slightly, smooth or fibrillose,
often with a few small scales, violet-black to violet-gray, the extreme apex usually pallid;
hollow; usually with white mycelial down at base. SPORE PRINT rosy-pinkish; spores
9.5-11.5 (13) x 7-8.5 microns, elliptical but angular. Cystidia absent.
HABITAT: Solitary, scattered, or in groups or tufts on ground in woods; widely dis¬
tributed. In our area it is fairly common in the late fall and winter under redwood, but is
not restricted to that habitat.
EDIBILITY: Unknown.
COMMENTS: This is one of several dainty, violet-tinged Leptonias. They are not as
numerous as the blue-black models (see L. parva), but are just as attractive. L. occidentalis
var. metallica (see L. carnea) is similar but has a more iridescent, densely fibrillose-scaly
stalk. Other species which characteristically exhibit violet tints include: L. convexa, whose
LEPTONIA 251

cap has erect scales on a bluish-gray, reddish-brown, or grayish-brown background;

L. zanthophylla, with pale yellow gills when young; and/,. diversa, whose cap is frequently
umbilicate in age and has purple-blue scales on a paler background, and whose gills are
often tinged bluish-gray. For violet-tinted, wood-inhabiting species, see comments under
L. carnea.

Leptonia parva (Blue-Black Leptonia)

CAP 0.7-4 cm broad, broadly convex becoming plane to shallowly depressed or umbili¬
cate; surface dry, finely fibrillose when young, often breaking up into small scales in age;
blue-black to black when fresh, often somewhat paler or grayer in age; margin often finely
striate or fibrillose-striate. Flesh thin, white or tinged cap color. GILLS very slightly
decurrent to adnate or adnexed, but sometimes seceding; whitish or with a slight tinge of
blue-gray, becoming pinkish in old age from spores. STALK 1.5-8 cm long, 2-4(7) mm
thick, more or less equal, smooth or in one form with minute bluish-black scales over
upper half; blackish-blue to bluish-gray, often becoming paler gray in age; base often with
whitish mycelial down. SPORE PRINT rosy-pinkish; spores 8-12 * 6-8 microns, elliptical
but angular. Cystidia typically absent on gill edges.
HABITAT: Scattered or in small groups in forest humus, widely distributed. This species
and its numerous look-alikes (see below) are especially common in coastal California in
redwood duff (often under ferns), and in the Pacific Northwest under western red cedar.
They occur throughout the mushroom season in our area, but are easily overlooked
because of their dark color.
EDIBILITY: Unknown, but too small to be of value.
COMMENTS: There are a number of petite blue to black or violet-tinted Leptonias that
will more or less fit the above description (David Largent recognizes over 100 Leptonia
species in his monograph). They are quite difficult to distinguish in the field, but you
needn’t know their exact identities to appreciate their beauty. Entoloma madidum, E. niti-
dum, and Leptonia carnea are similar in color but are larger and have thicker, fleshier
stems. L. parva is characterized by its whitish gills in youth and nearly smooth cap that
becomes minutely scaly (especially toward the center) in age. Some of its more common or
distinctive cousins include: L. decolorans, very similar and common, gills often bluish-
gray with cystidia on their edges; L. corvina, also similar and common, but with a cap that
fades to brown in age, except at the center;/,, rectangula, with a black cap that is nearly

Left: Leptonia nigroviolacea, a dainty violet-tinged species. Right: Leptonia parva, a common black
or steel-gray species.
Left: Leptonia parva, or a very similar species. Right: Gill detail in Leptonia serrulata, showing the
finely scalloped black edges characteristic of that species.

rectangular in profile when young and deeply depressed (but not umbilicate) in age, and
gray gills; L. nigra, a beautiful species with evenly blue-gray gills and minutely hairy blue-
black cap that is convex to plane but never umbilicate, fairly common; L. serrulata, gills
white to gray with dark blue to blue-black, minutely serrated edges (see above photo¬
graph), common and widely distributed; and L. cupressa, cap blackish but gills and
stalk brownish-gray, found mainly under cypress, not common. All of these are typically
terrestrial. For similar wood-inhabiting species or those with iridescent, fibrillose-scaly
stalks, see comments under L. carnea, and for small terrestrial species with a distinct violet
tinge, see L. nigroviolacea.

Leptonia gracilipes
CAP 1.5 -5 cm broad, convex becoming plane to shallowly depressed or umbilicate; sur¬
face dry, minutely scaly toward the center, the margin finely fibrillose or smooth; dark
yellow-brown to grayish-brown or dark brown, often fading in age to paler brown or even
orange-brown, the center usually darker. Flesh thin, fragile, grayish- or olive-tinted.
GILLS adnexed to adnate or with a slight decurrent tooth, white or pallid when young,
becoming pinkish in age from maturing spores, or sometimes brownish. STALK 2-8 cm
long, 1-5 mm thick, equal, often flattened or grooved, smooth, dark blue-gray fading to
blue-gray or gray in age. SPORE PRINT rosy-pinkish; spores 8-12 * 6.5-8 microns,
elliptical-angular.
HABITAT: Scattered or in small groups in humus under ferns, alder, and conifers; widely
distributed. I have found it several times in our area under redwood, in the fall and winter,
when few other mushrooms were out and about.
EDIBILITY: Unknown.
COMMENTS: The slender blue-gray stalk and yellow-brown to grayish-brown cap
characterize this rather forgettable species. Its stature is similar to that of L. parva, but the
cap is never bluish-black. Other species: L. asprella is similar, but has grayer gills and a
grayer cap; L. vinaceobrunnea has a blue-gray stalk and reddish-brown to vinaceous-
brown cap; L. fuligineo-marginata has a violet-brown to dark reddish-brown to grayish-
brown cap and stalk, but its gills are white with dark reddish-brown to grayish-brown
edges; L. undulatella is brown to yellow-brown with a striate cap; and L. exalbida has an
umbilicate, dark yellow-brown cap and white stalk.

A Iboleptonia sericella (Little White Leptonia)

CAP 1-3 (5) cm broad, convex becoming plane or centrally depressed; surface smooth,
dry, often silky or finely fibrillose; pure white to translucent white or tinged yellow (espe¬
cially in age), or sometimes even pinkish. Flesh very thin, whitish, fragile. GILLS adnate

252
 Wm.

Alboleptonia sericella is a nondescript fragile whitish mushroom with pinkish gills at maturity.

to slightly decurrent or notched, well-spaced, white, becoming rosy-pinkish in age.

STALK 1.5 -5 cm long, 1-4 mm thick, equal, smooth, fragile, white or discoloring like the
cap. SPORE PRINT bright flesh-color; spores 9-13 * 6-8 microns, elliptical but angular
(nodulose).
HABITAT: Scattered to gregarious in damp soil in woods, thickets, along trails, etc.;
widely distributed. Fairly common in our area in the fall and winter.
EDIBILITY: Unknown.
COMMENTS: Also known as Leptonia sericella and Rhodophyllus sericellus, this is the
only common small white mushroom with pinkish spores, a thin stem, and attached gills.
Its fragility makes it difficult to get specimens home in one piece. Camarophyllus species
and Inocybe geophylla are somewhat similar, but have white and brown spores,
respectively. Other species: A. adnatifolia is pure white with slightly smaller spores; A.
ochracea ages or bruises ochraceous.

PLUTEACEAE spores

FORMERLY called the Volvariaceae, these are small to medium-sized mushrooms with
a central stem, free gills at maturity, and smooth, pinkish to sordid reddish spores. In
the other major pinkish-spored family, the Entolomataceae, the gills are usually attached
to the stem and the spores are angular or longitudinally ridged under the microscope.
There are two principal genera in the Pluteaceae: Volvariella has a volva; Pluteus does
not. A third genus, Chamaeota, is extremely rare and not treated in this book. It lacks a
volva, but has a partial veil that usually forms an annulus (ring) on the stalk.
Because of the free gills and volva in Volvariella, the Pluteaceae are thought to be related
to the white-spored Amanitaceae. Besides the disparity in spore color, however, there is
a fundamental anatomical difference: the gill tissue is convergent in the Pluteaceae,
divergent in the Amanitas (see p. 19).
Key to the Pluteaceae
1. Universal veil and volva absent . 2
1. Universal veil present, typically forming a volva (sack) at base of stalk .4
2. Partial veil present, usually forming an annulus (ring) on the stalk . 3
2. Veil and annulus absent .Pluteus, p. 254
3. Spore print white or with only a slight pinkish tinge . (see Lepiotaceae, p. 293)
3. Spore print distinctly pinkish or deep flesh-color; mainly tropical; rare . Chamaeota

253
254 PLUTEACEAE

4. Spore print pale, with only a slight pinkish tinge; volva whitish .(set Amanita, p. 263)
4. Spore print distinctly pinkish to reddish-cinnamon; volva variously colored Volvariella, p. 258

PLUTEUS
Small to medium-sized, wood-inhabiting mushrooms. CAP convex to plane. Flesh usually soft.
GILLS typically close or crowded and free (at least at maturity), usually pinkish in old age. STALK
typically central and cleanly separable from the cap. VEIL and VOLVA absent. SPORE PRINT
flesh-colored to deep pinkish, pinkish-cinnamon, or sordid reddish. Spores smooth, usually
elliptical. Gill tissue convergent.

THESE pinkish-spored mushrooms have a central stalk and free, close gills and grow
almost exclusively on wood. The wood, however, may be buried or decomposed, making
the mushrooms appear terrestrial. They are most often confused with the pinkish, angular-
spored Entolomataceae, which are usually terrestrial with gills attached to the stem.
Pluteus is frequently encountered but rarely abundant. Most species have soft flesh and
decay rapidly. However, the larger types such as P. petasatus and P. cervinus are good
edibles when fresh and firm. P. salicinus and P. cyanopus may contain traces of psilocybin
and/ or psilocin; the edibility of many of the smaller species is unknown.
It is a fairly sizable genus, with over 50 species in North America and more than 20 in
California. As they are segregated primarily on microscopic features such as the structure
of the cap cuticle and the shape of sterile cells (cystidia) on the gills, only five species are
described here.
Key to Pluteus
l. Cap red to orange, fading to yellow .P. aurantiorugosus(see P. lutescens, p. 257)
1. Not as above (cap may be yellow but never red or orange) .2
2. Stalk yellow . 3
2. Stalk not yellow. 5
3. Cap velvety or granulose; stalk white or pinkish when young, yellowish only in age.
.P. flavofuligineus, p.258
3. N ot as above; stalk yellow even when young .4
4. Cap brown or olive-brown .P. lutescens, p. 257
4. Cap yellow or ochre .P. admirabilis(see P. lutescens, p. 257)
5. Cap white . P. pellitus (see P. cervinus, p. 255)
5. Cap not white (but may have colored fibrils or scales on a whitish background) .6
6. Gill edges dark brown to nearly black; usually found on conifers .
.P. atromarginatus (see P. cervinus, p. 255)
6. Not as above; gill edges same color as gills or paler (whitish, pinkish, etc.) . 7
7. Base of stalk blue- or greenish-stained in age or when bruised .
.P. salicinus & P. cyanopus (see P. longistriatus, p. 257)
7. Not as above . 8
8. Cap plushlike, velvety, or granulose but not conspicuously striate .
. P. flavofuligineus & others, p. 258
8. Cap not velvety or granulose, or if so, then also conspicuously striate .9
9. Cap 3-12 cm broad or more, not striate; stalk 4 mm thick or more . 10
9. Cap 5 cm broad or less, sometimes striate; stalk 5 mm thick or less . 11
10. Cap whitish or pallid with brownish fibrils or scales; often growing in clusters (and often
appearing terrestrial) .P. petasatus, p. 255
10. Cap dark brown to pale brown, grayish-brown, etc., sometimes with a fibrillose-streaked
appearance .P. cervinus & others, p. 255
11. Cap yellow or yellowish .P. leoninus(see P. lutescens, p. 257)
11. Cap not yellow.P. longistriatus & many others, p. 257
Pluteus petasatus is a robust species with small scales or fibrils on the cap. It often grows in clusters,
and like other species of Pluteus, has free crowded gills.

Pluteus petasatus
CAP 4-15 (20) cm broad, convex, or becoming broadly umbonate to plane in age; surface
usually not viscid, whitish with a brownish to grayish center or with darker (brown to
grayish-brown) fibrils or scales; margin usually whitish. Flesh fairly thick and firm, white;
odor mild or radishlike. GILLS crowded, broad, free at least in age, whitish for a long time,
then eventually pinkish. STALK 4-10 cm long, 0.7-3 cm thick, equal or swollen above or in
the middle; firm, whitish or discoloring below in age; sometimes streaked with fibrils.
SPORE PRINT pinkish to deep flesh-color; spores 6-10 * 4-6 microns, elliptical, smooth.
Cystidia on faces of gills with long necks and “horns.”
HABITAT: Gregarious or clustered on sawdust or wood chips in gardens, along roads,
etc. (often appearing terrestrial); widely distributed. It is fairly common in our area
during the mushroom season and sometimes even fruits in the summer.
EDIBILITY: Edible and very good—the best of the genus for the table. It is much firmer
and meatier than the better known P. cervinus.
COMMENTS: The robust stature, pale cap with darker fibrils or scales (or a darker
center), and tendency to grow in clusters distinguish this species from P. cervinus. Because
the gills remain white for so long it is liable to be mistaken for a white-spored mushroom,
but the free crowded gills and absence of a veil point to Pluteus. P. magnus (see P. cer¬
vinus) can also be robust, but has a dark brown to nearly black cap.

Pluteus cervinus (Deer Mushroom; Fawn Mushroom)

CAP 3-12(15) cm broad, obtuse or convex becoming broadly convex to broadly umbonate
or plane; surface smooth or radially streaked with fibrils, slightly viscid when moist and
often somewhat wrinkled when young; dark brown to pale brown to grayish-brown or
dingy fawn, the margin sometimes paler. Flesh soft, white; odor usually radishlike. GILLS
close or crowded, broad, soft, white becoming pinkish, finally dingy reddish or flesh-
colored; free at maturity. STALK 5-13 cm long, 0.5-2 (2.5) cm thick, equal or thicker at
base, dry, white or with grayish to brownish longitudinal fibrils. SPORE PRINT flesh-
colored to pinkish-brown; spores 5-8 * 4-6 microns, elliptical, smooth. Cystidia on faces of
gills with long necks and 2-4 “horns.”

255
Pluteus cervinus, often called the deer mushroom, is a common species with a brown cap and free
crowded gills that turn pinkish in old age.

HABITAT: Solitary or in groups on decaying wood, debris, sawdust piles, or humus rich
in lignin; widely distributed and common. In our area it usually appears with Pleurotus
ostreatus after the very first fall rains, then continues to fruit sporadically through the
remainder of the mushroom season. It is partial to (but not restricted to) hardwoods,
but P. atromarginatus (see comments) is common in our area on conifers.
EDIBILITY: Edible and quite good when fresh and firm, but the flaccid or waterlogged
specimens one usually finds are apt to be insipid.
COMMENTS: Also known as P. atricapillus, the deer mushroom is our most common
and conspicuous Pluteus. It is rather nondescript, but can be safely identified by its brown
cap, pinkish spores, close free gills, absence of a veil, and growth on wood. The cap color
is quite variable, but typically some shade of brown. Special care should be taken not to
confuse it with poisonous Entoloma species, which typically have attached (often notched)
gills and grow on the ground. P. cervinus, however, may appear terrestrial and have gills
which are slightly attached when young, so it’s a good precaution to eat only those that
are clearly growing on wood. There are several similar Pluteus species, all apparently
edible, including: P. magnus, more robust (stalk 1-3 cm thick) and with a dark, frequently
wrinkled cap, often growing in clusters on sawdust piles, etc.; P. atromarginatus, common
throughout much of the West on wood and debris of conifers, with a brown to dark brown
cap and dark brown to black gill edges (see photograph!); and P. pellitus, which has a
white cap (sometimes brown at center) and grows on dead hardwoods. The latter looks
like a destroying angel (A manita ocreata, A. virosa, etc.) from a distance, but lacksa volva
and has pinkish gills in age (and pinkish spores). It occurs in our area, but is rare.

Gill detail in Pluteus atromarginatus. This species resembles P. cervinus, but has dark-edged gills,
as shown here.
Left: Pluteus lutescens, a yellow-stemmed species. Right: Pluteus longistriatus, one of several
small and nondescript, infrequently-encountered species.

Pluteus lutescens (Yellow-Stemmed Pluteus)

CAP 1.5 -5 cm broad, convex becoming broadly umbonate or plane; surface sometimes
wrinkled at the center, not viscid, dark brown to olive-brown to yellowish-brown or
yellowish-olive. Flesh thin, white or pale yellow; odor mild. GILLS free at maturity, fairly
close, broad, whitish to pale yellow, but finally pinkish from ripening spores. STALK 2-7
cm long, 2-6 mm thick, more or less equal, straight or curved, fragile; pale yellow, the base
usually brighter yellow. SPORE PRINT pinkish to deep flesh-color; spores 6-7 * 5-6
microns, nearly round, smooth. Cystidia on faces of gills club-shaped to flask-shaped.
HABITAT: Solitary or in small groups on rotting hardwood logs, sticks, and debris;
widely distributed. Occasional in our area in the late fall and winter, especially on dead oak.
EDIBILITY: Said to be edible, but too small and infrequent to bother with.
COMMENTS: A fragile but beautiful Pluteus, easily identified by its yellow stem and
brownish to olive cap plus the free gills and pinkish spores. P. nanus var. lutescens is a passe
pseudonym. Other colorful species include: P. admirabilis, very similar but with a yellow to
ochre cap and yellow stalk, especially common in eastern North America; P. leoninus,
similar to P. admirabilis, but with a white stalk when young; and P. aurantiorugosus
(-P. coccineus), a beautiful but rare species with a bright red to orange-red cap that fades
to bright yellow as it ages, plus a white to yellow or orange-yellow stalk.

Pluteus longistriatus (Pleated Pluteus)

CAP 15 -5 cm broad, convex to plane or with a slightly depressed center; surface sometimes
minutely scaly or granulose, at least at the center, not viscid; gray to brownish-gray,
conspicuously striate in age. Flesh very thin, soft, pallid; odor mild. GILLS close, soft,
whitish becoming pinkish in age; free at maturity. STALK 2-8 cm long, 1.5-3 mm thick,
finely fibrillose-striate, pallid or tinged cap color, equal or with an enlarged base, straight
or curved, hollow. SPORE PRINT pinkish to deep flesh-color; spores 6-7.5 * 5-5.5
microns, nearly round, smooth. Cystidia on gills club-shaped or flask-shaped.
HABITAT: Solitary, scattered, or in small groups on decaying branches and sticks of
hardwoods; widely distributed but not common. I have found it several times in our area
on oak debris, and also on wet wood in a bathroom!
EDIBILITY: Unknown.

257
258 PLUTEACEAE

COMMENTS: This is one of many small, nondescript Pluteus species that are unlikely
to catch the eye of the average mushroom hunter except when they are sharing a log with
more spectacular fare (such as Pleurotus ostreatus). The conspicuously striate cap is the
principal fieldmark of this species. Others include: P. californicus, cap hazel to hazel-
brown or greenish-gray, with a striate margin when moist and a reddish-brown spore
print; P. seticeps, with a brown striate cap; P. cyanopus, with a chestnut- to cinnamon-
brown, faintly striate cap and hazel to grayish-olive stalk; andP. salicinus, with a grayish-
brown to greenish- or bluish-gray, non-striate cap and whitish stalk that stains blue at the
base. None of these are worth eating. P. chrysophaeus should also be mentioned.

Pluteus flavofuligineus
CAP 2-7 cm broad, convex becoming broadly umbonate or broadly convex to plane;
surface not viscid, appearing velvety to minutely granulose; dark brown or olive-brown
when young, developing yellow tones in age from the margin inward (eventually often
entirely yellow or ochre-yellow or yellowish with a brownish center). Flesh thin, pallid;
odor mild. GILLS free but sometimes appearing adnexed; close, whitish or tinged yellow
when young, becoming pinkish as spores mature. STALK 4-11 cm long, 4-6 (8) mm
thick, equal or tapering upward, smooth; whitish or pinkish when young, usually becoming
yellowish in old age. SPORE PRINT pinkish to deep flesh-color; spores 6-7 * 4.5-6
microns, nearly round, smooth. Cystidia on gills flask-shaped.
HABITAT: Solitary or in small groups on rotting hardwood logs, branches, etc.; widely
distributed, but not common in the West. I find it every winter in our oak woodlands.
EDIBILITY: Unknown.
COMMENTS: The velvety to granulose, brown or yellow cap combined with the free gills,
pinkish spores, and growth on dead wood are good field marks. The stalk is often rather tall
for a Pluteus—as much as three times as long as the diameter of the cap! P. lutescens is
somewhat similar, but has a clear yellow stalk even when young, and does not have a velvety
cap. Other species: P. granularis is also similar, but has a velvety brown cap and stem.

VOLVARIELLA

Small to medium-large, saprophytic or parasitic mushrooms found on wood, soil, humus, or other
mushrooms. CAP oval to convex or plane, sometimes viscid. GILLS free at maturity, close, pallid
becoming pinkish to flesh-colored or sordid reddish at maturity. STALK central, usually hollow,
cleanly separable from cap. VEIL universal, membranous, forming a saclike volva at base of stalk.
SPORE PRINT pinkish to deep flesh-color or sordid reddish. Spores smooth, usually elliptical.
Gill tissue convergent.

THE pinkish to reddish spores and presence of a volva separate Volvariella (formerly
Volvaria) from all other mushrooms. The volva is always saclike and there is no partial
veil or annulus. But for the spore color this genus resembles A manita, and young specimens
with pallid gills are often mistaken for that genus. Confusion with Pluteus is also possible
if the volva is overlooked or destroyed.
Several Volvariellas are edible. In fact, the paddy straw mushroom, V. volvacea, is to the
tropics what Agaricus bisporus is to the temperate zone—the principal mushroom of
commerce. It is cultivated widely in southern Asia—usually on straw in rice paddies—and
shipped abroad so you can pay exhorbitant prices for it in exotic food stores. I was
presented a can of it ten years ago, on some unforgettable occasion I can no longer
remember. I am sorry to say it has languished in the back of my cupboard ever since—
there are always so many fresh mushrooms in my refrigerator!
VOLVARIELLA 259

Volvariella is principally a tropical genus, and fewer than a dozen species occur in
temperate North America. They frequent forests, cultivated fields, gardens, straw heaps,
and greenhouses; a few grow on wood and one is parasitic on other mushrooms. Only
one species, V. speciosa, is truly common; several others are encountered infrequently.

Key to Volvariella
1. Growing on other mushrooms . V. surrecta(see V. bombycina, p. 261)
1. Growing on ground, wood, compost, in greenhouses, etc. 2
2. Cap typically 5-20 cm broad when mature . 3
2. Cap typically 2-5 (6) cm broad when mature . 5
3. Growing on wood; cap covered with silky fibrils; rare . V. bombycina, p. 261
3. Not as above . 4
4. Volva brown or grayish-brown (at least the upper portion or edge); cap not normally viscid;
found in tropics, subtropics, and warm environments V. volvacea{see V. bombycina, p,261)
4. Volva white to pale gray; cap viscid when moist; very common in temperate zone in cultivated
fields, gardens, roadsides, manure, etc.V. speciosa, below
5. Volva white, with long hairs; cap gray to grayish-brown V. villosavolva(see V. smithii, p. 261)
5. Not as above; volva without long hairs . 6
6. Cap gray to pinkish-gray; volva brown to grayish .V. taylori(see V. smithii, p. 261)
6. Cap whitish when fresh, but may discolor overall in age and center often tinged another color 7
7. Volva ochre-stained to brownish .V. smithii, p. 261
7. Volva white to grayish . 8
8. Stalk minutely hairy (pubescent); cap margin not striate . V. hypopithys, p. 260
8. Stalk not pubescent; cap margin often striate in age ... V. pusilla (see V. hypopithys, p. 260)

Volvariella speciosa (Common Volvariella) Color Plate 68

CAP 5-15 cm broad, at first oval, then convex to plane or broadly umbonate; surface
smooth or occasionally with patches of universal veil tissue, viscid when moist; dull white to
gray or grayish-brown (rarely fulvous), or often whitish with a darker center; often with a
metallic luster when dry; margin sometimes striate. Flesh soft, white. GILLS crowded,
broad, free, white becoming flesh-colored and finally sordid reddish. ST ALK 5-20 cm long,
1-2.5 cm thick, equal or thicker at base, dry, whitish, more or less smooth. UNIVERSAL
VEIL membranous, forming a white to pale grayish saclike volva at base of stalk; volva
often buried in soil, sometimes inconspicuous. SPORE PRINT deep flesh-color to
pinkish-brown; spores 11.5-21 * 7-12 microns, elliptical, smooth.
HABITAT: Solitary to scattered or gregarious in cultivated soil—gardens, vacant lots,
roadsides, manure, compost, straw heaps, fallow and planted fields, etc.—occasionally
also in the woods; widely distributed. It occurs year-round in our area, but is especially
common in the spring; I’ve seen enormous fruitings in Brussels sprouts fields. In the San
Joaquin Valley I’ve seen fallow fields littered with thousands of Volvariellas.
EDIBILITY: Edible, but mediocre. It was once thought to be poisonous, perhaps due to
confusion with Amanita. Should you try it, be sure to take a spore print—some
Amanitas can have pinkish gills in old age!
COMMENTS: This mushroom can cause quite a stir when it appears, bold and uninvited,
in the middle of your cabbage patch. It is our only common Volvariella, and is recognized
as such by its saclike volva, absence of a ring (annulus), and deep pinkish spores. It is quite
attractive when it first emerges, but quickly becomes flaccid and waterlogged. V. speciosa
var. gloicephala is said to differ from the typical variety in its darker (pearl-gray to fulvous)
pileus (cap) with a striate margin, and smaller spores. One dismal day of illegal trespassing
Left: A young specimen of Volvariella speciosa (see color plate for older ones). Right: Volvariella
hypopithys is a slender white species with silky hairs on the cap.

in a smelly old Brussels sprouts field yielded the following: twenty soggy specimens with
pale pileus and non-striate margin; six soggy specimens with dark pileus and striate
margin; five soggy specimens with dark pileus and non-striate margin; and eleven soggy
specimens with pale pileus and striate margin. If you can detect a meaningful mycelial
thread running through all of this, then you are a better mycologist than I... *

Volvariella hypopithys (Petite White Volvariella)

CAP 2-6 cm broad, at first somewhat bell-shaped, then convex to nearly plane; surface
dry, fibrillose (with fine silky hairs that sometimes form small scales), pure white or tinged
yellowish at the center, but sometimes discoloring overall in old age; margin usually
fringed with long, silky white hairs and not striate or only faintly so. Flesh thin, soft, white.
GILLS fairly close, free (at least at maturity), white becoming dingy pinkish or flesh-
colored. ST ALK 2-9 cm long, 2-5 mm thick, equal or slightly thicker below, fragile, white
or pallid; densely pubescent (clothed with short or long silky hairs) at least over the middle
and upper portion. UNIVERSAL VEIL membranous, forming a small, often incon¬
spicuous or barely visible volva at base of stalk; volva white, saclike, usually lobed.
SPORE PRINT pinkish or flesh-colored; spores 6-8.5 (10) * 3.5-6 microns, broadly
elliptical, smooth.
HABITAT: Solitary or in small groups in woodland humus; widely distributed but rare.
I have found it only once in our area, under live oak, in February.
EDIBILITY: Said to be edible, but much too small to be worth eating.
COMMENTS: This rare and beautiful Volvariella is easily recognized by its petite
dimensions, white color, silky-fibrillose cap, and white pubescent stalk. The inconspi¬
cuous white volva and slender stature (see photograph) are also distinctive, and help to
separate it from V. smithii, which is slightly stockier and has a more prominent, brownish-
stained volva. Another widely distributed but rare white species, V.pusilla(-V.parvula),
is even more petite than V. hypopithys. It has a slightly larger white or grayish volva, a
non-pubescent stalk, and a striate cap margin in age, and is more frequent in gardens,
greenhouses, lawns, and roadsides than in the woods. See also the species under V. smithii.

*Just be thankful, as I am, that I didn’t measure the spores!

260
Volvariella smithii is a small rare species with a prominent volva. Note size in relation to cypress
cone at upper right.

Volvariella smithii (Smith’s Volvariella)

CAP 2-5 cm broad, broadly conical to convex or plane; surface not viscid, usually some¬
what fibrillose at the margin; white, but often tinged buff to pinkish-buff at center. Flesh
rather soft, white. GILLS white becoming pinkish, broad, close, free (at least in age).
STALK3 -5 cm long, 3-7 mm thick, equal or enlarged at base, white, covered with fine white
hairs (pubescent). UNIVERSAL VEIL membranous, forming a large, prominent, saclike
volva at base of stalk; volva brownish to ochre or ochre-stained, usually lobed. SPORE
PRINT pinkish to sordid flesh-color; spores 4.5-7 x 3-4 microns, elliptical, smooth.
HABITAT: Solitary or in small groups in soil or humus in woods; known only from the
west coast, rare. The specimens in the photograph were growing under pine at New
Brighton Beach State Park near Aptos, California, in February.
EDIBILITY: Unknown, but too small and too rare for anyone to care.
COMMENTS: This is one of several small, uncommon, white to grayish Volvariellas. It is
also one of several mushrooms to be named after Alexander Smith, the foremostauthority
on North American mushrooms. The brownish to ochraceous-stained volva, whitish cap,
and small size distinguish it. Other small species include: V. taylori, widely distributed but
rare, with a gray to pinkish-gray cap and a brown to grayish volva; and V. villosavolva
of eastern North America, with a gray to grayish-brown cap and a white volva that has
long mycelial hairs. For small, pure white Volvariellas, see V. hypopithys.

Volvariella hombycina (Silky Volvariella)

CAP 5-20 cm broad, oval becoming bell-shaped or convex and in old age sometimes
nearly plane; surface dry and covered with long silky fibrils (usually more coarsely fibril¬
lose at margin), white to yellowish (often palest at margin). Flesh thin and rather soft or
flaccid, white. GILLS crowded, free, broad, white when young becoming flesh-colored or
pinkish as the spores mature. STALK 6-20 cm long, 1-3 cm thick, usually tapered upward
or enlarged below, often curved, smooth, white, firm. UNIVERSAL VEIL membranous,
often areolate or scaly, forming a thick, long (deep), saclike volva which sheathes the base
of the stalk; volva white to yellowish or dingy brown, often lobed. SPORE PRINT
pinkish to deep flesh-color; spores 6.5-10 x 4.5-6.5 microns, elliptical, smooth.

261
262 PLUTEACEAE

HABITAT: Solitary or in small groups on dead hardwoods or fruiting from wounds in

living trees; rare but conspicuous, found mostly in warm weather. It is widely distributed
east of the Mississippi and has also been found in riparian woodlands in New Mexico and
southern California. Elm and maple are among its favorite hosts; it also grows on beech,
oak, magnolia, and various other hardwoods. Our rainy season is apparently too cold for it.
EDIBILITY: Edible, and according to reports, choice. I haven’t tried it.
COMMENTS: It is easier to identify this striking mushroom than to find it. The silky
cap, pale color, deep saclike volva, and growth on wood make it unmistakable. The latter
feature plus the pinkish spores (and mature gills) separate it from Amanita, which also has
a volva. A yellow-capped variety has been found in Florida. Other species: V. volvacea,
the edible “Paddy Straw Mushroom,” sometimes turns up in greenhouses or straw and
compost in warmer climates, and is fairly common outdoors along the Gulf Coast. It
has a brown or partly brown volva and a gray to dark brown or blackish, often streaked
cap. V. surrecta is a rare but distinctive northern species that grows parasitically on
other mushrooms, particularly Clitocybe nebularis. It is small to medium-sized, with
a white to grayish cap and an ample volva that is usually lobed. Since its favorite host is
common in California, it may occur there rarely.

AMANITACEAE
MEMBERS of this family have white spores, white to pale-colored gills, a universal veil
that envelops the young mushrooms, and in most cases, a volva. Many are also furnished
with a partial veil and the stalk is cleanly separable from the cap.
There are two genera: In Amanita the universal veil is membranous, warty, powdery,
or cottony; in Limacella it is glutinous, manifesting itself as a layer of slime on the cap
and sometimes the stalk.
The Amanitaceae are most likely to be confused with the white-spored Lepiotaceae,
which have neither volva nor viscid cap. Microscopically the two families are distinct by
virtue of the amyloid or non-amyloid (but not dextrinoid) spores and divergent gill tissue
of the Amanitaceae, as opposed to the typically dextrinoid spores and parallel to inter¬
woven gill tissue of the Lepiotaceae (see p. 19).
Amanita is of paramount importance to toadstool-testers because it contains the
deadliest of all mushrooms, as well as some of the most delicious and beautiful. Limacella
is too rare to be of culinary value.

Amanita “eggs” resemble puffballs while still enveloped by the universal veil. However, when sliced
open lengthwise (perpendicular to the ground), they reveal the embryonic outline of cap, gills, and
stalk (left and center), while a puffball (right) is solid within. Note difference in shape between the
deadly poisonous A. phalloides “egg” (center) and the edible A. calyptrata “egg” (left).
AMANITACEAE 263

Key to the Amanitaceae

1. V olva present at base of stalk as a sack, free collar, or series of concentric rings A manita, below
1. Volva absent or indistinct..2
2. Cap usually viscid or slimy when moist, without warts or veil material; stalk sometimes slimy
also .Limacella, p. 291
2. Cap usually with warts or universal veil material, often dry or slightly viscid but not usually
slimy; stalk never slimy .Amanita, below

AMANITA (The Amanitas)

Medium-sized to large terrestrial fungi found mostly in woods. CAP smooth (bald) or with warts or
a cottony patch or other veil tissue. GILLS typically white, creamy, yellow, or pale gray, close,
attached or free. STALK central, usually hollow or stuffed in age and cleanly separable from cap.
PARTIAL VEIL often present and in most species forming a membranous ring on stalk.
UNIVERSAL VEIL present, usually forming a volva at base of stalk in the form of a sack, rim,
collar, or concentric scales. SPORE PRINT white. Spores smooth, amyloid or not amyloid. Gill
tissue divergent, at least when young.

LEARNING to recognize this genus should be an overriding priority for all mushroom
hunters, since Amanitas are responsible for 90% of mushroom-induced fatalities. The
outstanding attributes of any Amanita—what makes even a rotten Amanita not just
another rotten mushroom, but a rotten AMANITA (and therefore worthy of your
attention and respect)—are the white spores, pallid gills, and presence of a universal veil.
The universal veil completely envelops the young mushroom, but breaks as the stalk
elongates, usually forming a volva (sack or collar or scales) at the base of the stalk and often
depositing remnants on the cap in the form of a single large piece of tissue (volval patch) or
many smaller pieces (warts). Obviously, it is important to carefully dig up any unfamiliar
mushroom so as not to miss the volva, if it is present. It is also a good idea to examine the
surrounding soil to be doubly sure pieces of the volva aren’t left behind.
Most Amanitas—including the most dangerous ones—are also furnished with a partial
veil which, upon breaking, often forms a skirtlike ring (annulus) near the top of the stalk.
At one time those species without a partial veil were placed in a separate genus, Amanitop-
sis. A feature emphasized by most mushroom books is that the gills in Amanita are free.
This is not necessarily the case, however, and this feature is not stressed here.
The only other genus of agarics consistently equipped with a volva is Volvariel/a, which
has pinkish spores. Agaricus, Coprinus, and Cortinarius occasionally form a volva, but
have brown to black spores, while Lepiota superficially resembles Amanita but typically
lacks a volva and nearly always has free gills.
Amanita is divided into two large groups (subgenera) based on whether or not the spores
are amyloid. Half of the species described here (including the most dangerous ones!) have
amyloid spores. It should be emphasized, however, that some species with non-amyloid
spores are also poisonous. These two large groups are in turn subdivided according to the
type of volva (see p. 264). If the universal veil is membranous (skinlike), a loose sack or cup
is formed at the base of the stalk and the cap is usually bald or adorned with a volval patch.
If the universal veil is friable (easily crumbling), it manifests itself as a series of concentric
scales or rings around the base of the stem. If the universal veil is semi-friable and
interwoven with the base of the stalk, it will form a collar or free rim (as in A. pantherina),
but not a true sack. When the universal veil is friable or semi-friable, numerous pieces of
tissue (warts) are usually deposited on the cap. These warts are typically white, gray, or
yellow and with a few exceptions (e.g., A. magniverrucata) are readily removable—unlike
the colored scales of an Agaricus or Lepiota. In fact, they are often washed off by rain.
In some Amanitas, such as A. rubescens and A. silvicola, a distinct volva is not formed.
264

Different types of volvas in Amanita. Left to right: A. phalloides, A. pantherina, A. muscaria,

A. rubescens.

However, vestiges of the universal veil on the cap (in the form of warts or cottony tissue)
signify Amanita. In the rare instances in which neither volva nor universal veil remnants are
visible, Amanitas can still be recognized by their uncanny “Amanita aura.” They are so
unequivocally elegant and graceful that you quickly learn to tell an Amanita without
having to dig it up!
Amanita is a study in antithesis. At one extreme are the most poisonous of all
mushrooms—the death cap and destroying angels (A. phalloides, A. ocreata, A. virosa,
etc.). Every fungophile should learn the telltale signs of these deadly fungi (for more de¬
tails, see pp. 892-893). At the other end of the spectrum are three of the most exquisitely
flavored of the fleshy fungi—A. caesarea, A. calyptrata, and A. velosa. The rest of the
Amanitas fall somewhere between these extremes: several are hallucinogenic and/or
poisonous but not normally fatal (A. muscaria and A. pantherina); others are edible but
scarcely incredible (A. pachycolea and A. rubescens)’, still others are of unknown edibility
{A. aspera and A. magniverrucata).
I for one do not subscribe to the wholesale philosophy (as expounded by many
mushroom mentors) that Amanitas should not be eaten under any circumstances. In my
humble fungal opinion, it is just as easy to carelessly overlook the volva and mistake a
deadly A manita for an edible mushroom of another genus as to mistake a deadly Amanita
for the coccora (A. calyptrata) or grisette (A. vaginata). True, it is sheer stupidity to risk
your life for the sake of a single meal, however delectable it may be. But the key word here is
risk—and in the case of a few species such as A. calyptrata, A. caesarea, and A. vaginata, I
don’t consider it a risk for discriminating amateurs to eat them, provided they become
thoroughly familiar with their characteristics and those of their lethal counterparts.
Simplistic slogans or catchwords such as “Do not eat-a the Amanita" often accomplish the
precise opposite of what they intend. Rather than encouraging people to use their eyes and
nose and the gray mass between their ears, to approach each and every mushroom with
discrimination, intelligence, and respect, such adages reinforce people’s desire for
expediency by fostering an unhealthy, mindless reliance on shortcuts and glib generali¬
zations. Those who need simple rules should learn how to play dominoes or Scrabble
rather than eat wild mushrooms. Adages such as the above can even be misconstrued to
read: “If a mushroom isn’t an Amanita it won’t kill you”—a dangerous assumption!
AMANITA 265

Too many people eat and enjoy edible Amanitas for me not to recommend them. But at
the risk of being redundant, let me reiterate some rules of the trade. Unless you are
ABSOLUTELY, INDISPUTABLY, and IRREFUTABLY sure of your Amanita’s
identity, don’t eat it! (The one adage with which I wholeheartedly concur is: “When in
doubt, throw it out!”). If possible, have an experienced collector verify your identification,
and collect the species several times before venturing to eat it. Above all, don’t rely on a
single characteristic (such as striate vs. non-striate margin—see photo on p. 286) to dis¬
tinguish between edible and deadly poisonous species. Each individual mushroom is
subject to a different set of environmental and genetic factors—therefore each will be
slightly different. Only by using a combination of critical characteristics can you rest as¬
sured that you have a savory coccora or grisette instead of a death cap or destroying angel.
Furthermore, don’t assume that two or more Amanitas growing together are the same spe¬
cies. Judge each and every mushroom on its own merits. Finally, always keep in mind the
possibility of encountering a species not described in this book—or any other book!
My reason for lecturing on the Amanitas at such length is that they never fail to attract
attention and admiration. You certainly needn’t eat them to enj oy them, for they are among
the most beautiful and graceful of all fungi, the epitome of impeccability and elegance.
The fly agaric (A. muscaria), with its fiery red cap and white “stars” is the most
spectacular example, of course. Down through the ages it has been compared to bull
testicles and male genitalia and worshipped as the earthly incarnation of infinity, divinity,
and virility (more for its appearance, I suspect, than for its properties). It is one of the
commonest mushrooms of our pine forests, yet one never tires of finding it. The variation in
color, size, shape, and “constellations” is such that each and every one presents a new and
deliciously different feast for the eye. It’s hard to resist taking one or two home to show off to
impressionable neighbors or friends, but never is the ephemerality of life so emphatically
underscored as when they come over the next day to pay their respects, only to find a
writhing mass of beatific maggots where your blazing incarnation of the cosmos had been!
In contrast to the flamboyant splendor of the fly agaric is the subdued and radiant
warmth of the coccora (A. calyptrata). I say “warmth” because the huge eggs are so soft
and cottony that they look positively warm inside. Finding a family of them in rich red
madrone humus is like stumbling onto the nest of a rare and secretive woodland bird. And
watching a coccora “hatch”—the round, orange head emerging from its cottony cocoon—
is like watching the sun rise from a blanket of clouds, a quietly inspiring reaffirmation
of life best experienced alone.
M ost Amanitas are mycorrhizal. As a result, they are most common in the woods or near
trees. Some, however, grow in grass or open ground. Amanita attains its greatest
diversity in the warm temperate zone. In the southeastern United States, for instance, there
is a very diverse and bewildering Amanita flora that is beyond the scope of this book. The
west coast has fewer species but several are endemic. In our region two distinct floras can be
recognized. The first, comprised of northern species, occurs primarily with madrone and
conifers in the late fall and winter. A. muscaria, A. calyptrata, and A. silvicola are
prominent examples. The second group has a more southerly distribution and is associated
with oak. Some members of this group fruit in the fall (e.g., A. phalloides), others in the
late winter and spring (e.g., A. velosa, A. rubescens, A. ocreata—see photo on next page).
Amanita is a far more diverse genus than once thought. Despite a wealth of studies
conducted on the genus, no all-encompassing monograph has been published. (A volume
by David Jenkins is in press.) Particularly perplexing are the “Lepidellas”: those species
with amyloid spores and a friable universal veil that leaves remnants on the cap margin.
Several hundred species of Amanita occur in North America, including many poorly
known or unnamed ones; California has over 25 species. Twenty Amanitas are described
here and many others are keyed out. Their elegance and individuality make them a
fascinating and rewarding group to study, even for those not armed with a microscope. If
your “Amanita” doesn’t key out convincingly, try Cystoderma, Armillaria, and Lepiota.
Three species of Amanita that commonly fruit in the spring in association with live oak: Amanita
velosa (two at left), a small specimen of A. ocreata (top), and A. rubescens (three at right).

Key to Amanita
1. Volva saclike (i.e., forming a true sack that sheathes base of stalk as shown on p. 264); cap
usually bald or with a cottony or membranous patch of universal veil tissue or occasionally
with several patches or non-friable warts . 2
1. Volva collarlike (i.e., intergrown with base of stalk but with a free rim), scaly, warty, powdery, or
indistinct but not saclike (see p. 264); cap often with many small pieces of universal veil tissue
(warts), powder, etc., occasionally with larger pieces . 15
2. V olva tough, thick, large; cap and / or stalk often shaggy, fibrillose, or with cottony patches of veil
tissue; cap white or tinged brown (especially at center), often bruising brown or reddish, margin
often striate in age; stalk similarly colored; annulus (ring) absent; spores oblong or elliptical,
amyloid; fairly common in eastern North America, rare in West A. volvata& close relatives
2. Not with above features . 3
3. Margin of cap distinctly striate (at least when mature); spores not amyloid .4
3. Margin of cap not striate or only faintly so (occasionally striate in age); spores amyloid . . 12
4. Partial veil present when young, usually (but not always!) forming an annulus(ring) on stalk 5
4. Partial veil and annulus absent or rudimentary (but stalk sometimes scaly) . 8
5. Gills and stalk yellow to yellow-orange; cap bright red to orange (but may fade to yellow or
paler in age or sunlight) .A. caesarea group & others, p. 284
5. Not as above; gills and stalk typically white to creamy or very pale yellow .6
6. Volva often small and inconspicuous; cap brown to gray or sometimes nearly white; growing
in mixed woods and under hardwoods in eastern North America . A. spreta
6. Not as above; known only from western North America . 7
7. Cap brown to yellow or whitish, but if brown then usually with a yellow margin; partial veil typi¬
cally (but not always!) forming a prominent skirtlike annulus on stalk A. calyptrata, p. 284
7. Cap variously colored, but the margin not yellow; annulus (ring) usually pressed closely to the
stalk or poorly defined . 48
8. Cap dark brown to gray or grayish-brown . 9
8. Cap white to pale tan, beige, orangish, pinkish, orange-brown, or reddish-brown . 10
9. Fruiting body medium-sized to large; cap dark gray to dark brown when young, often paler
in age and often developing a darker band near inner edge of striations; gill edges usually brown;
known only from the West .A. pachycolea, p. 290
9. Fruiting body medium-sized to rather small and slender; cap usually gray, but sometimes
grayish-brown or brown; gill edges not brown; widely distributed .49
10. Cap white with long striations; widespread, but rare in West A. alba (see A. vaginata, p. 288)
10. Not as above; if cap white then with shorter striations and usually found near western oaks in
winter and spring (or even summer); common . 11
Cap pinkish-tan to orangish, beige, or paler .A. velosa & others, p. 286
Cap orange-brown to reddish-brown, tawny, etc.; common in eastern North America, infrequent
in West .A.fulva, p. 287
12. Cap greenish to yellow-green, brownish-olive, grayish-olive, or nearly white when young,
often duller (dingy tan, etc.) or with a metallic luster in age .A. phalloides, p. 269
12. Not as above; cap usually white or whitish when fresh (but may discolor by maturity) ... 13
AMANITA 267

13. Cap white, but discoloring pinkish, brownish, or yellowish (at least centrally) in age; asso¬
ciated with oak; found in California and the Southwest and Texas.A. ocreata, p. 271
13. Cap usually remaining white or found elsewhere. 14
14. Cap white; partial veil usually forming a distinct annulus (ring) on stalk (which may disappear
in age!); very common in eastern North America, also found in the Pacific Northwest .
.A. virosa& others (see A. ocreata, p. 271)
14. Not as above; partial veil absent or evanescent; found in eastern North America (do not eat!)
.A. peckiana & others
15. Universal veil remnants yellow to grayish-yellow (check cap for warts and base of stalk for
volva); cap not whitish . 16
15. Universal veil remnants not yellow; cap may or may not be whitish .22
16. Partial veil absent; gills yellow; found in eastern North America .
. A. parcivolvata (see A. caesarea, p. 284)
16. Not as above . 17
17. Cap bright red to orange-red (but may fade in age); stalk white .A. muscaria, p.282
17. Cap orange to yellow, yellow-brown, or dark brown; stalk white or yellow. 18
18. Lower stalk sheathed with shaggy scales . (see Armillariak. Allies, p. 189)
18. Not as above . 19
19. Cap salmon to salmon-pink when fresh; found in mountains of eastern U.S.A. wellsii
19. Not as above . 20
20. Base of stalk staining reddish in age or where bruised; found in eastern North America .
.A. flavorubescens (see A. aspera, p. 278)
20. Not with above features (if staining as above, then found in West) . 21
21. Cap yellow to yellow-brown to dark brown; stalk usually white; common along the Pacific
Coast .A. aspera, p. 278
21. Cap yellow-orange to yellow; stalk often colored similarly; fruiting body rather small; common
in eastern North America, rare in the West .A.flavocortia, p. 278
22. Cap pale yellow-green to pale yellow to nearly whitish with thin grayish, whitish-buff, or pinkish
to lavender-gray warts (which may wash off); cap margin not striate; stalk with an abrupt, soft,
rounded bulb at base; spores amyloid; common in eastern North America, especially under
hardwoods but also with conifers .A. citrina (see A. porphyria, p. 279)
22. Not as above . 23
23. Cap brown to olive-brown or paler; stalk lacking grayish patches, terminating in an abrupt basal
bulb that is usually split or chiseled longitudinally; flesh usually staining reddish-brown;
spores amyloid; common in eastern North America (especially under hardwoods); also
reported from the Pacific Northwest (but rare) . . A. brunnescens(see A. porphyria, p. 279)
23. Not as above .24
24. Cap brightly colored (red, orange, or yellow); partial veil present, usually forming an annulus
(ring) on stalk . 25
24. Not as above (cap may be sordid reddish or reddish-brown, pinkish-tan, etc.) .26
25. Volva usually a series of concentric rings at apex of bulbous stalk base, but sometimes only a
single ring or collar; cap medium-sized to large and bright red to orange, apricot, yellow-
orange, or yellow (yellow form rare in coastal California, but common in the Sierra Nevada
and most of eastern North America) .A. muscaria, p. 282
25. Volva usually a single collarat top of basal bulb or often indistinct, but sometimes consisting of
several rings; cap small to medium-sized (occasionally large), usually pale yellow (but some¬
times brighter), at times completely covered by veil material; widespread A. gemmata, p. 281
26. Partial veil absent (check young specimens if possible); cap margin distinctly striate; spores not
amyloid . 27
26. Partial veil present, or if absent, then cap margin not striate; spores amyloid or not .31
27. Cap gray to grayish-brown, brown, or darker . 28
27. Cap pale yellow, orange-buff, salmon, pinkish, tan, or nearly white. 30
28. Cap powdery-mealy; volva if present also mealy A. farinosa(see A. sp. (unidentified), p. 275)
28. Not as above . 29
268 AMANITACEAE

29. Cap gray or sometimes grayish-brown, with or without warts; upper limb of volva usually well-
developed (but falls off easily); common in California .A. constricta, p. 289
29. Cap gray to brown to dark brown to nearly black, usually with warts; upper limb of volva not
well-developed; widespread, but rare in California . . . A. inaurata(see A. constricta, p. 289)
30. Cap yellow to creamy to whitish; volva usually collarlike (with free rim) . A. gemmata, p. 281
30. Cap orange-buff to pale pinkish-orange to pinkish-tan, beige, or sometimes whitish; volva
not typically collarlike; often growing in the open (but near trees) .A. velosa, p. 286
31. Cap entirely brown when young, breaking up into large brown scales in age; flesh in stalk usually
staining orange or saffron (and eventually reddish) when cut (see Lepiota rachodes, p. 297)
31. Not as above . 32
32. Cap with erect, often pyramidal brown warts which usually come off easily; stalk and/ or under¬
side of veil with similar warts; spores usually dextrinoid, not amyloid . (ste Lepiota, p. 293)
32. Not with above features (but may have some of them); common . 33
33. Some part of fruiting body usually with sordid reddish stains (especially the stalk); flesh slowly
staining dingy reddish when bruised or cut; maggot tunnels also reddish A. rubescens, p. 276
33. Not as above .. 34
34. Fresh fruiting body white or whitish (but may age buff, yellowish, brownish, pinkish, etc.) 35
34. Not as above; cap distinctly colored or with colored veil material even when young.42
35. Stalk terminating in a fairly conspicuous bulb, not typically with a rooting portion below the
bulb . 36
35. Stalk without bulb, or if with a bulb, then also with a tapered rooting base below the bulb 40
36. Volva typically present as a distinct free rim (collar) or series of concentric rings at apex of basal
bulb; spores not amyloid .37
36. Not as above; spores amyloid . 38
37. Volva typically consisting of a single tight-fitting collar around bulb apex; cap often tinged
yellowish or brownish at center; often rather slender A. cothurnata(see A. pantherina, p. 280)
37. Volva usually a series of concentric rings; cap white to grayish-white or tinged buff; not
unusually slender.A. muscaria, p. 282
38. Cap surface with rather soft and cottony universal veil tissue, lackingconspicuous warts; known
only from western North America .A. silvicola, p. 273
38. Not as above (if cap cottony, then found elsewhere) . 39
39. Cap usually with brown warts; stalk often rather stout (up to 8 cm long); known only from
California, associated with live oak. A. sp.(unidentified)(see A. rubescens, p. 276)
39. Not as above .40
40. Cap without warts or warts obscure; growing in sand .A. baccata, p. 273
40. Not as above; cap usually with distinct, well-developed warts .41
41. Cap covered with large, exaggerated warts; fairly common with oak and pine, known only from
California .A. magniverrucata, p.274
41. Found elsewhere, or if found in California then warts smaller and often concentrated at center
of cap .A. cokeri & many others (the “Lepidellas”) (see A. magniverrucata, p. 274)
42. Stalk arising from a well-developed cylindrical tojug-shaped, sometimes hollow, underground
“tuber”; rare . (see Squamanita, p. 197)
42. Not as above (but stalk may root deeply or have a bulbous base) .43
43. Cap yellow to creamy, the margin usually striate or tuberculate-striate . . A. gemmata, p. 281
43. Not with above features .44
44. Volva indistinct, powdery, or scaly; cap margin not normally striate .45
44. V olva usually present as a free rim or collar on basal bulb and/ or margin of cap striate in age 47
45. Found in California .46
45. Found in eastern North America (especially common in Southeast) .50
46. U sually found in open ground (pastures, etc.); partial veil often disappearing; cap grayish-white
to gray to brownish-gray, small .A. sp. (unidentified), p. 275
46. Associated with oak; partial veil usually forming a persistent, prominent annulus; cap white
to brownish . A. sp.(unidentified)(see A. rubescens, p. 276)
47. Warts gray and/or stalk gray or with grayish patches; spores amyloid . . A. porphyria, p. 279
47. Stalk white; warts usually white or pallid; spores not amyloid .A. pantherina, p. 280
AMANITA 269

48. Cap brown . A. calyptratoides(see A. calyptrata, p. 284)

48. Cap orangish to salmon, pinkish-tan, buff, or even whitish .A. velosa, p. 286
49. Lower part of volva tightly constricted around stalk, the upper part flaring outward (see photo
on p. 289) .A. constricta, p. 289
49. Not as above; volva saclike .A. vaginata, p. 288
50. Cap pallid beneath a thin coating of pinkish-tan to brownish-orange universal veil material and
often with several large chunky warts at center; found in woods in Southeast . A. roseitincta
50. Not as above . 51
51. Cap distinctly pinkish when fresh; usually found in open; southern .A. salmonea
51. Not as above . 52
52. Cap with concentrically arranged brown to grayish-brown universal veil remnants (scales);
partial veil evanescent; known only from the Southeast .A. hesleri
52. Not as above; cap usually grayish or with grayish warts . 53
53. Cap pale brown to dark grayish-brown, with large grayish warts or patches; partial veil usually
forming a distinct annulus (ring) on stalk .A. spissa (see A. pantherina, p. 280)
53. Not as above; cap grayish .... A. onusta&A. cinereoconia (see A. sp. (unidentified), p. 275)

Amanitaphalloides (Death Cap) Color Plate 50

CAP 4-16 cm broad, at first nearly oval, then convex to plane; surface smooth, viscid or
tacky when moist, often shiny when dry or with a metallic luster; color variable: green to
brownish-olive, yellow-green, yellowish, or sometimes white, often darker toward center
and paler at margin, and often fading in age (to grayish-green, light brown, olive-buff,
dull yellowish, etc.); sometimes with one or more patches of thin, silky, white universal veil
tissue; margin typically not striate. Flesh white; odor at first mild, but later quite pungent
or nauseating (like raw potatoes or chlorine). GILLS adnate to adnexed or free, close, white
or tinged faintly greenish. STALK 5-18 cm long, 1-3 cm thick, tapering upward or equal
with an enlarged base; white or tinged cap color, smooth or with minute scales and fibrils;
solid or hollow. PARTIAL VEIL membranous, white or tinged yellow-green, forming a
persistent but fragile, superior, skirtlike ring which may disappear in age. UNIVERSAL
VEIL membranous, white, forming a saclike volva that sheathes base of stalk; volva thin
and rather fragile, usually buried inground and sometimes disintegrating. SPORE PRINT
white; spores 7-12 * 6-9 microns, broadly elliptical to nearly round, smooth, amyloid.

Amanita phalloides, showing the classical features of the deadly Amanitas: presence of a sack (volva)
at the base of the stalk and a skirt or ring (annulus) near the top of the stalk, plus white or whitish gills.
(Don’t expect every specimen to be so “classical,” however!)
Fruiting body development in the deadly Amanita phalloides. At first it is enclosed by the universal
veil, which ruptures to form a volva at the base of the stem as the stalk elongates.

HABITAT: Solitary, scattered, or in groups or troops in woods or on lawns near trees;

widely distributed. In our area it is very common under live oak in the fall and early winter
and may even turn up in the summer if moisture is sufficient. In the late winter and spring,
however, it is largely supplanted by A. ocreata. Perhaps an adventitious (but hardly
advantageous!) introduction from Europe, it has taken a fancy to our native oaks and
spread like the plague, so that it is now the most abundant Amanita of our live oak
woodlands. Heavy fall rains often elicit a stupendous crop—I have counted as many as one
hundred specimens under a single oak! I’ve never seen it growing without live oak in the
vicinity, but in southern Oregon it grows with other oaks and in eastern North America it
has turned up in numerous localities under conifers as well as hardwoods. Apparently it can
form mycorrhiza with a wide range of hosts!
EDIBILITY: DEADLY POISONOUS! Learn to recognize this species before eating any
mushroom with gills! It is particularly dangerous because the symptoms are delayed, not
appearing for from 6 to 24 hours after ingestion, by which time there is relatively little
modern medicine can do except to treat the victim symptomatically. In the last decade
there have been one or two deaths in California every time a bumper crop has appeared.
However, none of the victims were knowledgeable fungophiles, let alone “mushroom
experts” (as they are often called by the press). The flavor, incidentally, is described by
survivors as excellent—despite the awful odor which develops in old specimens. For an
account of symptoms and treatment, see pp. 892-893.
COMMENTS: There is no rational reason why anyone should mistake the death cap for
an edible mushroom—but since when were human beings completely rational? The telltale
signs are: (1) white gills (2) white spores (3) partial veil covering the gills, then breaking to
form a skirtlike ring or annulus near the top ofthestalk(4) membranous white sack (volva)
at the base of the stalk (5) margin of the cap not striate (however, I have seen mature
specimens with small striations on the margin where the cap tissue had apparently

270
Maturing specimens of Amanita phalloides. Note how cap opens out and the partial veil breaks to
form an annulus (ring) high up on the stalk. Cap color is variable: green, yellow, brownish, even white.
The cap is usually bald, but sometimes has a thin white patch of universal veil tissue (as shown here).

collapsed against the gills!). The ring is sometimes obliterated and the volva can be
carelessly overlooked, so just to be safe, don’t eat mushrooms with any two of these
characteristics unless you are absolutely sure what they are. Your life is at stake! The
cap color—usually greenish, but extremely variable—and the pungent odor in age (it
reeks of death) plus the association with live oak (in our area) are good secondary field-
marks. In eastern North America, A. phalloides can be confused with A. citrina and A.
brunnescens (see the key to Amanita and comments under A. porphyria).

Amanita ocreata (Destroying Angel; Death Angel)

CAP 4-15 cm broad, nearly round or oval becoming convex and finally plane; surface
viscid when moist but soon dry, smooth, white when young but in age often discoloring
pinkish, buff, yellowish, or brownish, especially toward the center; sometimes with a very
thin white patch of universal veil tissue; margin usually (but not always!) not striate. Flesh
thick, firm when young, white; odor mild, becoming disagreeable in old age. GILLS at first
adnate or adnexed, sometimes free in age, close, white. STALK 6-20 cm long, 1-3 cm
thick, tapered upward or equal with an enlarged base, white, often finely powdered or scaly
at apex or occasionally throughout; hollow or solid. PARTIAL VEIL membranous, white,
forming a very fragile superior or apical skirtlike ring or shredding into pieces or
disappearing entirely. UNIVERSAL VEIL membranous, white, forming a saclike volva
that sheathes the base of the stalk; volva often large and ample, but sometimes thin. SPORE
PRINT white; spores 9-14 * 7-10 microns, broadly elliptical to nearly round, smooth,
amyloid. Flesh turning bright yellow in KOH (potassium hydroxide).
HABITAT: Solitary to widely scattered or in small groups on ground under oaks, known
from Marin County in California east to the Sierra Nevada foothills and south to Arizona
and Texas. In our area it is associated with live oak and is common in the winter and spring,

271
Amanita ocreata. This deadly poisonous Amanita is pure white when young but usually discolors
(pinkish, brownish, ochre, etc.) on the cap as it ages. Note the fragile partial veil, voluminous volva,
and white gills. See Color Plate 53 for its eastern counterpart.

usually after A. phalloides has finished fruiting. It is quite numerous, but doesn’t normally
produce the huge crops typical of the latter species. Related “destroying angels” are abun¬
dant under hardwoods and conifers in eastern North America, and to a lesser extent, the
Pacific Northwest (see comments).

EDIBILITY: DEADLY POISONOUS! It doesn’t enjoy the same notoriety as A. phal¬

loides, but is just as dangerous, and in some regions (such as southern California), it is the
more common of the two. It recently caused several deaths near San Diego. The victims
were apparently starving illegal aliens who ate the mushrooms out of desperation.

COMMENTS: This elegant, pristine-pure, lethal-looking Amanita is our only white

mushroom with both an annulus (ring), a true sack (volva) at the base of the stalk, and a
non-striate (or only rarely striate) cap margin. The species epithet ocreata means
“sheathed,” a reference to the voluminous volva which may extend as much as halfway up

Amanita ocreata varies greatly in stature, as shown here. Left: A robust specimen that could easily
be mistaken for the pale form of A. calyptrata (and is just as big). Right: A slender, graceful mature
specimen that could be mistaken for a washed-out A. velosa.
AMANITA 273

the stalk, though it is often much smaller. The partial veil is quite fragile, however, and
often turns to shreds rather than forming a distinct ring (annulus). The sinister name
“destroying angel” also embraces three closely related, deadly poisonous, pristine-white
“veiled threats”—A. virosa (COLOR PLATE 53), A. verna, and A. bisporigera. All three
are common in eastern North America — especially under hardwoods — and have been
reported from the Pacific Northwest, but not from California. They closely resemble A.
ocreata, but are often more slender and do not discolor as much in age; also, A. verna
supposedly doesn’t yellow in KOH. A. virosa is distinguished by its round spores and scalier
stalk, while A. verna has elliptical spores and A. bisporigera has 2-spored basidia and
round spores. A. alba and the pale (white to creamy) form of A calyptrata are easily con¬
fused with these species, but always have a striate cap margin and non-amyloid spores. Be
very careful—A. ocreata sometimes has a striate cap margin (see photo on p. 286)! Other
species: A. mutabilis is an eastern species with a white or pinkish-tinged cap. It has a sac-
like volva and amyloid spores, stains pinkish when bruised, and often smells like anise.

A manita silvicola (Western Woodland Amanita)

CAP 5-12 cm broad, convex to plane; surface dry or slightly viscid when moist, white (but
occasionally discolored in age), covered with flattened cottony or fluffy-powdery universal
veil tissue; margin often hung with veil remnants and extending beyond the gills, not
striate. Flesh white; odor mild or slightly soapy. GILLS white, close, adnate to adnexed or
free, edges finely powdered or cottony. STALK 5-12 cm long, 1.5-2.5 cm thick, usually
rather stout, terminating in a basal bulb which is up to 5 cm broad; white (or sometimes
brownish-stained), usually powdery or with cottony scales. PARTIAL VEIL white,
delicate, forming a slight ring or fibrillose zone or disappearing entirely. UNIVERSAL
VEIL cottony, white, forming a scaly or indistinct volva consisting of cottony white zones
or patches at base of stalk which often disintegrate or remain in the ground. SPORE
PRINT white; spores 8-12 * 4.5-6 microns, elliptical, smooth, amyloid.
HABITAT: Solitary or in small groups in mixed woods and under conifers(e.g., Douglas-
fir); known only from the Pacific Northwest (where it is fairly common in the fall, parti¬
cularly in campgrounds) and California. I have found it only once in our area, in December.
EDIBILITY: Unknown. Do not experiment!
COMMENTS: This attractive Amanita can be told at a glance by its cottony white cap,
rather stout stature (for an Amanita), and enlarged stem base(see photo on next page). The
cap lacks the prominent warts characteristic of A. magniverrucata, A. cokeri, etc., and the
volva is not saclike as in the destroying angels (A. ocreata, A. virosa, etal). Also, the stalk
lacks the tapered, pointed, rooting base so characteristic of A baccata. The attached gills
and indistinct volva suggest Armillaria, but it has that ineffable Amanita “aura.” Also
see A. smithiana (under A. magniverrucata), a larger version of A. silvicola.

Amanita baccata (Sand Amanita) Color Plate 54

CAP 4-10(12) cm broad, convex becoming plane; surface dry or slightly tacky, white (but
often dirty and sometimes discolored buff in age), with obscure mealy to powdery warts or
flattened universal veil remnants which may disappear in age; margin often hung with
veil remnants, not striate. Flesh white, soft, fragile; odor mild or slightly pungent. GILLS
close or crowded, white, becoming dingy yellowish in age, usually adnate or adnexed, but
sometimes free. STALK 5-18 cm long, 0.5-2.5 cm thick, with a long, tapered, often
pointed rooting base below a slight to distinct bulb; dry, white, sometimes with yellowish or
buff stains; usually somewhat fibrillose or with delicate ragged or powdery scales;
apex often striate. PARTIAL VEIL white, cottony, very fragile; disappearing or forming
a thin, poorly defined superior ring on stalk. UNIVERSAL VEIL friable, white, forming
Left: Amanita silvicola lacks the saclike volva of A. ocreata and the tapered rooting base of A. baccata.
Right: Amanita baccata grows in sand; note pointed rooting base (see color plate for view of cap).

an indistinct volva or a scaly zone just above the bulb. SPORE PRINT white; spores
10-15 x 4-6.5 microns, more or less cylindrical, smooth, amyloid.

HABITAT: Solitary or in small groups in sand or sandy soil (often buried), associated with
oak and/or pine; distribution spotty—reported from southern Europe, northern Africa,
and Michigan—frequent in our area in the fall and winter, but never in large numbers.
EDIBILITY: Unknown.
COMMENTS: Also known as A. boudieri, this is an inelegant but oddly charming
Amanita. The tapered, rooting stem base, soft flesh, absence of prominent warts on the
cap, cylindrical spores, and habitat in sand form a most distinctive combination of
features. The cap may scarcely poke above the ground and along with the rooting base, is
usually covered with dirt or sand (see photo). The fruiting body decays rapidly and is hard
to keep in one piece. For these reasons and many more, it is one of my favorite Amanitas.

A manita magniverrucata (Pine Cone Amanita) Color Plate 55

CAP 7-20 (30) cm broad, nearly round becoming broadly convex or plane; surface dry or
slightly viscid when moist, covered with large (up to 2 cm broad and 1 cm high!), persistent,
strongly-attached warts, the warts pyramidal at first becoming truncated or more flattened
in age; color white to creamy-white becoming yellowish-buff or tan in old age (sometimes
with darker stains); margin usually with cottony veil remnants and extending beyond the
gills, not striate. Flesh thick, firm, white; odor unpleasant in age (like chlorine or dirty
socks). GILLS adnate to adnexed or free, close, white or creamy, delicately powdered.
STALK 7-12 (20) cm long, 1-4 cm thick at apex, rooting deeply in ground; equalabove or
tapering upward from a thicker base or bulb (up to 6 cm broad but sometimes incon¬
spicuous), the rooting portion below the bulb tapered downward; white throughout or
with brownish to yellowish-buff stains; firm, rather tough. PARTIAL VEIL white, mem¬
branous, usually forming a fragile, superior, skirtlike ring on stalk. UNIVERSAL VEIL
warty and friable, forming a scaly volva consisting of concentrically arranged rows of
warts or scales at apex of bulb, these sometimes disappearing in age. SPORE PRINT white;
spores 8.5-12.5 * 5.5-8.5 microns, elliptical to nearly round, smooth, amyloid.
HABITAT: Solitary to gregarious under live oak and pine; known only from California,
but related species (see comments) are widespread. In our area it is not uncommon in the
fall, winter, and spring. It develops slowly and persists for weeks without decaying.

274
Left: Amanita magniverrucata, top view of cap. Note the exaggerated warts. Right: Amanita cokeri
is a somewhat similar whitish species with smaller warts on the cap.

EDIBILITY: Unknown—do not experiment! It belongs to the “Lepidellas,” a subgroup

of Amanita that contains both poisonous and harmless species.
COMMENTS: One of our most spectacular mushrooms—the large, erect warts on the cap
set it apart, making it look like a white pine cone or a glob of meringue (buttons resemble
the Sierran puffball, Calvatia sculpta). It is one of several large, white, warty Amanitas
that for years have passed under the names A. strobiliformis and A. solitaria. However, our
species is quite distinct because of its exaggerated warts, and has recently been rewarded
with a name of its own. It has several large counterparts in eastern North America with an
unpleasant, often chlorine-like odor and more fragile warts, including: A. chlorinosma,
A. polypyramis (especially large), and A. rhopalopus (with a large, deeply rooting base);
A. ravenelii, with brownish imbricate (shingled) warts or scales; and A. daucipes, which
often has a pinkish- or orange-tinged stalk. A. cokeri is a common eastern oak- and pine-
loving species that also occurs in California. It has smaller (to 5 mm broad) but firmly-
attached warts mainly at the center of the cap, a mild odor, a rooting bulb on the stalk, and
an annulus (ring) and volva (see photo above). A. smithiana grows under conifers in the
Pacific Northwest. It looks something like a matsutake or large A. silvicola with its poorly
developed cottony warts, unpleasant odor, and ragged or shaggy stalk with an enlarged but
scarcely pointed base. There are many other closely related white or pallid species (the
“Lepidellas”), particularly in southeastern North America(e.g., A. abrupta, with a broad,
abrupt basal bulb; A. longipes, usually odorless with a pointed, often flattened, rooting
bulb; A. thiersii, with a shaggy stalk; A. atkinsoniana and A. cinereopannosa, with a
grayish veil; and A. praegraveolens, with a whitish to pinkish-tan, scaly cap and non-
bulbous stalk). “Lepidellas” grow on lawns as well as in the woods, and are often difficult
to identify without a microscope. They should not be eaten.

Amanita sp. (unidentified) (Anonymous Amanita)

CAP 2.5-5 (8) cm broad, convex becoming plane; surface dry, whitish to gray, covered with
darker (gray to brownish-gray) mealy or powdery warts which are easily obliterated;
margin not striate. Flesh white, odor rather pungent in age. GILLS usually adnate or
adnexed (rarely free), close, creamy-white becoming dingy yellowish or yellowish-orange
in old age. STALK 2-4 (7.5) cm long, 0.4-1 cm thick, equal or tapering downward, some¬
times with a short, rooting base or swollen slightly above the base; white and striate above
the ring, dingy whitish or tinged cap color below and somewhat scaly. PARTIAL VEIL
membranous but very fragile, disappearing or forming a superior, median, or even basal
ring on stalk. UNIVERSAL VEIL friable, mealy-powdery, forming an indistinct or scaly
volva in the form of obscure grayish scales or powdery-mealy warts over lower portion of
stalk. SPORE PRINT white; spores 7-11 x 5-8.5 microns, elliptical, smooth, amyloid.
 :: >i :

::

Amanita sp. (unidentified). An odd, unimposing species that is locally abundant in pastures and open
ground. Note the small size, stocky stature, and grayish warts on the cap. The fragile partial veil and
volva are easily obliterated.

HABITAT: Scattered to gregarious in pastures, open fields, hard-packed ground, under

trees, etc. Fairly common in our area in the fall and winter and sometimes very abundant.
It usually grows in the open, often mingling with Agaricus campestris and Agaricus
cupreobrunneus, which it superficially resembles. Apparently it is not mycorrhizal.
EDIBILITY: Unknown. Do not experiment!
COMMENTS: This anomalous, anonymous Amanita is quite unamanitalike with its
short stem, compact mealy-warty, grayish cap, and predilection for growing in meadows.
The volva is so fragile that it may disappear, but the warts on the cap signify Amanita.
A. farinosa is a similar species (cap small, powdery-mealy, gray to brownish-gray, etc.),
but has non-amyloid spores, a striate cap, no partial veil, and grows in forests or at their
edges. It is widespread but rare in the West. Two similar grayish species, A. cinereoconia
and A. onusta, are larger and apparently restricted to eastern North America; the latter
usually grows in sandy soil.

Amanita rubescens (Blushing Amanita; The Blusher)

CAP 4-12 (20) cm broad, convex becoming plane or shallowly depressed; surface slightly
viscid or dry, at first covered with white, pinkish, brownish, or grayish warts, the back¬
ground white at first, then flushed sordid reddish, pinkish, reddish-brown, brown, etc.;
margin usually not striate. Flesh firm, white, slowly reddening when bruised; odor mild,
taste mild or latently bitter. GILLS adnate to adnexed when young, sometimes free in age;
close, white or pallid, sometimes stained reddish. STALK5-14(20)cmlong, 1-3.5 cmthick,
equal or enlarged downward to a swollen base (bulb); at first white, soon stained sordid
reddish, reddish-brown, or pinkish below the ring and often somewhat scaly; white or
tinged pinkish above. PARTIAL VEIL membranous, white or tinged reddish, forming a
fragile, superior, skirtlike ring on stalk. UNIVERSAL VEIL friable, forming an indistinct
or scaly volva (i.e., disappearing or leaving sordid reddish scaly zones at base of stalk).
SPORE PRINT white; spores 7.5-10.5 * 5-7 microns, elliptical, smooth, amyloid.
HABITAT: Solitary to scattered or in groups in woods and under trees, partial to oak but
also found with conifers; common and widely distributed. In our area it is monogamous
with live oak and usually fruits twice—a small flush after the first fall rains, and a larger
crop in the late winter and spring (February-April) when A. velosa and A. ocreata
“bloom.” It can also turn up in the summer—in fact, I’ve found it every month of the year!
EDIBILITY: Not recommended. It is edible when cooked, but indigestible or even
poisonous raw, and it is easily confused with poisonous species. It is highly esteemed in
Europe (chiefly France), but our local version does not have a good flavor.

276
Amanita rubescens is extremely variable in size, stature, and color, but always stains dingy reddish
(often slowly). These whitish-capped specimens are small; more can be seen below and on p. 266.

COMMENTS: The “blushing” of the cap, stem, and flesh is the one infallible fieldmark
of this fickle fungus. The blushing process is slow to manifest itself and is best seen on the
lower stalk or around the edges of maggot tunnels (see photograph below). In other
respects it is an exasperatingly variable Amanita. Young specimens may be pure white
(in our form), while older individuals usually develop strong reddish or brownish tones.
The warts may be evenly disposed over the entire cap surface, or concentrated at the center,
or more prevalent toward the margin, or completely absent (especially in rainy weather).
Mature specimens are sometimes mistaken for A. pantherina (if they are brownish), or even
A. muscaria (if they are reddish), but the indistinct volva, reddish stains, and amyloid
spores separate it. Pure white buttons, on the other hand, resemble the A. strobiliformis
group (see A. magniverrucata), but lack the tapered, rooting base of those species. They can
also be confused with the “false blusher” (see below). The attached gills and absence of a
volva can lead to confusion with Armillaria, but once again the reddish stains distinguish it.
The “true” A. rubescens that is so common in eastern North America differs from our
form in several respects. Its universal veil is grayish to dirty pinkish and its cap is soon
reddish-brown to flesh-color to brown or olive-tinged, while ours is often white. Also, it is
a much larger, taller, and more stately fungus, and is often parasitized by a pallid to pinkish
mold (see photo onp. 884), whereas ourform is not. Infact,aboutall that ourversionhasin
common with the “true” A. rubescens is its blushing behavior—and it may eventually
prove to be a distinct variety or species. (Perhaps we should appoint an ad hoc committee
to investigate the problem!)
Also common under live oak in our area is the “false blusher”—an unidentified and
probably unnamed species that closely mimics our A. rubescens in every respect save one:
it does not “blush.” Its bulb is often brownish-stained, however, and like A. rubescens it
is whitish when young, browner in age, and has warts on the cap. Its edibility is unknown
—another good reason not to eat A. rubescens!

Amanita rubescens. Left: Mature brown-capped specimens; note warts on cap and indistinct volva.
Right: Even the maggot tunnels stain reddish, as shown in this close-up.
278 AMANITACEAE

A manita aspera (Yellow-Veiled Amanita) Color Plate 56

CAP 4-12(15) cm broad, nearly round to convex, then plane; surface viscid when moist,
dark brown to grayish-brown, yellow-brown, or bright yellow, covered with yellow mealy
or powdery warts which become flattened and grayish to dingy buff in age or occasionally
disappear; margin not striate or only faintly so. Flesh white or tinged yellow, soft. GILLS
white or creamy-yellow, close, adnate to adnexed or free. STALK 5-15 (20) cm long,
0.7-2.5 cm thick, equal or tapering upward, the base often enlarged; white to pale yellow
above the ring, white to yellow, buff, or grayish-tinged below; base often with orangish,
reddish, or brown stains(on exterior or interior). PARTIAL VEIL membranous, white or
pale yellow above, bright yellow to grayish-yellow on underside; forming a superior, skirt¬
like ring on stalk. UNIVERSAL VEIL powdery, friable, forming a scaly volva (scaly
zones and/or yellow to grayish-yellow powdery scales) at base of stalk, but volva easily
obliterated. SPORE PRINT white; spores 8-12 * 6-8 microns, elliptical, smooth, amyloid.
HABITAT: Solitary to scattered or in small groups under both hardwoods and conifers;
common in our area and throughout the Pacific Northwest, but seldom in large numbers.
It often grows with A. muscaria in our coastal pine forests and the two make a colorful pair;
it is also frequent in our oak-madrone woodlands. Rather than fruiting in one large,
spectacular burst like A. calyptrata and A. muscaria, it usually keeps a lower profile, fruit¬
ing rather sporadically throughout the mushroom season.
EDIBILITY: To be avoided. Chemical analysis has failed to reveal the presence of
amanita-toxins, but this does not mean it is edible. True, the European form is edible
(according to one source), and if our form were poisonous we would probably know by now
—but why tempt fate when there are so many other safe, savory mushrooms available?
COMMENTS: This elegant Amanita is easily recognized by its powdery yellow universal
veil and yellow partial veil. Fresh specimens are among our most lovely mushrooms. The
cap color ranges from yellow through yellow-brown to dark brown and is apt to bewilder
the color-conscious beginner. A. pantherina and A. gemmata are often confused with A.
aspera, but have non-amyloid spores and white veils plus a collarlike volva. A. flavo-
rubescens (-A. flavorubens) is a very similar if not identical species with a yellow-orange
to yellow or yellow-brown cap. It is common in eastern North America, especially under
oak. A.flavoconia is also similar, but is smaller and never brown and is rare in the West.

A manita flavoconia
CAP 2.5-7.5 (10) cm broad, nearly oval becoming convex or plane; surface viscid when
moist, bright orange to yellow-orange or yellow, or orange at the center and yellow at the
margin; adorned with scattered yellow warts which may disappear in age or rainy weather;
margin not striate or only faintly so. Flesh rather thin, white. GILLS adnate to adnexed or
free, close, white, sometimes with yellow edges. STALK 4-10 cm long, 0.5-1.5 cm thick,
tapering upward or equal with an enlarged base; smooth or somewhat scaly, yellow or
sometimes white. PARTIAL VEIL membranous, forming a superior, skirtlike ring which
usually has a yellow underside. UNIVERSAL VEIL friable, forming a scaly volva consis¬
ting of powdery yellow scales and patches at base of stalk which wear off easily or disappear
in age. SPORE PRINT white; spores 7-11 * 3.5-5 microns, elliptical, smooth, amyloid.
HABITAT: Solitary to scattered or gregarious in woods (mainly under hardwoods, but
also with conifers); often abundant in summer and early fall in eastern North America, but
rare in the West. (It occurs in Arizona and has been found in northern California.)
EDIBILITY: Unknown. Do not experiment!
COMMENTS: This beautiful Amanita looks like a miniature A. muscaria, and is often
mistaken for the yellow-orange form of that species. However, it does not have concentric
AMANITA 279

rings at the base of the stalk and is much smaller. The yellow veil remnants are reminiscent
of A. aspera, but the cap is orange to yellow rather than yellow to brown, and the stalk is
often yellow as well. Other species: A.frostiana of eastern North America is similar, but
has whitish to buff warts, a more distinct volva, and non-amyloid spores.

Amanita porphyria (Booted Amanita)

CAP 3-12 cm broad, convex to plane or broadly umbonate; surface slightly viscid when
moist, gray to grayish-brown, often with a subtle purplish cast; usually adorned with
scattered grayish (or sometimes whitish) warts, these sometimes merging to form a patch
or often wearing off or washing away; margin not striate or only faintly so. Flesh white;
odor often turniplike in age. GILLS adnate to adnexed or free, close, white or sometimes
aging or bruising grayish. ST ALK 5-18 cm long, 1 -1.5 cm thick at apex, equal above with
a large, soft, abrupt, rounded or flattened bulb at the base; bulb often cleft; white or pale
gray above the ring, gray or with grayish to grayish-brown or purple-gray patches below.
PARTIAL VEIL membranous, forming a superior, skirtlike ring which often collapses
or disappears in age; ring gray or grayish-yellow. UNIVERSAL VEIL somewhat mem¬
branous but also friable; forming a collarlike volva (free rim) around the apex of bulb or
leaving scattered grayish patches or disappearing. SPORE PRINT white; spores 7-10
microns, round, smooth, amyloid.
HABITAT: Solitary, scattered, or in small groups under conifers; widely distributed in
northern North America. It is fairly common in the Pacific Northwest but rather rare in
California. I have yet to find it in our area.
EDIBILITY: Poisonous?? Do not experiment!
COMMENTS: The abrupt, soft, rounded, basal bulb and grayish color distinguish this
conifer-loving Amanita. The volva may have a free margin but is not truly saclike. In
eastern North America there are two related species with a prominent basal bulb:
A. citrina (-A. mappa) is very common (mainly under hardwoods) and rather slim, with
a pale yellow to yellow-green to whitish cap and buffy-white to pinkish or lavender-gray
warts; A. brunnescens, also fond of hardwoods, has a brown to olive-brown or paler cap,
a stalk that stains reddish-brown, and a longitudinally split or chiselled basal bulb; it has
also been reported from the Pacific Northwest. Neither of these Amanitas should be eaten.

Amanita porphyria. Note large basal bulb and grayish patches on stalk. Cap is purple-gray to brown.
Amanita pantherina, young button at right, mature specimen at left, intermediate stages between.
Note the collarlike volva (i.e., the free rim of the basal bulb).

A manita pantherina (Panther Amanita)

CAP 5-15 (25) cm broad, nearly round or convex becoming plane to slightly depressed;
surface viscid when moist, color variable: dark brown to light brown, tan, dull yellowish,
or paler (see comments), often darker at center and paler toward margin; adorned with
many white to pale buff universal veil remnants (warts), but these often washed off by rain;
margin usually striate. Flesh firm, white. GILLS adnate to adnexed or free, close, white or
pallid. STALK 5-15 (20) cm long, 1-3 cm thick, tapering upward or equal with a basal
bulb; dry, white or aging buff; usually smooth above the ring, often scaly below. PARTIAL
VEIL membranous, white, forming a superior or median skirtlike ring whose margin is
often ragged or toothed. UNIVERSAL VEIL friable, white, usually forming a collarlike
volva (i.e., adhering to the bulb except for a free rim at apex of bulb), sometimes also with
scaly or ragged zones above the free rim, or sometimes indistinct (no free rim). SPORE
PRINT white; spores 9-13 * 6.5-9 microns, elliptical, smooth, not amyloid.
HABITAT: Solitary to scattered or gregarious on ground in woods; widely distributed,
but especially common under conifers from the Rocky Mountains westward. In our area it
fruits from fall through spring and is quite common under pine, oak, and Douglas-fir,
though rarely in the large numbers characteristic of A. muscaria. In the Pacific Northwest
and Sierra Nevada it is perhaps the most omnipresent of all the Amanitas.
EDIBILITY: Poisonous! It contains the same toxins as A. muscaria, but apparently in
higher concentrations—large doses can be fatal! It is said to be one of the most common
causes of mushroom poisoning in the Pacific Northwest.
COMMENTS: The dark brown to tan or dull yellowish cap with whitish warts and the
free rim or collar at the top of the basal bulb are the fieldmarks of this ubiquitous species. As
is often the case in Amanita, there is considerable variation in cap color as well as in size.
Yellowish forms are difficult to distinguish from A. gemmata—generally they are duller
and often somewhat browner at the center, but the two species appear to intergrade (see
comments under A gemmata). The cap is never red or bright orange-yellow as in A.
muscaria, and the stalk lacks the grayish patches characteristic of A. porphyria. A. spissa
of eastern North America is somewhat similar, but has amyloid spores and a gray universal
veil that leaves a powdery-scaly rather than collarlike volva. A. cothurnata is a slender
eastern species with a whitish cap that is often tinged yellowish or brown at the center. A
mushroom meeting this description occurs in our area, but is rare. Some amanitologists
consider A. cothurnata to be a variety of A. pantherina.
AMANITA 281

Amanita gemmata (Gemmed Amanita; Jonquil Amanita)

CAP 4-10(14) cm broad, rounded becoming convex to plane; surface slightly viscid when
moist, creamy to pale yellow, golden-yellow, or buff (but see comments), often slightly
darker at the center; covered with whitish universal veil remnants (warts) which may merge
to form a patch or may disappear in age; margin striate or not. Flesh white, fairly thick.
GILLS adnate to adnexed or free, close, white. STALK 5-13 cm long, 0.5-2 cm thick,
tapered upward or equal with an enlarged base; dry, white or tinged yellowish, usually
smooth above the ring and sometimes scaly below. PARTIAL VEIL typically present
(but absent in var. exannulata), membranous, white; forming a fragile, superior to
median, skirtlike ring on stalk, or disappearing. UNIVERSAL VEIL friable to somewhat
cottony, white, usually forming a collarlike volva (free rim) at apex of basal bulb,
sometimes also with scaly zones above rim, or sometimes forming a thin sheath instead
of a rim, or sometimes the volva more or less indistinct. SPORE PRINT white; spores
8-13 * 6-9 microns, elliptical, smooth, not amyloid.
•»

HABITAT: Solitary, scattered, or in groups in woods or along forested paths and roads;
widely distributed. I n our area it occurs in mixed woods and under live oak shortly after the
first fall rains and less commonly thereafter. In the Sierra Nevada and Pacific Northwest
it is common under conifers in the spring as well as the summer and fall. The mountain form
differs slightly from the typical form and may eventually be classified as a separate species.
EDIBILITY: Poisonous! Some books list it as edible, but other sources say it contains
the same toxins as A. pantherina and A. muscaria.
COMMENTS: Also known as^4. junquillea, this attractive Amanita can be distinguished
from typical A. pantherina by its yellow to creamy cap and more modest size. (A. pan¬
therina ranges from small to very large.) However, a confusing series of “hybrids” exist
whose cap color and degree of toxicity are intermediate between the two species. A. aspera
and A. flavoconia differ by having yellow veil remnants, while the yellow-capped form of
A. muscaria is larger, with a volva composed of concentric rings. Sometimes the universal
veil of A. gemmata forms a continuous layer over the entire cap, but does not peel off easily
like the volval patch of A. calyptrata. The latter also differs in its large, thick, saclike volva.
Other species: A. russuloides of eastern North America is a similar, slimmer version of
A. gemmata and may just be a regional variation. A. breckoniihas a double, nearly basal
annulus (ring) and a short, tapered, rooting base beneath the bulb. It occurs in our area
under pine and perhaps oak, but is rare. A small species with an orange cap (sometimes
yellow toward margin) and no partial veil also occurs rarely in our area. It may be a form
of var. exannulata or it may be an undescribed species.

Amanita gemmata. Left: Typical specimens; note collarlike volva. Right: A vernal mountain form
that may actually be a distinct species. It also has a yellow cap, but is often stockier. It usually
appears when the morels are out.
282 AMANITACEAE

Amanita muscaria (Fly Agaric; Fly Amanita) Color Plates 58,59

CAP 5-30 (40) cm broad, round becoming convex and finally plane or slightly depressed;
surface viscid when moist, color variable: bright red to blood-red, scarlet-red, or orange-
red when fresh (var. muscaria and var. flavivolvata), but often fading to orange, yellow-
orange, or paler; bright yellow-orange to yellow, then fading (var. formosa), yellow with a
peachy center (var. persicina); or white to buff to silvery-grayish-white(v«r. alba)', covered
at first with a dense coating of universal veil fragments (warts) which are usually white(but
are yellow in var. flavivolvata and often buff or tan in var. alba)', warts flattened in age, often
wearing away or washed off by rain; margin usually at least somewhat striate. Flesh firm
when young, soft in age; thick, white. GILLS adnatetoadnexed or free, close, broad, white.
ST ALK 5-20 (30) cm long, 1 -3 (4) cm thick at apex, tapering upward or equal with a basal
bulb up to 6 cm broad; white or whitish, or somewhat discolored in age; smooth or
somewhat ragged-scaly below the ring; often fragile in old age. PARTIAL VEIL
membranous, usually forming a thin, persistent, median to superior, skirtlike ring on stalk
which may collapse in age; ring white or with yellow patches, margin often torn or toothed.
UNIVERSAL VEIL friable, forming a scaly volva at apex of bulb consisting of one or more
(usually 2-4) concentric rings. SPORE PRINT white; spores9-13 x 6.5-9 microns, broadly
elliptical, smooth, not amyloid.
HABITAT: Solitary or scattered to densely gregarious or in large rings in forests and at
their edges, also with planted trees. Common throughout most of the northern
hemisphere—its favorite mycorrhizal mates include pine, spruce, fir, birch, and aspen. The
bright red form with white warts (var. muscaria) is the common one in western North
America as well as Europe and Asia. In our area it fruits along with the yellow-warted
variety from fall to early spring and is often abundant in our coastal pine forests and along
freeways where pines have been planted. It also turns up occasionally under oak and
madrone. In the Rocky Mountains it is common in late summer with spruce and in the
Pacific Northwest in the fall with various conifers. The red-capped form with yellow warts
(var .flavivolvata) has a southern distribution but is quite common in our area. The form
with a bright yellow to yellow-orange cap (var .formosa) is the dominant one in eastern
North America. In California it is common in the Sierra Nevada but rather rare in the Coast

Amanita muscaria. Left: A button completely covered with white warts (the universal veil). Right:
“Bloody mirror of the galaxy.” This large button was frisbee-sized when fully expanded. (J oel Leivick)
 ■A

•L>
■

Amanita muscaria var. alba closely resembles the more common red-and yellow-capped varieties,
but has a white to grayish-buff cap.

ranges. The grayish-white form (var. alba) is more common in northern regions. It has been
found in northern California, but is rare. The peach-centered variety (var. persicina) is
most common in the S outheast. It is interesting to note that wherever A. muscaria grows it
is often accompanied by Boletus edulis, and can serve as a “red flag” indicator for that
species. I’ve even seen a very large ring of A. muscaria locally that numbered over 100
fruiting bodies, with three bulky B. edulis growing right in the middle of it!

EDIBILITY: Poisonous and hallucinogenic—esteemed by both maggots and mystics.

Fatalities are extremely rare, but it is undoubtedly dangerous in large or even moderate
amounts. The effects vary from person to person, mushroom to mushroom, and from
region to region and season to season, so that there is no way to determine in advance
what one’s reaction will be. Too many people have had unpleasant experiences for me to
recommend it (for an account of its effects, see p. 894). Tales of “getting off’ by nibbling
on a piece of the veil or licking the “stars” off the cap are frivolous. However, the skin ap¬
parently contains greater concentrations of the toxins than the rest of the fruiting body.
The name “fly agaric,” incidentally, is derived from the ancient practice of using the mush¬
room (often mixed with milk) to stupefy flies. It is also notable for its ability to concentrate
vanadium—a rare, malleable, ductile metal used to add tensile strength to steel—from the
soil.
COMMENTS: The brilliant red form of this mushroom needs no introduction—it is
known to every myopic middle-class “mystic” in America. Its caricature appears on key
chains, incense holders, posters, candles, curtains, calling cards, and calendars. Large
plastic reproductions can be found on lawns and in display windows, amusement parks,
and fantasy decor. The irony of it is that few people realize that such a mushroom actually
exists! The color, of course, is its outstanding fieldmark, plus the numerous warts on the
cap, which look like curds of cottage cheese, or to the more cosmically inclined, like “stars.”
(The warts may be washed off by rain, however.) The white gills, presence of an annulus
(ring), and scaly volva separate it from other bright red mushrooms( notably the Russulas),
and, unless you’re color blind, it is difficult to confuse it with any of our other Amanitas.
The cap may fade drastically as it ages, especially in direct sunlight or after a soaking rain,
but will usually retain at least some vestige of its original splendor. The other color forms,
particularly the white one, are not quite so distinctive, and are best recognized by their
volva, which usually consists of a series of scaly concentric rings above the basal bulb. In
our area the fly agaric grows bigger than any other Amanita—\ have seen “monsters”
nearly two feet broad! In such specimens the stalk has difficulty supporting the weight of
the cap and will break at the slightest provocation, or even topple over of its own accord.

283
284 AMANITACEAE

A manita caesarea group (Caesar’s Amanita) Color Plate 60

CAP 7-20 (25) cm broad, nearly oval to convex, becoming broadly convex or plane; surface
slightly viscid when moist, smooth, or sometimes with one or two pieces of thick white
universal veil tissue; bright red to orange-red or orange, often fading to yellowish or even
paler in sunlight or heavy rain; margin conspicuously striate. Flesh thick, yellow under
the cuticle, otherwise white. GILLS close, broad, adnate to adnexed or free, bright chrome-
yellow or at times egg-yellow, or pale yellow with darker yellow edges. STALK 5-15 (20)
cm long, 1.5-3 cm thick, equal or slightly thicker below (but may appear bulbous from the
thick volva); smooth or with small scales, same color as gills or slightly paler; stuffed with a
pith or jelly, eventually hollow. PARTIAL VEIL membranous, forming a persistent,
superior and often striate, skirtlike ring which is colored like the gills. UNIVERSAL VEIL
thick, membranous, white, forming a large, rather tough, lobed, saclike volva which is
attached only to the base of stalk and may have a flange within. SPORE PRINT white or
faintly yellow; spores 8-12 x 6-8 microns, elliptical, smooth, not amyloid.
HABITAT: Scattered to gregarious or sometimes in large rings, in pine and oak woods in
southern Europe and much of the warm temperate zone. Its North American distribution
parallels that of Lactarius indigo: it is common in the summer under ponderosa pine in
Arizona and New Mexico, as well as with various pines in Mexico and Central America,
and a slightly different variety (see comments) occurs in the southeastern United States
north to Quebec. It has not been recorded from California but is to be looked for in the
warmer parts of the state (it is apparently supplanted by A. calyptrata). I have seen
enormous fruitings near the Grand Canyon, in August and September.
EDIBILITY: Edible and highly prized in Europe, where it is considered among the very
best of all mushrooms (it is said to have been a favorite of the Caesars). In America,
however, it is not so highly regarded. Its flavor is very similar to that of A calyptrata, which
is to say, not to everyone’s liking. Because of its brilliant color it is by far the safest of the
Amanitas for the table (but not necessarily the best—see A. velosaf).
COMMENTS: This resplendent prince of the Amanitas is easily distinguished by its bright
yellow gills and stalk, brilliant red to orange cap with striate margin, and thick, volumi¬
nous, white saclike volva. It is a robust fungus that calls to mind A. calyptrata. In fact,
it may mimic the latter in having one or more thick patches of veil tissue on the cap,
especially in dry climates. The variety in eastern North America has a slimmer stalk
(sometimes with reddish fibrils) plus a usually bald, sometimes umbonate cap that is often
yellow toward the margin in age. It is now considered a distinct species, A. hemibapha
(formerly A. umbonata). Another common eastern species, A. parcivolvata, is similar in
color(red to yellow-orange striate cap and yellow gills), but lacks a partial veil and does not
have a saclike volva; it is harmless anyway. The poisonous A. muscaria is easily dis¬
tinguished by its warted cap, white gills and stalk, and scaly rather than saclike volva.

Amanita calyptrata (Coccora; Coccoli) Color Plates 61,62,63

CAP 7-25 cm or more broad, rounded becoming convex and finally plane; surface viscid
when wet, usually with a large, thick, central, cottony whitish patch of universal veil tissue;
otherwise typically dull orange to orange-brown, bronze, yellow-brown, or dark brown,
usually with a paler (yellow) margin, or pale yellow to creamy or whitish (the pale form)
or occasionally greenish; margin usually clearly striate. Flesh firm, thick, white orcreamy;
odor mild becoming slightly unpleasant in age. GILLS broad, close, adnate becoming
adnexed or free, white to pale creamy-yellow. ST ALK 7-25 cm long, 1-3.5 cm thick, equal
or tapering upward, smooth, creamy to yellowish (typical form) or creamy to white (pale
form) or white to greenish (greenish form); hollow, the cavity often filled with a cottony
or jellylike substance. PARTIAL VEIL membranous, colored like stalk or slightly darker,
Amanita calyptrata (-A. calyptroderma), mature specimens. Note striate cap margin. (Joel Leivick)

forming a large but fragile, superior, skirtlike ring on stalk which is easily obliterated.
UNIVERSAL VEIL membranous, white, forming a large saclike volva that sheathes the
stalk base; volva thick, ample, felty or cottony, often lobed, often with a collar or flange
within. SPORE PRINT white; spores 8-11* 5-6 microns, elliptical, smooth, not amyloid.
HABITAT: Solitary to widely scattered or gregarious on ground in woods; known only
from the west coast. The distribution of the typical (darker) form parallels that of its
favorite mycorrhizal mate, madrone: it is common in northern California and southern
Oregon, where madrone is also common, less frequent in Washington, where madrone is
likewise less numerous, then common again on Vancouver Island in British Columbia,
where madrone is abundant. In our area it usually fruits after the first fall rains, sometimes
in tremendous quantity. The pale form, on the other hand, is a late winter and spring
mushroom. In our area it favors oak, but never fruits in the large quantities characteristic
of the fall (typical) form. In the Sierra Nevada the pale form is prevalent under pine and
other conifers, often forming “mushrumps” (especially in the spring).
EDIBILITY: Edible and popular, but with a rather strong, fishy flavor that doesn’t appeal
to everyone. Too many people eat and enjoy the typical form for me not to recommend it.
However, be absolutely sure of your identification, review the comments on pp. 264-265,
and avoid the pale and greenish forms, which are easily confused with poisonous species
(see photo on next page!). The caps are superb stuffed and then broiled. The hollow stems
can be sliced crosswise (to make rings) and marinated. The flesh does not keep well, so use
what you pick as soon as possible. Italian-Americans stalk it with a passion, undoubtedly
because of its resemblance—in both appearance and flavor—to their beloved A. caesarea
of Italy. “Coccora,” “coccoli,” and “cocconi” (another nickname) are presumably derived
from an Italian word for cocoon—a very apt description of the large, soft, cottony “eggs.”
COMMENTS: For many years this species has been called A. calyptroderma, but the
name A. calyptrata, first applied to the greenish form, may be the correct one according
to the International Code of Botanical Nomenclature (whose purpose it is to dispel con¬
fusion) because it was published first. The typical(darker) formand the pale formarequite
different in color, as evidenced by the color plates. I n my opinion they merit at least varietal
status (just like the varieties of A. muscaria) because they do not appear to intergrade and
are ecologically distinct (see comments above). The rare greenish form, on the other hand,
may be environmentally-induced (mycologist Harry Thiers suggests it is caused by cold
temperatures). The typical form of this magnificent mushroom is distinguished by its
(1) large size (2) orange to brown or yellow-brown cap with a yellow margin (3) cottony
285
Left: The pale winter-spring form of Amanita calyptrata. Right: Close-up of cap in Amanita ocreata
(p. 271-272). The pale form of A. calyptrata is easily confused with the deadly A. ocreata (see photos
on p. 272). A. ocreata usually lacks striations (fine lines) on the margin of the cap, but as evidenced
here, striate or partially striate specimens do occur—some of them with volval patches!

white volval patch or “skullcap” on the cap (4) striate cap margin (5) creamy or pale yellow
tints to the stalk and veil (6) thick, voluminous, saclike volva(7) non-amyloid spores. Even
beginners have little trouble recognizing the typical (darker) form once they’ve seen it
several times. In the deadly poisonous A. phalloides the cap is usually (but not always)
greenish to greenish-yellow or paler, there is no volval patch or if one is present it is very
thin, the cap margin is only very rarely striate, the spores are amyloid, and an unpleasant
pungent odor often develops in age. Veteran toadstool-testers can differentiate them at a
glance by their color, but beginners should not place undue emphasis on such a capricious
character. The veil tissue on the cap of A. gemma ta and A. pantherina can mimic a“patch,”
but does not peel off easily, and the volva in those species is collarlike or indistinct, not sac¬
like. The pale form of A. calyptrata is easily confused with poisonous Amanitas such as
A. ocreata and A. gemmata, and I cannot recommend it to any but the most seasoned and
intrepid toadstool-tester. The same goes for the greenish form. Another variety, originally
called A. calyptratoides, may be a distinct species. I have found it only once in our area, but
it is quite common in southern California under oaks. It sometimes approaches A. velosa in
size and stature, and has a very poorly formed, appressed or evanescent annulus (ring), a
brown cap without a yellow margin, and a modest volval patch. It is edible, but care must be
taken in identification!

Amanita velosa (Springtime Amanita) Color Plate 64

CAP 3-12(15) cm broad, at first nearly oval, soon convex and finally plane; surface viscid
when moist, smooth, pinkish-tan to orange-buff to salmon, beige, “the color of a brown
hen’s egg,” or sometimes much paler (even white), often fading; usually with a white patch
of universal veil tissue or several large, thick pieces; margin distinctly striate. Flesh
fairly thick, white, odor rather pungent in age. GILLS adnate to adnexed or free, close,
white, sometimes dull pinkish in old age. STALK 5-12(15) cm long, 0.5-1.5 (2.5) cm thick,
equal or tapering upward; white or tinged cap color, apex often powdery or striate,
lower portion smooth or broken into rings or scales. PARTIAL VEIL usually absent, but
a rudimentary ring (annulus) sometimes present as a cottony or fibrillose-scaly zone.
UNIVERSAL VEIL membranous, forming a saclike volva which sheathes the stalk base;
volva white or tinged cap color, usually buried, ample but sometimes disintegrating or
obscure. SPORE PRINT white or tinged very slightly pinkish; spores 8.5-12.5 * 6-10
microns, nearly round to elliptical, smooth, not amyloid.
AMANITA 287

HABITAT: Widely scattered to gregarious, common in winter and spring, associated in

our area with live oak but often growing out in the open (in fields and around their edges,
brushy areas, lawns, etc.), up to 40 feet away from its mycorrhizal host (which is often
stunted by poor soil conditions). It also occurs in the aspen-conifer forests of the Sierra
Nevada and with various oaks in the Sierra Nevada foothills and southern Oregon. It is
apparently endemic to the west coast and more common in our area than anywhere else.
With A. rubescens and A. ocreata it forms a striking triumvirate of springtime Amanitas
that are monogamous locally with live oak. It is interesting to note that when all three form
mycorrhiza with the same tree, they frequently occupy distinctly different zones or niches:
A. velosa typically grows in the open at the outer fringes of the host’s roots; A. ocreata
grows in deep shade near the trunk; and A. rubescens occupies the intermediate zone or
“shade border’’ near the perimeter of the branches.

EDIBILITY: Edible and incredible! Most “objective’’ fungophiles rate it far superior to
the better-known A. calyptrata and A. caesarea. At any rate, it is far sweeter. As a bonus,
it fruits in the spring when there is a paucity of other collectable delectables. However,
it is not a good choice for beginners. Faded specimens should be avoided, as they are
easily confused with A. ocreata and other poisonous Amanitas.

COMMENTS: The pinkish-tan to orange-buff, beige, or paler cap with striate margin and
volval patch or large warts, absence of a partial veil, saclike volva and habit of fruiting in the
open (unusual behavior for an Amanita!) are the fallible fieldmarks of this handsome but
variable fungus. In our area it is the last of its clan to appear—often not showing up until
February—but as if to make up for its tardiness, it lingers on through April or even May.
A. calyptrata and A. calyptratoides both have volval patches but are differently colored,
tend to grow in the woods, and have a partial veil. However, A. velosa often has a
rudimentary ring and one specimen I found had a full-fledged membranous partial veil that
covered the gills when young and broke to form an annulus (ring). This lends credence to
the modern trend toward de-emphasizing gross structural (“Friesian”) features and paying
more attention to chemical and anatomical (microscopic) similarities. The obsolete genus
Amanitopsis (Amanitas without a partial veil) was incorporated into Amanita onjust such
a pretext. The trend may bode ill for the multitudes without microscopes, but it better
reflects natural relationships. A. crocea, an eastern and Rocky Mountain species, is similar
and equally delicious, but has an oranger, usually bald cap and orangish scales on the stalk.

Amanitafulva (Tawny Grisette)

CAP 4-10 cm broad, oval becoming convex to plane or umbonate; surface slightly viscid
when moist, smooth or rarely with large whitish fragments of universal veil tissue;
orange-brown to reddish-tan, tawny, or tan; margin deeply grooved (striate). Flesh thin,
white, odor mild. GILLS close, adnexed or free, white or creamy. STALK 7-16 cm long,
0.4-1.5 cm thick, more or less equal, usually rather long, white or tinged cap color; fibrillose
or with a few scales, hollow in age. PARTIAL VEIL absent. UNIVERSAL VEIL
membranous, forming a large, loose, persistent, lobed, saclike volva which sheathes the
stalk base; volva or “bag” white to pale tan or rusty-stained. SPORE PRINT white; spores
8-10 microns, round, smooth, not amyloid.
HABITAT: Solitary to scattered or gregarious under both hardwoods and conifers, often
in boggy areas; widely distributed but not yet reported from California. I have seen large
fruitings in the aspen-conifer forests of the southern Rocky Mountains in late summer.
EDIBILITY: Edible, but be sure of your identification!
COMMENTS: This species is essentially a “fulvous” version of the grisette, A. vaginata,
and for many years was regarded as a mere color form of that species. The orange-brown to
Amanita fulva. Note conspicuously striate (grooved) cap and absence of a partial veil (annulus).
Cap color ranges from orange-brown to reddish-tan to tan.

reddish-tan, markedly grooved cap, absence of a partial veil, large “bag” at the base of the
stem, and rather tall, slender stature distinguish it. It is quite handsome when fresh. Other
species: A. crocea is similar but has an oranger cap and pale orange scales on the stalk.

Amanita vaginala (Grisette)

CAP 3-10 cm broad, at first oval, then convex and finally plane or with a slight umbo;
surface slightly viscid when moist, gray to grayish-brown, smooth or sometimes with a
white patch or patches of universal veil tissue; margin grooved (deeply striate). Flesh soft,
white to grayish, thin. GILLS white or tinged gray, adnate to adnexed or free, close.
ST ALK 7-15 (20) cm long, 0.5-2 cm thick, usually rather long and slender; equal or tapering
upward, smooth and white or often covered with delicate grayish to grayish-brown scales.
PARTIAL VEIL absent. UNIVERSAL VEIL membranous, forming a saclike volva that
sheathes the stalk but is attached only at the very base; volva white or tinged gray
(occasionally rusty-stained), loose, lobed. SPORE PRINT white; spores 8-12 microns,
round or nearly round, smooth, not amyloid.
HABITAT: Solitary to scattered or in small groups in woods or under trees; common and
widely distributed. In our area it occurs under both hardwoods and conifers, but is largely
supplanted by A. pachycolea and A. constrict a.
EDIBILITY: Edible when cooked, and fairly good. Though prized in France, it tends to be
flaccid, thin-fleshed, and does not refrigerate well. It is one of the safest Amanitas for the
table, but see comments on pp. 264-265 before eating it!
COMMENTS: The combination of deeply striate gray to gray-brown cap, white gills,
absence of a ring, and membranous sack or “bag” at the base of the stem typifies a group of
Amanitas collectively called A. vaginata. The group is especially complex in the South¬
west, where a number of forms with white to gray or brown caps are common in ponderosa
and pinyon pine forests after summer thundershowers. Several color forms of A. vaginata
have been described, some of which are now considered distinct species. One is A. fulva
(see description); another is A. alba, which has a pure white, grooved (striate) cap, a saclike
volva, and no partial veil. Although edible, A. alba should not be eaten because of its
resemblance to the destroying angels (A. ocreata, A. virosa, etc.). I have found it only twice
in our area, but it is said to be fairly common in some regions. A. vaginata has also been
called Amanitopsis vaginata and Vaginata plumbea.
Left: Amanita vaginata, mature specimens. The cap does not necessarily feature the small patch of
universal veil tissue shown here; note saclike volva. Right: Close-up of the constricted, flaring volva
of Amanita constricta (which otherwise resembles A. vaginata).

A manita constricta (Constricted Grisette)

CAP 4-13 cm broad, oval becoming convex or plane to slightly umbonate; surface slightly
viscid when moist, smooth or covered with a large patch of white to buff or grayish universal
veil tissue which often separates later into several pieces; color beneath the veil tissue (if
present) gray, or sometimes brownish-gray. Flesh fragile, rather thin, white to grayish.
GILLS adnate to adnexed or free, close, white or grayish. ST ALK 10-16 cm long, 1 -2 cm
thick, equal or tapering upward, stuffed or hollow in age; smooth or more often belted with
numerous delicate grayish scales. PARTIAL VEIL absent. UNIVERSAL VEIL mem¬
branous, gray to white or buff, often bruising reddish when wet; forming a constricted,
flaring volva (i.e., lower portion of volva constricted around the stalk, upper margin
flaring outward) which may disintegrate in age. SPORE PRINT white; spores 9.5-13 *
8-10.5 microns, elliptical to nearly round, smooth, not amyloid.
HABITAT: Solitary to scattered or in groups in woods, associated mainly with oaks;
apparently endemic to California or at least the west coast. Common in our area
throughout the mushroom season but most abundant in the winter.
EDIBILITY: Edible, but see comments on pp. 264-265 before eating it!
COMMENTS: The gray, conspicuously striate cap, absence of a partial veil, and peculiar
constricted volva are the hallmarks of this Amanita. It is the most common grisette
in our area. It is easily distinguished from both A. pachycolea and A. vaginata by its
constricted rather than saclike volva, and frequent presence of universal veil tissue on the
cap. A. inaurata(-A. strangulata, A. ceciliae) is a closely related species “complex” whose
volva is also “strangled,” usually forming a belt of grayish tissue around the stalk base.
It has a gray to grayish-brown to blackish cap decorated with gray to charcoal-gray warts
(which may wear off in age) and it has round spores(10-14 microns). It is fairly commonin
the Pacific Northwest as well as in eastern North America, but I have yet to find it in our
area.

289
Amanita pachycolea is easily told by its large size and dark brown to grayish-brown striate cap.
The cap is often darkest at the inner edge of the striations (a feature also shown in the color plates).

Amanita pachycolea (Western Grisette) Color Plates 65,66

CAP 7-20 (25) cm broad, at first nearly oval, then convex or somewhat bell-shaped, finally
plane or with uplifted margin and often a low, broad umbo; surface smooth, viscid when
moist, dark brown when young, brown to grayish-brown or paler in age (usually darker
at the center and paler toward margin or with a darker brown band at inner edge of the
striations); sometimes completely washed out in old age, occasionally with a thick patch or
patches of universal veil tissue; margin grooved (deeply striate) for 1.5-3 cm. Flesh white,
rather soft; odor sometimes unpleasant in age. GILLS close, broad, adnate becoming
adnexed or free; white with dark brown edges when fresh, sometimes discoloring dingy
orange or yellow-orange in old age. STALK 12-30 cm long or more, 1-3 cm thick, equal
or tapering upward gradually, usually covered with fine, delicate, grayish-brown to brown
particles or fibrillose scales on a pallid background; stuffed or hollow in age. PARTIAL
VEIL absent. UNIVERSAL VEIL membranous, forming a very large, loose, lobed,
saclike volva which sheathes lower portion of stalk for up to 12 cm but is attached only at
the very base; volva ample, thick, white or rusty-stained. SPORE PRINT white; spores
11-14.5 * 10-12.5 microns, broadly elliptical to round, smooth, not amyloid.
HABITAT: Solitary, scattered, or in Small groups in mixed woods and under conifers;
apparently restricted to the Pacific Coast. It is fairly common in our area in the fall and
winter with pine and oak, but rarely fruits in large numbers.
EDIBILITY: Edible, but not choice. Though meatier than A. vaginata, it develops a rather
strong fishy taste in age. See comments on pp. 264-265 if you plan to eat it.
COMMENTS: This lofty Amanita is easily recognized by its brown, deeply striate cap,
absence of a ring (partial veil), and huge sheathing “bag” at the base of the stem. It is one of
the most strikingly beautiful of all mushrooms, differing from its close relatives in the
A. vaginata group by its larger size, brown rather than gray cap, and frequent presence of
rusty stains on the huge volva. The volva is so voluminous that even washed out specimens
are easily recognized. The stalk in mature individuals is so long and fragile that it is difficult
to transport them home in one piece. Other species: A. umbrinolutea is similar, but is
said to have white gills, smaller spores, and a wide distribution.

290
 291

LIMACELLA
Medium-sized, mostly terrestrial fungi. CAP usually smooth and viscid or slimy. GILLS free or
nearly free, close, white or pallid. STALK typically central, hollow or stuffed, dry or viscid.
PARTIAL VEIL present, sometimes forming a ring, sometimes evanescent. UNIVERSAL VEIL
slimy, not forming a volva. SPORE PRINT white. Spores smooth, not amyloid (but rarely dex-
trinoid). Gill tissue divergent, at least when young.

THE viscid to slimy cap, presence of a veil, typically free gills, and absence of a volva typify
this small, rather rare genus. Many Limacellas used to be placed in Lepiota, but the
divergent gill tissue suggests a closer relationship to Amanita. The stature of the fruiting
body is somewhat like Amanita, but the universal veil takes the form of a layer of slime
that coats the cap and often the stem, and does not form a volva.
About a dozen species are known from North America. They are woodland fungi with
whimsical fruiting habits. Generally they are rare, but every so often there is a large,
localized fruiting. Little is known of their edibility. Five species are keyed here and two are
described.
Key to Limacella
1. Stalk distinctly viscid or slimy .2
1. Stalk not viscid (but patches of slime from cap may drip onto it) .4
2. Partial veil forming a distinct membranous annulus (ring) L. roseicremea(see L. illinita, p.292)
2. Veil slimy, not forming a membranous annulus . 3
3. Cap (and slime) entirely brown to bright reddish-brown to golden-brown.6
3. Cap white to creamy, or brownish at center and white toward margin .L. illinita, p. 292
4. Partial veil fibrillose, disappearing or forming a slight ring or ragged zone on stalk; cap reddish-
brown, chestnut-brown, pinkish-brown, or orange-brown .L. glioderma, below
4. Not as above; partial veil forming a distinct membranous (flaring or skirtlike) ring .5
5. Cap white or pale buff .L. solidipes(see L. illinita, p. 292)
5. Cap dull ochre to pale tan to creamy-pink; gills developing olive-gray stains in age or upon
drying in one variety, not developing them in another .L. guttata (-L. lenticularis)
6. Cap and stalk bright reddish-brown .L. glischra(see L. glioderma, below)
6. Cap and stalk more or less golden-brown.L. kauffmanii{see L. glioderma, below)

Limacella glioderma
CAP 3-8 cm broad, convex to plane or broadly umbonate; surface viscid to nearly dry,
bright to dull brick-red, reddish-brown, cinnamon, chestnut-brown, or at times orange-
brown, sometimes fading in age to pinkish-tan; cuticle often breaking up into small fibril¬
lose scales or pulling away from the margin, revealing the pinkish flesh underneath; margin
often hung with veil remnants. Flesh thin, tinged pinkish; odor distinctly but pleasantly
farinaceous. GILLS adnexed or notched, or in age becoming free; close, whitish or tinged
cap color. STALK 4-12 cm long, 0.5-1 (1.5) cm thick, often rather slender and fragile,
equal or slightly thicker at either end; dry or occasionally with a few patches of slime from
the cap; whitish or pallid above the ring, with cap-colored scales, patches, and/ or fibrils
below. PARTIAL VEIL fibrillose, whitish, forming a slight superior ring or ragged zone
on stalk or disappearing entirely. UNIVERSAL VEIL evanescent (usually not visible),
not forming a volva. SPORE PRINT white; spores 3-5 microns, round, smooth, not
amyloid. Gill tissue divergent.

HABITAT: Solitary, scattered, or in groups on ground in woods; widely distributed, and

the most common Limacella in our area. I find it regularly in the late fall and winter with
oak, madrone, and conifers.
EDIBILITY: Unknown, and like myself, likely to remain so.
Limacella glioderma. Note ragged or patchy stalk and evanescent partial veil.

COMMENTS: The viscid reddish-brown cap, evanescent veil, and dry fragile stem
distinguish this species from other Limacellas. Because the gills may be slightly attached
to the stalk, this fungus has been placed in Tricholoma and Armillaria. However, the
divergent gill hyphae (a microscopic feature) indicate Limacella. The prevalent form in
our area has only a slightly viscid cap, but fairly glutinous specimens are also encountered.
L. glischra is a widespread but rare species with a bright reddish-brown cap and stalk, both
of which are very slimy when moist. L. kauffmanii is a southern species with a more or less
golden-brown, viscid-slimy cap and stalk. Neither of these is worth eating.

Limacella illinita (White Limacella) Color Plate 67

CAP 2-8 cm broad, at first rounded or oval, then convex, finally plane or broadly umbo-
nate; surface smooth, very slimy or at least viscid, white to creamy-white in typical variety,
brownish at the center and white at the margin in var. argillacea', margin often hung with
slimy veil remnants. Flesh thin, soft, white. GILLS notched or free, white, close, ST ALK
5-9 (13) cm long, 0.5-1 cm thick, equal or tapering upward, slimy or viscid, white in typical
variety, but tinged tan or buff in var. argillacea. PARTIAL VEIL fibrillose beneath a layer
of slime, evanescent, not forming a distinct ring. UNIVERSAL VEIL slimy, coating stalk
and cap but not forming a volva. SPORE PRINT white; spores 4.5-6.5 x 4-6 microns,
broadly elliptical to round, smooth, not amyloid. Gill tissue divergent.
HABITAT: Scattered or in groups in woods, widely distributed; probably the most
common Limacella in North America. In our area the typical variety turns up occasionally
after heavy rains in the fall and winter in mixed woods and under Douglas-fir. I have seen
var. argillacea only once—a large fruiting under tanoak. The typical variety is also said
to occur in sand dunes. In the South it is said to be common on lawns as well as in woods.
EDIBILITY: Unknown. The slime is a formidable deterrent.
COMMENTS: The slimy white or brown-tinged cap and stem combine with the free gills
to set this slippery mushroom apart. Hygrophorus eburneus is just as glutinous, but has
adnate to decurrent gills and is much more common. L. solidipes is a widely distributed
but rare species with a viscid white cap, large flaring to skirtlike ring, and dry white stalk.
It also occurs in mixed woods in our area and could be mistaken for Lepiota naucina,
but the viscid cap and woodland habitat distinguish it. Other species: L. roseicremea,
known only from Washington, is similar but has a creamy or rose-tinged cap plus a dis¬
tinctly membranous partial veil and annulus (ring).

292
 spores
LEPIOTACEAE (Parasol Mushrooms)
Saprophytic, mostly terrestrial mushrooms of variable size. CAP typically dry or only slightly
viscid, often scaly with a smooth center when mature. GILLS typically free, white to pallid or
yellow, close. STALK cleanly separable from cap, central, base often enlarged. VEIL present,
usually forming a ring (annulus) on stalk, or if not then stalk usually scaly below the veil. VOLVA
typically absent. SPORE PRINT white to pale buff or dull greenish. Spores smooth, usually
dextrinoid, but not amyloid. Gill tissue parallel or interwoven.

THIS family is defined here to include one very large and diverse genus, Lepiota, plus a
single green-spored mushroom, Chlorophyllum molybdites, which could just as well be
considered an aberrant species of Lepiota. The “splitters” have erected several additional
genera for Lepiotas with thick-walled spores (e.g., Leucocoprinus, Leucoagaricus, and
Macrolepiota), but until one system of classification is firmly agreed upon, it seems best
for the purposes of this book to retain them in Lepiota.
Among the white-spored genera, Lepiota is most likely to be confused with Cystoderma
and Limacella—both of which were originally included in Lepiota. Cystoderma, however,
has attached rather than free gills plus a granulose cap and stalk, while Limacella has a
sticky or slimy cap. Lepiota also bears a superficial resemblance to Amanita, but lacks a
volva.* There are also fundamental microscopic differences: the gill tissue is parallel to
interwoven in Lepiota, divergent in Amanita (see p. 19), and the spores are typically
dextrinoid in Lepiota, while in Amanita they are frequently amyloid but never dextrinoid.
Aside from spore color, Lepiota closely resembles the common genus Agaricus (which
has chocolate-brown spores) in having free gills, a veil, and no volva. (Some taxonomists
even consider the Lepiotas to be a subfamily of the Agaricaceae.) There are ecological
similarities between Lepiota and Agaricus, as well as morphological ones. Both are large
genera centered in the warm temperate zone and tropics. (Lepiota is one of the largest and
most bewildering genera of tropical agarics.) And both are at their best in moist, relatively
warm weather. Furthermore, both are largely if not exclusively saprophytic—in contrast to
the Amanitas, which are mostly mycorrhizal. The larger Lepiotas or “parasol mushrooms”
frequent roadsides, waste places, lawns, gardens, pastures, and open woods. Many of the
smaller species occur in both heavily forested and cultivated areas. In our area, cypresses
are unusually rich in Lepiota (and Agaricus) species—some of them unclassified.
The large, white-spored parasols are among our most delicious edible mushrooms.
However, allergic reactions are reported for virtually every edible Lepiota, and extreme
care must be taken not to confuse them with poisonous species. L.procera of eastern North
America is the best of the best, L. rachodes and L. barssii are the best in the West, while L.
naucina is the best of the rest. The dozens of smaller Lepiotas should be strictly avoided.
Not only are they devilishly difficult to differentiate, but several (e.g., L. josserandii,
L. helveola, and L. castanea) are said to contain potentially fatal amanita-toxins! The
green-spored parasol, Chlorophyllum molybdites, can cause severe gastrointestinal
distress, and the powdery yellow greenhouse species, L. lutea, is also said to be poisonous.
At least 200 species of Lepiota are thought to occur in N orth America. Many have very
limited or erratic distributions. Only some of the larger or more distinctive species are
presented here. The diligent Lepiota-lover will doubtlessly uncover many species that
cannot be identified, particularly in California and the southern United States.

Key to the Lepiotaceae (Lepiota & Allies)

1. Fruiting body medium-sized to large; stalk usually (but not always) at least 6 mm thick; veil
usually forming a distinct collarlike or sleevelike annulus (ring) on stalk which may or may not
be thick, ragged, and double-edged . 2
1. Fruiting body small to medium-sized; stalk usually less than 6 mm thick; veil disappearing, or
if forming an annulus then the annulus not typically thick and double-edged . 10

* L. rachodes, however, may have a volva-like rim on its basal bulb.

294 LEPIOTACEAE

2. Both scales and background of cap white or whitish, the center often tinged yellow or yellow-
brown; margin distinctly striate at maturity; common in Gulf Coast region in many habitats;
spore print white; (may be very poisonous!) .L. humei
2. Not as above . 3
3. Cap (and often stalk or underside of veil) covered with pointed, often pyramidal warts (at least
when young) that usually rub off easily . 11
3. Not as above; scales on cap absent, or if present then caused by the breaking up of the cap cuticle
and not rubbing off easily. 4
4. Spore print greenish or grayish-green; usually growing in grass or gardens; fruiting in warm
or hot weather .Chlorophyllum molybdites, p. 295
4. Spore print white or pallid, not greenish or grayish . 5
5. Fruiting body tall (12-40 cm), cap 7-25 cm broad; stalk brown or breaking up into delicate brown
scales or granules below the ring (which may wear off); found in woods, old pastures, etc., in
eastern North America (also in southern California and the Southwest?) L. procera, p. 298
5. Not as above . 6
6. Cap smooth when young, but soon breaking up into brown to reddish scales or fibrils .... 7
6. Not as above; cap white to gray, grayish-brown, or buff . 9
7. Cap red to reddish-brown, pink, pinkish-orange, or cinnamon-buff, at least at center; fruiting
body not staining when bruised or in age; stalk typically up to 1 cm thick L. rubrotincta, p. 305
7. Not as above; stalk interior usually staining yellow-orange to reddish when cut or bruised . 8
8. Fruiting body aging or drying dark reddish-brown to vinaceous; stalk often spindle-shaped
(i.e., swollen above the base) .L. americana, p. 301
8. Not as above; stalk terminating in a swollen base or bulb .L.rachodes, p.297
9. Cap covered with flattened gray to grayish-brown fibrils when young, sometimes scaly in age
.L. barssii, p. 303
9. Cap white to gray or buff, usually smooth (but sometimes breaking up into small scales in age
and typically without fibrils when young) .L. naucina, p. 299
10. Cap (and sometimes stalk) covered with small, erect, pointed or pyramidal scales which rub off
easily; cap not striate. 11
10. Not as above . 13
11. Stalk with pointed scales below the veil . 12
11. Stalk with few if any scales (but underside of veil often has them); mainly found ineastern North
America and the Southwest .L. acutesquamosa (see L. eriophora, p. 303)
12. Veil usually forming a distinct annulus (ring) on stalk; stalk 1-2 cm thick .
.L. asperula (see L. eriophora, p. 303)
12. Veil disappearing in age; stalk 4-8 mm thick . L. eriophora, p. 303
13. Cap conspicuously striate, at least in age, the surface usually powdery or mealy or with small
scales when young; flesh thin and fragile; veil usually (but not always) forming a distinctive
sleevelike or collarlike annulus (ring) on stalk (examine several specimens) . 14
13. Not with above features; cap not normally striate . 16
14. Fruiting body yellow or mostly yellow, at least when fresh .L. lutea & others, p. 302
14. Fruiting body not yellow (but may stain yellow) . 15
15. Cap and base of stalk purplish to pinkish-brown or purple-brown when young, the cap surface
breaking up into purplish scales; found in nurseries and greenhouses.
. L. lilacinogranulosa (-Leucocoprinus lilacinogranulosus)
15. Not as above . L. cepaestipes& others, p. 301
16. Stalk shaggy or cottony below the veil; cap margin often shaggy also; growing in woods; spores
12-20 microns long .L. clypeolaria & others, p. 309
16. Not with above features . 17
17. Fruiting body whitish or with a brownish cap, the stalk typically staining reddish; common
in grazed pastures (but not on dung) in Florida . L. sp. (unidentified)*
17. Not as above . 18
18. Veil usually forming a distinct collarlike annulus (ring) on stalk (examine several specimens) 19
18. Veil usually disappearing or forming only a fibrillose or cottony zone or obscure annulus 29

* Popularly known as “Peele’s Lepiota” because it was discovered by Stephen Peele of Florida, this species is said
to be hallucinogenic; it should not be eaten until better known, because several similar species are poisonous!
LEPIOTA& ALLIES 295

19. Cap, stalk, and/or flesh staining bright orange to red (or staining yellow and then orange to
reddish) when bruised or rubbed, then often discoloring to dark brown or purplish.20
19. Not as above . 24
20. Fruiting body aging or drying dark reddish to vinaceous; stalk usually spindle-shaped (i.e.,
swollen above or at the base); cap often with coarse scales; growing in compost, sawdust,
around old stumps, etc., usually in groups or clusters .L. americana & others, p. 301
20. Not as above . 21
21. Tropical or found along Gulf of Mexico; staining yellow before reddening or darkening . 22
21. Not as above; not staining yellow . 23
22. Gills tinged pale yellow; center of cap and scales on cap blackish-brown .L. sanguiflua
22. Not as above .L. tinctoria & others (see L. americana, p. 301)
23. Gills bruising pinkish or red-orange . . . . L. roseifolia & others (see L. flammeatincta, p. 304)
23. Gills not staining when bruised .L. flammeatincta & others, p. 304
24. Cap black or dark gray at center, usually with scattered gray to greenish-gray scales .
...L. atrodisca, p. 304
24. Not as above; cap scales not gray or black . 25
25. Cap white to yellowish or pale tan . 26
25. Cap purple, red, pink, reddish-brown, or brown, at least at the center . 27
26. Taste strongly acidic; cap typically with small scales in age; usually growing with alder.
. L. pulcherrima
26. N ot as above; cap usually smooth, yellowish to tan or grayish at center and paler toward margin
.L. sequoiarum & others, p. 307
27. Cap or fibrils on cap purplish . L. roseilivida(see L. rubrotincta, p. 305 & L. cristata, p. 306)
27. Not as above; cap (or scales) some shade of red, pink, orange, brown, etc.28
28. Cap 3-8 cm broad, typically with radiating fibrils .L. rubrotincta & others, p. 305
28. Cap usually 5 cm broad or less, with small scattered or concentrically arranged scales ... 32
29. Gills yellow .L. luteophylla (see L. lutea, p. 302)
29. Gills not yellow . 30
30. Cap and stalk tinged lavender to lilac, without scales; odor unpleasant; growing in woods;
rare . L. bucknallii
30. Not as above . 31
31. Cap small (usually 2.5 cm broad or less), white or tinged pinkish to pinkish-cinnamon; smooth
or minutely powdery but without contrasting colored scales .L. seminuda, p. 307
31. Not as above . 32
32. Stalk typically without scales; cap small (usually 1-5 cm broad); annulus (ring) often present
on stalk .L. cristata & others, p. 306
32. Stalk typically with scales below the veil or cap larger; distinct annulus usually lacking . . 33
33. Stalk and often the cap developing strong ochraceous to rusty-orange tones in age or after
handling; fruiting body small (stalk typically 1-3 mm thick) . L. castanea, p. 307
33. Not as above (but reddish tones may develop); fruiting body small to medium-sized . 34
34. Scales on cap and stalk blackish to dark brown.L. felina (see L. josserandii, p. 308)
34. Not as above; scales brown to reddish .L. josserandii & others, p. 308

Chlorophyllum molybdites (Green-Spored Parasol)

CAP (5) 10-30 (40) cm broad, oval or nearly round, then convex to broadly cone-shaped,
plane, or umbonate; surface dry, at first smooth but soon breaking up into light brown to
brown or pinkish-brown scales on a white background; scales flat or curled, usually rather
few at maturity and concentrated toward the center. Flesh thick, white, soft in age; not
staining when bruised or bruising slowly reddish or occasionally bruising orange in the
stalk. GILLS free, broad, close, white to dingy yellowish, then slowly becoming grayish
to greenish in old age. STALK 5-25 cm long, 1-2.5 cm thick at apex, equal or thicker at base;
smooth, firm, white or brownish-stained. VEIL membranous, white, formingapersistent,
superior, double-edged ring on stalk; ring becoming brownish on underside and usually
Chlorophyllum molybdites is best distinguished from other parasol mushrooms by its greenish spore
print. Note how the gills are still white in the fully expanded specimen in background.

movable in age. SPORE PRINT grayish-olive to greenish; spores 8-13 * 6.5-9 microns,
elliptical, smooth, thick-walled, with an apical pore, dextrinoid. Cap cuticle composed
of narrow, interwoven, mostly repent hyphae (but often upright at center of cap).
HABITAT: Solitary to scattered or in groups or large rings on lawns and other grassy
places, also in gardens; fruiting in summer or during warm weather and widely distributed
in the tropics and warm temperate zone. It is common in most of eastern and southern
North America as well as inland northern California. I have seen enormous fruitings on
lawns in Fresno, Los Angeles, Palo Alto, and San Diego, but have never seen it on the
central California coast, perhaps because of the cool summers.
EDIBILITY: Poisonous! Some people eat it without ill effect but many suffer severe
gastrointestinal distress. It is probably the most common cause of mushroom poisoning in
the U nited States, a tribute to its growth on lawns, tempting size(it borders on being irresis-

Chlorophyllum molybdites growing on a lawn. N ote broadly cone-shaped cap of mature specimen at
top right, and drumstick shape of buttons in foreground. Gills at far right have started to darken.
CHLOROPHYLLUM 297

tible) and resemblance to Lepiota rachodes, Coprinus comatus, Agaricus species, etc.
COMMENTS: This distinctive summer mushroom always attracts attention because of
its large size and handsome appearance (frisbee-sized specimens are commonplace). In
much of inland and southern California and the Southwest, it is one of the commonest
urban lawn mushrooms or “toadstools.” It so closely resembles other large Lepiotas that
some lepiotologists merely consider it an aberrant species with greenish spores (i.e.,
Lepiota molybdites). It is most likely to be mistaken for Lepiota rachodes, especially in the
button stage. It differs, however, in its habitat (usually grass), fondness for hot weather, less
pronounced staining reactions (though this character is variable), and less bulbous stem.
The only completely reliable feature, however, is the spore color. But beware—the gills may
remain whitish well into maturity, so that making a spore print is the only sure way to
determine the spore color! Young buttons look like drumsticks when they first emerge from
the ground. They may not show any white on the cap, whereas adults may be almost entirely
white. Lepiota morgani is an older name for it.

Lepiota rachodes (Shaggy Parasol) Color Plate 69

CAP 5-20 cm broad, oval to convex or marshmallow-shaped, becoming plane in age or
slightly umbonate; surface dry, at first pale brown to reddish-brown, cinnamon-brown,
or brown, soon breaking up into large, coarse or shaggy scales as the cap expands, revealing
the white to dingy buff background, the center usually remaining smooth and brown;
margin usually fringed or shaggy. Flesh thick, white, fairly firm, typically bruising yellow-
orange to orange and then reddish or brown when cut. GILLS broad, free, white, close,
sometimes dingy brownish in old age or when handled. STALK 5-18 cm long, 1-3 cm thick
at apex, enlarged below (var. rachodes), or with a large, abrupt basal bulb which may have
a raised rim {var. hortensis); white when young, developing brownish stains in age or upon
handling; smooth, dry; interior usually staining bright orange or saffron and then reddish
to dark brown when cut. VEIL membranous, white or with a brown, ragged margin;
forming a large, thick, collarlike, double-edged, superior ring on stalk which is usually
movable in age. SPORE PRINT white; spores 6-13 * 5-9 microns, elliptical, smooth, with
a large apical pore, thick-walled, dextrinoid. Cap cuticle composed of enlarged, erect cells.
HABITAT: Usually in groups or rings on ground under trees (particularly conifers) and
bushes, in gardens and compost piles, near stables, on ant hills, along roads and other
disturbed places, even in basements and greenhouses, sometimes also in open fields or in

Lepiota rachodes var. hortensis differs from the typical variety (shown in color plate) by having a
more abrupt and pronounced, even rimmed (volva-like) basal bulb. N ote how cap cuticle is smooth at
first, then cracks into scales. Specimen at far right has been sliced to show the staining of the flesh.
298 LEPIOTACEAE

the woods; widely distributed, but most common in western North America. It fruits
whenever conditions are favorable, i.e. moist and mild, and often produces several crops a
year. “Patches” are not particularly numerous, but can be very prolific—I’ve seen several
hundred specimens under a single tree! Variety hortensis is quite common in our area,
especially under planted conifers (e.g., cypress) and in sandy soil along the coast (I have
found it on Ano Nuevo Island in sand and elephant seal dung, and in the splash zone at
Pebble Beach!). The typical variety is more common in the Pacific Northwest.
EDIBILITY: Edible and excellent, with caution. A number of people have had severe
“allergic” reactions to it, particularly on the west coast, and it is easily confused with
Chlorophyllum molybdites. It has an exceptionally strong, nutty flavor but the water
content is high. For the best and safest results, fry it on high heat in an open pan. Delicious!

COMMENTS: The outstanding features of this outstanding mushroom are the large
brown scales on the cap, free white gills, prominent collarlike ring (not skirtlike as in
Amanital), basal bulb or thickened stem base, and brusing of the flesh to orange and then
reddish. (The latter feature is best seen by cutting the stem.) The brightness and duration
of the color changes vary according to the moisture content and age of the mushroom. In
variety hortensis (the common one in our area), the basal bulb may have a raised rim which
can be mistaken for a volva. However, the brown cap scales and collarlike ring point to
Lepiota. The poisonous Chlorophyllum molybdites looks very similar, but has dull
greenish spores. L. rachodes (sometimes spelled L. rhacodes and also known as Macro-
lepiota rachodes and Leucoagaricus rachodes) can also be mistaken for an Agaricus
(especially A. augustus), but the white spores and gills distinguish it.

Lepiotaprocera (Parasol Mushroom)

CAP 7-25 cm broad or more, at first oval, then expanding to convex, plane, or umbonate;
surface dry, at first smooth and brown, soon breaking up into brown scales and patches
except for the smooth, dark central umbo; flesh between the scales at first white but soon
weathering to buff, grayish, or brownish and usually with a shaggy or torn-up appearance.
Flesh white or tinged reddish, but not staining orange or red when cut; soft in age. GILLS
free, broad, close, white when young, but sometimes discoloring to pinkish, tan, or dingy
brownish in old age. STALK 12-40 cm long, 0.8-1.5 cm thick, typically very long and
relatively slender with an enlarged base; pallid but the surface below the ring covered with
numerous small brown scales or flakes which often separate into belts or wear away in
age. VEIL membranous, forminga thick, superior, brownand white, double-edged ring on
stalk; ring collarlike and movable. SPORE PRINT white; spores 12-18 x 8-12 microns,
broadly elliptical, thick-walled, with an apical pore, smooth, dextrinoid.
HABITAT: Solitary to widely scattered or in small groups in open woods and at their
edges, in old pastures, along trails, etc.; fairly common in the summer and fall in eastern
North America (especially New England and the South) and Mexico. It (or something
similar) also occurs in southern California, and it may very well grow in southern Arizona
and New Mexico, where a number of so-called “eastern” species (e.g., Lactarius indigo,
Strobilomycesfloccopus) occur.
EDIBILITY: Edible and one of the very best of all agarics! Prized by connoisseurs for its
strong, meaty-nutty flavor, it is now being grown commercially in Europe. It does not rot
readily and is usually free of maggots. The leathery, sun-dried specimens frequently found
in dry weather can be just as good as fresh ones if treated properly.

COMMENTS: Also known as Macrolepiota procera and Leucoagaricus procerus, this

lofty, imposing mushroom is the most distinctive of all the Lepiotas. It is also the safest
and the tastiest, although “allergic” reactions have been reported. It is much taller than L.
The parasol mushroom, Lepiota procera. Left Side view of mature specimen. Right: T op view of two
mature specimens. The fruiting bodies look like long drumsticks before the cap opens out.

rachodes, and doesn’t stain red or orange when cut. The spore print is white (not greenish
as in Chlorophyllum molybdites), and the brown stalk that breaks up into delicate
branlike scales (which may wear away) is also distinctive. There is no volva at the base of the
stalk as in Amanita, and the solid brown “nipple” or umbo at the center of the cap is also
distinctive. The stalk is so slender in relation to its height that it is not uncommon for large
specimens to topple over in old age from the weight of the cap.

Lepiota naucina (Smooth Parasol; Woman On Motorcycle)

CAP 4-10 (15) cm broad, oval to nearly round when young, then broadly convex to plane;
surface dry, typically dull white but at times gray, buff, or creamy, the center sometimes
tinged pinkish-buff; usually smooth, but sometimes with numerous small branlike par¬
ticles, or breaking up into scales in age; one form staining yellow when handled, another
brown. Flesh thick, white, not bruising; odor mild, or in one form unpleasant. GILLS free,
close, white, but often becoming buff, pinkish, or grayish-pink in old age, and finally
brownish. STALK 5-15 cm long, 0.5-1.5 cm thick, equal or with an enlarged base; dry,
white, without scales, sometimes staining yellow when bruised, and usually discoloring
brownish in age or upon handling. VEIL membranous, white, forming a distinct,
persistent, superior, double-edged, collarlike or sleevelike ring on stalk which is usually
movable in age (and may fall off). SPORE PRINT white or very faintly pinkish; spores 7-9 x
5-6 microns, broadly elliptical, thick-walled, smooth, with an apical pore, dextrinoid.
HABITAT: Solitary to scattered or gregarious in grassy areas (lawns, pastures, etc.),
sometimes also along roads, freeways, and in other disturbed places, and sometimes
under trees or even in the woods; widely distributed and common. In our area it is often
abundant in the fall and early winter, and occasionally encountered in the spring and
summer as well.
EDIBILITY: Edible and very good, but not recommended. Either some persons are sen¬
sitive to it, or certain variants (perhaps the yellow-staining or ill-smelling ones) are toxic.
According to one source, it is one of the most frequent causes of mushroom poisoning in the
Pacific Northwest, yet it is listed as “edible” in every mushroom book! One former friend
of mine who had eaten it previously was made quite ill by a cream-of-L. naucina soup
which I had painstakingly prepared. There is also the more serious danger of carelessly
confusing it with a deadly Amanital
Lepiota naucina in its favorite milieu: grass. Note free gills and annulus. (Nancy Burnett)

COMMENTS: The white to grayish cap, membranous annulus (ring), free gills, white
spores, and fondness for grass are the hallmarks of this cosmopolitan mushroom. Its
appearance on lawns and in cemeteries marks the beginning of our fall mushroom season.
A good way to become acquainted with it is to bicycle around town: there it will be—tall,
white, stately, in graceful groups on lawns. Later it appears in pastures. It is a beautiful
mushroom when young, the smooth unexpanded cap being reminiscent of a motorcycle
helmet. The shape, though difficult to describe, is very distinctive (see photographs).
Though the stalk base is often enlarged, there is never a sack (volva) as in the deadly
Amanitas, and the ring is not skirtlike, nor is the cap viscid as in Limacella. The staining
reactions are also an important fieldmark. In some specimens the cap and stem stain yellow
quite dramatically in the tradition of Agaricus xanthodermus. Usually, however, they
discolor yellow-brown to brown after handling and in age—especially on the stem. The
frequent darkening of the old gills to pink or brown leads to confusion with Agaricus,
and once again points out the folly of equating gill color with spore color. I have also seen
buttons of Agaricus californicus interspersed with L. naucina that were virtually indis¬
tinguishable. A final hint: If your “L. naucina” fails to turn brown when cooked, double¬
check your identification, because you may have something else! Lepiota leucothites,
L. naucinoides, and Leucoagaricus naucinus are synonyms.

Lepiota naucina, sometimes called the “ Woman on Motorcycle” because of the helmet-shaped young
caps.
Lepiota americana, a distinctive reddening species with a spindle-shaped stalk.

Lepiota americana (American Parasol)

CAP 3-15 cm broad, oval becoming convex, plane, or broadly umbonate; surface dry,
smooth at first, soon breaking into coarse vinaceous- to reddish-brown or pinkish-buff
scales, the center usually remaining smooth; background white, but often reddening with
age. Flesh white, bruising yellow to orange when young and fresh (especially in stalk), but
aging or drying reddish to vinaceous. GILLS free, close, white, but may stain or age like
the flesh. STALK 7-14 cm long, 0.5-2 cm thick, enlarged at base or often spindle-shaped
(swollen at or below the middle, with a narrowed base); smooth, at first white, but aging or
drying reddish to dark vinaceous. VEIL membranous, forming a white, double-edged,
superior ring on stalk which may disappear in age. SPORE PRINT white; spores 8-14 * 5-
10 microns, elliptical, smooth, with an apical pore, thick-walled, dextrinoid.
HABITAT: In groups or clusters in sawdust and compost piles, around old stumps, in
waste places, rich soil, etc. Widely distributed and fairly common in eastern North
America, but rare in California. I have found it only once in our area, growing with
Agaricus subrufescens, in the summer and early fall.
EDIBILITY: Edible, but be sure of your identification! I haven’t tried it.
COMMENTS: This species is easily recognized by its tendency to darken to reddish or
burgundy as it ages or dries, and the yellow to orange staining of fresh specimens (best
seen by cutting the stalk). In addition, the shape of the stalk is quite unusual—often very
slender at the apex and blatantly bulbous at or toward the base (see photo).L. badhamiiof
Europe is very similar, but is said to be poisonous. There are also several closely related
species in the southeastern U.S., including one unnamed variant with pale green spores
and several species (e.g., L. tinctoria) with smaller, paler, and more numerous cap scales.

Lepiota cepaestipes (Onion-Stalk Parasol)

CAP 2 -8 cm broad when expanded, oval becoming broadly conical or bell-shaped to nearly
plane or umbonate, eventually drooping; surface dry, powdery or mealy becoming some¬
what scaly or fibrillose-scaly in age, white to pale pinkish (but may be darker when young
and yellowish to brownish in age); margin clearly striate at maturity. Flesh thin, white,
sometimes bruising yellowish. GILLS white, crowded, free. STALK 4-14 cm long, 3-6
mm thick, equal or swollen in places, with an enlarged base; smooth or powdery, slender,

301
302 LEPIOTACEAE

white, but may discolor yellowish when handled. VEIL white, forming a persistent,
superior but easily detachable ring on stalk. SPORE PRINT white; spores 6-10 x 5-8
microns, elliptical, with an apical pore, smooth, thick-walled, weakly dextrinoid.
HABITAT: In groups or clusters in rich soil, wood chips, around old stumps, straw piles,
gardens—in other words, in decomposing organic matter of almost any kind; widely
distributed, but much more common in eastern North America than in the West. In our
area it fruits in the summer or during warm, moist weather.
EDIBILITY: Not recommended. Though traditionally listed as edible, it has adverse
effects on some people and is very thin-fleshed besides.
COMMENTS: Also known a.s Leucocoprinus cepaestipes, this species is easily recognized
by its mealy, whitish, striate cap. It is essentially a whitish version of L. lutea, and the two
used to be considered color forms of the same species. There are many closely related tropi¬
cal and southern species, including: L. breviramus, similar but with soft warts on the cap
(including the center) and usually growing scattered or tufted; L. longistriatus, with a tan
to pale tan fibrillose cap; and L. brebissonii, with a small (2-3 cm) white cap with a dark
gray to brownish center. I have found the latter on lawns in Berkeley, California, in the
summer. Because of their mealy, striate caps, all of these species are placed in their own
genus, Leucocoprinus, by many mycologists.

Lepiota lutea (Yellow Parasol; Flower Pot Parasol) Color Plate 70

CAP 2 5 -6 cm broad when expanded, oval becoming broadly conical or bell-shaped, then
eventually umbonate or plane and finally drooping; surface dry, powdery, mealy, and/or
minutely scaly, usually scalier and less powdery in age; bright yellow to greenish-yellow
or pale yellow, the center sometimes brown or buff; fading quickly after it matures or
becoming browner; margin conspicuously striate nearly to center at maturity. Flesh very
thin, yellow. GILLS free, crowded, yellow or pale yellow. STALK 3-10 cm long, 1.5-5 mm
thick, usually slender and enlarged somewhat at or toward base base; dry, smooth or
powdery like the cap, yellow. VEIL yellow, forming a small, superior, collarlike ring on
stalk which may disappear. SPORE PRINT white; spores 8-13 x 5.5-8 microns, elliptical,
with an apical pore, thick-walled, smooth, dextrinoid.
HABITAT: Solitary, tufted, or in groups in flower pots, greenhouses, and planter boxes, or
if it is warm enough, outdoors (in lawns, gardens, etc.); widely distributed, fruiting indoors
most anytime, outdoors mainly in the summer. The specimens in the color plate were
growing in a planter box in front of the Bank of America in downtown Los Angeles.
EDIBILITY: Poisonous to some people, according to some sources.
COMMENTS: This brilliant yellow greenhouse mushroom boasts a plethora of pseudo¬
nyms, including Leucocoprinus luteus, L. birnbaumii, and L. cepaestipes var. luteus. It
can be the object of considerable consternation to plant lovers when it pokes up in one of
their flower pots. However, it won’t hurt the plant (or you, unless you eat it). If one should
appear, consider yourself lucky and take advantage of the situation—sprinkle it lightly
as you would any other houseplant, and watch new individuals develop from tiny “pin¬
heads” within a few days. The bright yellow color and striate cap set apart L. lutea from
other Lepiotas. It has a Coprinus-like stature, but the spores are white and it doesn’t deli¬
quesce (though like many tropical fungi, it withers quickly). Bolbitius vitellinus is also
yellow, but has a slimy-viscid cap. Other species: L. flavescens is another yellow green¬
house species with smaller, nearly round spores (5-7 microns broad); L.fragilissimus has a
very thin, fragile cap and pale yellow to white gills and is a tropical and subtropical wood¬
land species. Lepiota luteophylla has a brownish, non-striate cap and bright yellow gills.
It was originally found in California but has also turned up in Michigan.
LEPIOTA 303

Lepiota eriophora (Sharp-Scaled Parasol)

CAP 2-7.5 cm broad, oval or convex when young, broadly convex to umbonate or plane at
maturity; surface dry, white to buff, covered with small, erect, pointed brown scales which
rub off easily. Flesh white, rather thin, not staining. GILLS free, white to buff or tinged
pinkish, close. STALK 2-10 cm long, 3-7 (10) mm thick, equal or enlarged slightly at base,
dry, smooth at apex, covered with brown scales (like cap) below, but these often wearing
away in age. VEIL evanescent, not forming a membranous ring on stalk, but sometimes
leaving a fibrillose zone. SPORE PRINT white; spores 3-6 * 2-3 microns, oblong, smooth,
dextrinoid.
HABITAT: Solitary, scattered, or in small groups in woods and in rich soil, widely
distributed. Occasional in our area in the fall and winter, especially under cypress.
EDIBILITY: Unknown.
COMMENTS: This is one of a number of confusing, Amanita-like Lepiotas with erect,
pointed scales on the cap. It was described in the first edition under the namel. hispida,
a possible synonym. Closely related species include: L. asperula, with an annulus (ring) and
stalk 1 -2 cm thick; L. scabrivelata, a small subtropical species with yellow warts on the cap
and stalk; and L. acutesquamosa, medium-sized to large (cap 5-15 cm) with large brown
warts on the cap but few if any on the stalk, and veil often with brown scales on its under¬
side but not necessarily forming an annulus (ring). The latter, which is said to be edible,
is fairly common in eastern North America and also occurs in the Southwest.

Lepiota barssii (Gray Parasol)

CAP 4-15 cm broad or more, nearly round to convex, becoming broadly convex to plane or
umbonate; surface dry, covered with fine, flattened gray to brownish-gray radiating fibrils,
the center often darker; sometimes becoming scaly in age or direct sunlight. Flesh fairly
thick, white to grayish, not staining when bruised. GILLS close, free, white or staining
dingy yellowish to brownish. STALK 5-13 cm long, 0.5-2.5 cm thick, equal or more often
swollen below, but with a tapered base (no bulb!); smooth; white or stained brownish.
VEIL membranous, white, forming a superior, collarlike ring on stalk which may be
movable or drop off at maturity. SPORE PRINT white; spores 7-11 * 5-6 microns,
elliptical, smooth, dextrinoid.
HABITAT: Scattered to gregarious in cultivated or composted soil, lawns, gardens,
pastures, and plowed fields; known only from the west coast. It is fairly common in
Washington and Oregon, but rather infrequent in our area, where it fruits mainly in the
summer and fall.
EDIBILITY: Edible and choice—but be sure of your identification!
COMMENTS: The grayish fibrillose cap, membranous ring, free white gills, absence of
a basal bulb on the stalk, and habitat in cultivated ground are the telltale traits of this
fine fungus. The stalk is often buried deep in the soil, especially if it is growing in mulch.
S pecimens growing on lawns or hard soil tend to be smaller, with a shorter stem. The flesh
does not stain noticeably when cut as in L. rachodes. The veil is persistent but may be
obliterated, in which case the free gills are an important fieldmark. It is much stouter and
paler than L. atrodisca, and does not grow in the woods. It is darker and more fibrillose
than L. naucina, and it can also be confused with an unidentified Amanita(p. 275) which
has warts on the cap, amyloid spores, and yellowish gills in old age. In the W illamette Valley
in Oregon L. barssii is quite common and frequently attains a large size; most of the speci¬
mens I’ve seen in California, however, have been smaller, averaging 4-10 cm broad. Other
species: L. excoriata is vaguely similar, but has a cuticle that recedes from the margin as
the cap expands.
Lepiota atrodisca has black to greenish-gray cap scales and a smooth stem. Note sleevelike annulus.

Lepiota atrodisca (Black-Eyed Parasol)

CAP 1 -5 (6.5) cm broad, oval or convex becoming broadly umbonate or plane, the margin
sometimes uplifted in age; surface dry, white with flattened black, gray, or greenish-gray
scales or fibrils, the center usually darker(blackish). Flesh thin, white, not bruising. GILLS
free, white or creamy, close. STALK 2.5-10 cm long, 3-7 mm thick, usually slender but
sometimes rather stout, enlarged somewhat at the base; smooth, dry, white or discoloring
slightly upon handling. VEIL membranous, forming a fragile, sleevelike, white or black-
edged ring at or above middle of stalk, or sometimes disappearing. SPORE PRINT white;
spores 6-8 * 3-5 microns, elliptical, smooth, dextrinoid.
HABITAT: Solitary, scattered, or in small groups on ground or rotting wood under both
hardwoods and conifers, fall and winter, apparently endemic to the west coast. During
cold, dry weather I have seen enormous numbers in woodland thickets (oak-manzanita-
hazel nut)—an unusual abode for a Lepiota—but it is not restricted to that habitat.
EDIBILITY: Unknown—do not experiment! It is too small to be of importance anyway.
COMMENTS: This attractive little woodland Lepiota is easily recognized by its unusual
grayish-black scales. L. felina (see comments under L. josserandii) has dark brown to
blackish scales on both the cap and stem, and an evanescent veil; it is poisonous.

Lepiota flammeatincta (Flaming Parasol)

CAP 1.5-5 (7.5) cm broad, convex when young, then plane or with uplifted margin or
sometimes broadly umbonate; surface dry, with nearly black to dark purple-brown to
brown or reddish-brown fibrils or scales (sometimes very sparse) on a whitish background;
surface quickly staining scarlet to scarlet-orange when bruised, then slowly turning dark
brown or dark purple-brown. Flesh white, staining slightly pinkish, reddish, or orange
when bruised, then fading. GILLS free, close, white, not staining when bruised. STALK
3-10 (15) cm long, 2-6 mm thick, equal or slightly thicker below, slender, white above the
ring, fibrillose (like cap) below and quickly bruising scarlet to scarlet-orange like the cap
surface, then discoloring dark brown. VEIL membranous, white, forming a median to
superior, sleevelike ring on stalk, or disappearing. SPORE PRINT white; spores 6-8.5
* 4-5 microns, elliptical, smooth, dextrinoid.
HABITAT: Solitary or in small groups in woods and under trees, along trails, etc.; not
uncommon along the Pacific Coast. In our area it fruits in the fall and early winter, but
seldom in large numbers. L. roseifolia(see comments) is also fairly common.

EDIBILITY: Unknown.
COMMENTS: This is one of our most striking Lepiotas because of the spectacular scarlet-
304
 Vm

Lepiota flammeatincta. Surface of cap and stalk stain bright reddish-orange when rubbed, but gills
do not. A very similar species, L. roseifolia (not illustrated) has pinkish-staining gills.

staining (“flaming”) of the cap and stem. The dark brown color that wounded areas sub¬
sequently assume is also characteristic. L. roseifolia is a very similar species whose gills
turn pinkish or red-orange when bruised (within five minutes); it favors cypress in our area.
An unidentified local species with a lovely purple to vinaceous-pink cap when young
(browner in age) also occurs, usually under oak. Its cap and especially the stalk sometimes
stain orange or orange-red when rubbed. L. brunnescens and L. roseatincta are two
eastern scarlet-staining species; the latter has a pinkish-red cap.

Lepiota rubrotincta (Red-Eyed Parasol)

CAP 3-8 cm broad, oval or rounded becoming convex, finally plane or broadly umbonate
or with an uplifted margin; surface dry, at first uniformly pinkish-brown or reddish, then
breaking up into flat, radially arranged fibrils or scales which vary in color from cinnamon-
buff to coral-pink, reddish, or pinkish-orange; background whitish and the center remain¬
ing smooth and usually darker (deep red to chestnut); margin often splitting in age. Flesh
Lepiota rubrotincta. Cap has fibrils and a smooth dark center; common in the fall. (Joel Leivick)
306 LEPIOTACEAE

thin, white, not bruising. GILLS free, white, close, not bruising. STALK 4-16 cm long,
0.4-1 cm thick, usually rather slender and equal or thicker below, often extending fairly
deep into the humus; smooth, white, discoloring somewhat in age and becoming hollow.
VEIL membranous, white, forming a thin, fragile but persistent ring on stalk; ring median
to superior, typically sleevelike above and flaring below. SPORE PRINT white; spores
6-10 * 4-6 microns, elliptical, smooth, dextrinoid(?).
HABITAT: Solitary to scattered or in small groups in humus, usually in woods; widely
distributed and sometimes common in our area, especially after the first fall rains.
EDIBILITY: Not firmly established. Katy Caldwell of Santa Cruz claims to have eaten
a small quantity without ill effect, but it is easily confused with other Lepiotas, some of
which are poisonous.
COMMENTS: This beautiful mushroom is one of our more common and easily identified
woodland Lepiotas. It is larger and taller than L. cristata, and has a more persistent ring.
The color of the cap fibrils varies considerably, but usually has a reddish tone. The smooth,
dark center is suggestive of a human breast. Other species: L. roseilivida is somewhat
similar but smaller, has a purplish cap, and occurs under various western conifers;/,, glat-
felteri has a vinaceous-brown fibrillose cap and has been found in rich soil and under
cypress in Santa Barbara. Also similar is the beautiful unidentified species mentioned
under L. flammeatincta; its cap is purple or pink at first and its stalk often bruises orange.

Lepiota cristata (Brown-Eyed Parasol)

CAP 1-5 (7) cm broad, convex becoming broadly convex to plane or broadly umbonate;
surface dry, soon breaking up into small tawny to brown or red-brown scales (often
concentrically arranged) on a white background, the center usually remaining smooth and
darker(brownto red-brown). Flesh thin, white, not bruising; odor mild or sweet and fruity
or pungent. GILLS free, white to buff, close. ST ALK 2-8 cm long, 2-5 mm thick, equal or
thicker below, slender, white to pinkish-buff and often darker toward base; more or less
smooth, fragile. VEIL white, sometimes disappearing, at other times forming a thin,
fragile, median to superior ring on stalk. SPORE PRINT white to pale buff; spores 5-8 *
3-5 microns, bicornute (wedge-shaped and spurred at one end), smooth, dextrinoid.
HABITAT: Scattered or in groups on ground in woods, under trees (especially redwood
and cypress) or shrubs, on lawns, etc. Widespread; common in our area in fall and winter.
EDIBILITY: To be avoided—perhaps poisonous.

Left: Lepiota cristata. Note absence of prominent scales on stalk. Right: Lepiota seminuda, our
smallest Lepiota. Note the free whitish gills and fragile veil that leaves remnants on cap margin.
LEPIOTA 307

COMMENTS: The chestnut to dark brown cap center, fragile ring, small size, and smooth
stem characterize a number of Lepiotas which can only be differentiated microscopically.
L. castaneidisca, for instance, has elliptical rather than bicornute spores but is otherwise
identical. In my experience it is just as common in our area as L. cristata. There are many
other similar species too numerous to mention. They include: L. decorata, with reddish to
pink scales; L. roseilivida, with a purplish or purplish-pink, fibrillose cap; andL. “tomen-
todisca,” with pale to dark reddish-brown scales. In all three of these species the center
of the cap is minutely hairy (tomentose), a character best seen with a hand lens.

Lepiota sequoiarum (Boring Lepiota)

CAP 1.5-4 cm broad, oval becoming convex or finally plane; surface dry, smooth, without
distinct scales, yellowish to tan at the center, pallid toward the margin. Flesh thin, white,
not bruising. GILLS white, free, close. STALK 2-7 cm long, 2-5 mm thick, equal or thicker
below, slender, white, smooth. VEIL membranous, white, forming a superior ring on stalk
which often collapses in age. SPORE PRINT white; spores 7-9 * 3.5-4 microns, elliptical,
smooth, dextrinoid.
HABITAT: Scattered or in small groups in woods and planted areas, infrequent. I have
found it several times in the fall and winter under redwood. It is one of several small
Lepiotas apparently endemic to the west coast.
EDIBILITY: Unknown—do not experiment!
COMMENTS: The smooth cap, small size, membranous ring, free white gills, and white
spores are the decisive features of this lackluster Lepiota. The less said about it the better.
A somewhat similar unknown species with a grayer cap when young occurs on wood chips.

Lepiota seminuda (Lilliputian Lepiota)

CAP 1-2 (4) cm broad, conical or convex becoming plane; surface dry, white or tinged
pinkish (especially at center), smooth or minutely powdery-mealy; margin often with veil
fragments. Flesh very thin, white. GILLS free, white to pale pinkish, close. STALK 2-5
cm long, 1.5-3 mm thick, equal or slightly thicker below, white to dingy pinkish or tinged
cinnamon toward base, thin and fragile, smooth or minutely mealy like the cap. VEIL
evanescent, leaving remnants on the cap margin, but not a distinct ring on stalk. SPORE
PRINT white; spores 3-5 * 2-3 microns, elliptical, smooth.
HABIT AT: S olitary or scattered in humus under hardwoods or conifers; widespread. Oc¬
casional in our area in the fall and winter (e.g., under redwood) but easily overlooked.
EDIBILITY: Unknown. It is hardly worth troubling with such a trifle.
COMMENTS: Also known as L. sistrata, this is our smallest and most delicate Lepiota.
Its lilliputian dimensions might lead to confusion with Mycena or Collybia, but it has free
gills and a veil when young. It is smaller thanL. sequoiarum, and the veil does not normally
form an annulus (ring) on the stalk (see photo at bottom of p. 306).

Lepiota castanea (Petite Parasol)

CAP 1-3 (4) cm broad, convex-umbonate to nearly plane; surface dry, covered with rusty-
ochraceous to chestnut-brown to cinnamon-brown scales on a pallid to yellow-brown or
ochraceous background (scales often densest at center). Flesh very thin, yellowish-buff;
odor sometimes faintly sweet. GILLS close, usually free, white to buff or rusty-stained.
STALK 3-8 cm long, 1-3 mm thick, very fragile and slender, equal or enlarged slightly at
base; smooth above the veil, covered with rusty-ochraceous to dark chestnut-brown scales
(like those on cap) below, often staining orange or yellow-orange when handled or in age.
 Two deadly poisonous Lepiotas. Left: Lepiota castanea. Note scales on lower stalk. Right: Lepiota
josserandii is easily confused with L. cristata (p. 306), but usually lacks an annulus (ring) on stalk.

VEIL fibrillose, evanescent, not forming a distinct ring on stalk, but sometimes leaving
remnants on cap margin. SPORE PRINT white; spores 9-13 x 3.5-5 microns, bullet¬
shaped, smooth, dextrinoid.
HABITAT: Solitary to widely scattered or in small groups in rich humus in woods, espe¬
cially under conifers. It has a wide distribution and is not uncommon in our area in the fall
and winter, but never seems to fruit in large numbers and is likely to be overlooked.
EDIBILITY: POISONOUS! Like L. josserandii, it contains deadly amanita-toxins.
COMMENTS: The rusty-orange to chestnut-colored scales on cap and stem, free or
nearly free gills, and lack of a distinct annulus (ring) typify this petite Lepiota. It
might be mistaken for a Cystoderma, but the gills are usually free and the spores are
dextrinoid. It is quite fragile and difficult to transport home in one piece.

Lepiota josserandii (Deadly Parasol)

CAP 2-5 (7) cm broad, convex to plane or sometimes umbonate; surface dry, with cinna¬
mon-brown to pinkish- or reddish-brown scales, the center darker and the background
whitish to ochraceous, but reddish or rosy tints often developing in age or upon drying;
margin often fringed with veil remnants. Flesh thin, pallid; odor faintly sweetish or musty,
especially if several are left in a closed container. GILLS free or adnexed, close, white to
creamy-yellow, not bruising. STALK 3-7 cm long, 0.3-1 cm thick, equal or enlarged
downward, pallid at apex, fibrillose-scaly to near smooth and cap-colored (or slightly
pinker) below. VEIL fibrillose, evanescent, not forming a distinct ring on stalk, but
sometimes leaving a hairy zone. SPORE PRINT whitish; spores 6-8 x 2.5-4.5 microns,
elliptical, smooth, dextrinoid.
HABITAT: Solitary, scattered, or in groups in cultivated ground, under bushes and trees
(including cypress), on lawns, etc.; widely distributed and not uncommon in California.
I have found it several times in our area in the summer and fall.
EDIBILITY: POISONOUS! Deadly amanita-toxins have been isolated in this species and
several relatives. Fortunately, they are unlikely to be eaten because of their small size and
infrequent occurrence. However, L. josserandii has been implicated in at least one fatality
(in Albany, New York).
308
LEPIOTA 309

COMMENTS: This undistinguished little Lepiota{see photo at top of p. 308) is described

here because it poses a threat to those who sample mushrooms wantonly, or think that only
the Amanitas are deadly poisonous. It belongs to a large group of small, poorly-known,
difficult-to-identify Lepiotas with no annulus (ring) or only a slight one, and a more or less
fibrillose-scaly or cottony stem. Until they are better known, none should be eaten. Several
occur in our area under cypress, including a beautiful pinkish-hued species that might be
L. subincarnata, another poisonous species. Other deadly poisonous species include:
L. helveola, very similar to L. josserandii, but with larger spores (7-10 microns long) and
a slightly more membranous veil; and L. felina, with blackish to dark brown scales on
both the cap and stalk. See also L. castanea. Other species:/,, cortinariusis a medium-sized
species (cap 3-10 cm broad; stalk 0.6-2 cm thick) that grows under northern conifers such
as spruce. Its cap cuticle soon breaks up into numerous concentrically-arranged small
brown to reddish-brown scales, its veil is evanescent, and its gills are remote from the stalk
or attached to a collar. It does not age or stain reddish or pinkish; its edibility is unknown.

Lepiota clypeolaria (Shaggy-Stalked Parasol)

CAP 2-8 cm broad, oval or bell-shaped becoming convex to nearly plane with a low, broad
umbo; surface dry, soon breaking up into yellow-brown or brown scales except for the
smooth, darker center; often yellower toward margin, which is soft and ragged from
cottony veil remnants. Flesh white, not staining appreciably; odor sometimes pungent.
GILLS free, close, white or creamy. STALK 4-12 (18) cm long, 3-7 (10) mm thick, usually
slender and about equal, fragile; sheathed with soft, shaggy or cottony scales below the
veil, usually yellow or colored like cap. VEIL cottony, leaving remnants on cap margin or a
slight cottony-fibrillose ring on stalk. SPORE PRINT white; spores 12-20 * 4-6 microns,
fusiform (elongated), smooth, dextrinoid.
HABITAT: Solitary, scattered, or in small groups in woods, widely distributed. In our area
it favors conifers such as Douglas-fir, and is fairly common in the fall and winter. A related
species or variant (see comments) occurs under oak in the winter and spring.

Lepiota clypeolaria, mature specimens. The shaggy stalk plus the smooth dark cap center typify this
beautiful woodland mushroom. The cap is oval or convex before it expands.
Lepiota clypeolaria (form “nabiscodisca”). This form is not as shaggy as the one shown on the previous
page, and does not show as much yellow. In our area it favors oak.

EDIBILITY: Said to be poisonous—all slender woodland Lepiotas are best avoided.

COMMENTS: The ragged or shaggy appearance of the cap margin and stalk, smooth
“eye” at the center of the cap, absence of a distinct annulus (ring), free whitish gills, and long
narrow spores are diagnostic. In prime condition it is one of our most beautiful and
exquisitely adorned mushrooms. In age, however, it assumes a rather decrepit appearance
—the stalk is weak and collapses easily. In North America L. clypeolaria is probably a
“collective” species—that is, there are several varieties which may be distinct species. In
addition to the form described above, we have in our area an oak-loving variety with
fusiform spores, a more evenly colored and less ragged cap, and a whitish, only slightly
shaggy stalk. I call it var. “nabiscodisca” because of its cookie-colored cap cuticle, but it
may prove to be the “true” L. clypeolaria of Europe (and the L. clypeolaria described
above may beL. ventriosospora—but such issues are best left to licensed lepiotologists).
Other species: L. clypeolarioides of the Pacific Northwest lacks yellow tones, is not as
shaggy, and has elliptical spores (6-9 microns long).

AGARICACEAE (Agaricus)
Mostly medium-sized to large, terrestrial, saprophytic mushrooms. CAP smooth or scaly, typically
neither brightly colored nor viscid. Flesh usually white. GILLS close, free or nearly free (at least at
maturity), pallid or pinkish when young, chocolate-brown to blackish-brown in age. STALK
central, fleshy, cleanly separable from cap. VEIL present, membranous or cottony, usually forming
an annulus (ring) on stalk. VOLVA typically absent (except mA. bilorquis and relatives). SPORE
PRINT chocolate-brown. Spores smooth, mostly elliptical or almond-shaped.

FROM both an economic and gastronomic standpoint this is unquestionably the most
important group of gilled mushrooms, for it includes the familiar cultivated or “grocery
store” mushroom as well as a large number of other delectable collectables. As defined here,
the Agaricaceae include only one common genus, Agaricus (sometimes listed in older
books as Psalliota). It is a remarkably clearcut, “natural” genus, and as such is a cinch to
recognize: the stalk is typically furnished with an annulus (ring) but lacks a volva, the gills
are pinkish or pallid when young but become chocolate-brown and are free at maturity, and
the spore print is always chocolate-brown (the color of dark chocolate, not milk chocolate).
Stropharia is somewhat similar to Agaricus, but usually has a viscid cap and attached gills
that are never pink, while Amanita has white spores and white or pallid gills plus (usually) a
volva. Lepiota and Chlorophyllum are very similar in general appearance, but have white
and greenish spores respectively. (The Lepiotas and several miscellaneous smaller genera
are included in the Agaricaceae by many mycologists.)

310
These photographs show the key fieldmarks of any Agaricus: presence of a veil that usually forms
an annulus on the stalk plus gills that are free and chocolate-brown at maturity. Left: Close-up of a
mature Agaricus augustus (p. 337). Right: Mature specimens of A. arorae{an unusual species—see
p. 325). Note how cap separates easily from stalk.

Because so many of its species are edible, one often hears Agaricus characterized as a
“safe” genus. This is hardly the case, however.True, no Agaricus is deadly poisonous, but
some species produce mild to severe vomiting and diarrhea in most people, and most of the
edible species (including the cultivated mushroom, A. bisporus) produce mild to severe
vomiting and diarrhea in some people. In fact, Agaricus species are the most frequent cause
of mushroom poisoning in our area, if not in California—not a very good track record for a
“safe” genus! It therefore behooves you Agaricus-eaters to sample each and every species
cautiously to determine your reaction to it, and to be absolutely sure of your identification!
Unfortunately, being “absolutely sure of your identification” is easier said
than done, because Agaricus species are perplexingly polymorphic (variable in size, shape,
color, etc.), and as a result, devilishly difficult to differentiate from each other. In view of the
frequency with which Agaricus species are eaten, and the fact that there is no single “Simple
Simon” rule for distinguishing the edible ones from the poisonous, it seems prudent to
detail some of the more important characters used in separating them:
1) Staining reactions should be noted when fresh on the surfaces of the cap (near the
margin) and stalk, the flesh in both the cap and stalk, and the flesh in the extreme base of the
stalk. Some species do not stain at all; others are rufescent, (i.e., they stain red to orange or
vinaceous when cut) or lutescent (i.e., they stain yellow to yellow-orange when bruised or
rubbed repeatedly); still others are latently lutescent (i.e., they stain yellow only when a
drop of 10% potassium hydroxide (KOH) or sodium hydroxide (NaOH) is applied to the
surface of the cap near the margin).* Actually, potassium and sodium hydroxide drama¬
tize the yellow-staining of all the naturally lutescent species, but in our area are especially
useful for distinguishing the edible, non-staining A. campestris from the inedible or
poisonous, latently lutescent A. californicus.
2) Odor should be noted by gently crushing the cap tissue as well as the flesh in the very
base of the stalk. Three odors are especially prevalent in Agaricus: mild (or “fungal”) to
faintly fruity, as in the cultivated mushroom, A. bisporus; sweet (like anise or almond
extract) as in A. augustus; and phenolic (an unpleasant chemical odor reminiscent of
phenol, carbolic acid, creosote, library paste, ink, tar, or bleach), as in A. xanthodermus.
However, these odors, when present, are not always obvious and it sometimes takes an
experienced nose to detect them. It is interesting to note that there is often a correlation
between the degree of lutescence and strength of odor—as a rule, the more quickly and
brightly an Agaricus stains yellow, the stronger it will smell!
*Many household cleaning agents contain potassium or sodium hydroxide and can be substituted successfully.
Drano (one teaspoon per % cup water) and Lysol both work, but don’t eat the tissue tested with these chemicals!
Veil detail in Agaricus. Left to right: A. campestris, with a thin cottony veil; A. californicus, with a
thicker membranous veil and inrolled, lobed cap margin; and A. arvensis, whose membranous veil
shows a cogwheel pattern of patches on its underside. (Ralph Buchsbaum)

3) Veil characteristics: The veil in Agaricus is actualy composed of two layers of tissue.
In some species only one layer is clearly visible, while in others the lower layer (universal
veil) can be seen as distinct patches of tissue on the underside of the upper layer (partial
veil), as shown in the photograph above. The type of annulus(ring) formed by the ruptured
veil is also significant: whether skirtlike (pendant), sheathlike(peronate), or intermediate
between the two (see illustrations on this page).
4) Spore size is extremely useful in delimiting species, as are reactions to other chemicals
besides potassium and sodium hydroxide. However, these features are not stressed here
since most people will be unable to ascertain them. The best spores for measuring purposes,
incidentally, are those that are deposited naturally (in the wild) on the annulus or stem.

Different types of rings in Agaricus. Left to right: A. silvicola, A. praeclaresquamosus, A. bitorquis.

What makes identification of Agaricus species so difficult is that all of the above charac¬
ters with the exception of spore size are easily influenced or altered by the environment, as
are grosser features such as the size, shape, color, and degree of scaliness of the fruiting
body. Specimens growing intheopen,forinstance,can be differently colored orsquatteror
more scaly than specimens of the same species growing in deep shade. Dry or old specimens
are apt to manifest their staining reactions and odors much more slowly or subtlely than
young, fresh individuals, and soggy specimens may not display their normal odor and
staining reactions at all! It is obvious, then, that habitat and environmental conditions
are among the first things to be taken into account when you are attempting to identify an
unfamiliar Agaricus.

312
AGARICUS 313

One practical way for beginners to overcome some of the vagaries and difficulties dis¬
cussed is to approach the genus Agaricus at the level of section (subgroup) rather than
species. The overwhelming majority of North American species will fit into one of seven
groups or “sections.” A few, such as A. subrutilescens and A. arorae, do not fit con¬
veniently into any of them, and may well represent “bridges” between them. The seven
sections are:

Section Agaricus: Not staining; annulus thin and slight or even absent (when present
usually intermediate); KOH-negative; edible.
Section Hortenses: Not staining or rufescent; annulus skirtlike or intermediate; stature
usually rather squat; KOH-negative; edible.
Section Bitorques: Not staining or rufescent; annulus sheathlike; flesh hard; stalk
solid; KOH-negative; edible or sometimes too tough to eat.
Section Sanguinolenti: Rufescent; annulus skirtlike; stature usually erect; KOH-
negative; edible.
Section Xanthodermati: Latently or strongly but fleetingly lutescent (i.e., staining
yellow, then eventually discoloring brownish or vinaceous); annulus skirtlike or
intermediate; odor phenolic (sometimes faint!); KOH-positive; poisonous!
Section Arvenses: Latently or strongly but persistently lutescent; annulus skirtlike;
odor usually sweet; veil typically with two distinct layers; KOH-positive; edible.
Section Minores: Very much like section Arvenses and sometimes grouped with it, but
with a small, usually slender and fragile fruiting body, and veil often with only one
distinct layer.

Learning to recognize these “sections” has several advantages. The most obvious is
that it hones critical skills while dealing with only seven entities instead of dozens.
Moreover, you learn to group species according to their chemistry, which is precisely what
edibility is all about! For instance, it appears that the phenol-smelling species (section
Xanthodermati) poison a majority of people who eat them. The phenol odor is not always
evident in the field, but usually becomes quite pronounced if the mushrooms are cooked.
(The taste of these species is rather astringent or metallic, yet there are people who eat them
not only with impunity, but with gusto!) It is also interesting to note that people who have
an allergy to an edible species are likely to have allergies to other members of the same
section, but not necessarily to species in other sections . For instance, someone allergic to
A. augustus is likely to be adversely affected by A. arvensis also. Similarly, someone who
can eat the cultivated mushroom (A. bisporus) will probably not have trouble with other
species in the section Hortenses. Thus it clearly pays for prospective Agaricus-eaters to
learn something of the chemistry and interrelationships of the common species, in addi¬
tion to their critical fieldmarks.
The best known—and perhaps the most mediocre—of the edible Agaricus species is
undoubtedly the cultivated mushroom, A. bisporus. The meadow mushroom or
“champignon,” A. campestris, is also very popular, but the best species are somewhat lesser
known—A. augustus, A. subrufescens, A. arvensis, A lilaceps, and A bernardiiare choice
mushrooms if there are any, and A. bitorquis has few peers. Agaricus species are also
notable for their beauty. Some rival the Amanitas for stateliness and elegance, while
others are as plump and meaty as boletes.
Agaricus species are saprophytic and are among the most conspicuous urban and
suburban mushrooms. Not only is the cultivated or “button” mushroom, A. bisporus, to be
found in practically every grocery store and produce stand, but its untamed (and in some
cases unnamed) cousins fruit prolifically on lawns (where they are often kicked over like
other “toadstools”), in cemeteries, under hedges and shrubbery, along roads and sidewalks,
in gardens, and on compost piles. Many species are rural as well, fruiting by the bushel in
pastures and fields, and some also favor the woods, but comprise a much smaller
314 AGARICACEAE

percentage of the fleshy fungi found there. Few, if any, are mycorrhizal. It is interesting to
note that in our area cypresses—particularly old ones—harbor a remarkable wealth of
species, including both cosmopolitan and rare or endemic ones.
Agaricus species fruit whenever conditions are favorable—that is, moist and mild. In our
area the greatest variety can be found shortly after the first fall rains, but another crop
appears in the spring, and if it is mild enough, in the winter. Some species also fruit on
watered lawns in the summer, and A. augustus is a common summertime mushroom in
the coastal fog belt.
Agaricus is a large genus centered in the tropics and subtropics, hence one would expect
to find more species in the southern United States than in the north. About 200 species
are estimated for North America, but no all-encompassing critical study has been made.
In California the total number of known species is about 60, most of which occur in our
area. Many of the 24 species described here have wide distributions, but some are presently
known only from California.* Readers outside “our area” will undoubtedly enounter
several species they can’t identify, but with a good nose and a keen eye they should be able to
assign most of them to one of the seven “sections” already described.

Key to the Agaricaceae (Agaricus)

Note: As already pointed out, many of the characters used in this key are subject to
environmental influence—particularly the intensity (or even presence) of staining reac¬
tions and odor. It is therefore imperative to have several specimens of each species in
hand, preferably collected in conditions neither exceptionally wet nor unusually dry.

1. Very base of stalk giving off an unpleasant odor (like phenol or library paste) when crushed;
base sometimes also staining yellow when cut or crushed . 2
1. Not as above; odor not phenolic; base of stalk typically not staining yellow (but may stain orange
or yellow-orange), or if staining yellow then odor sweet . 7
2. Extreme base of stalk staining bright yellow when cut or nicked; other surfaces often staining
yellow also . 3
2. Base of stalk not staining bright yellow (but may stain faintly yellow) .4
3. Cap with inky-gray to grayish-brown to brown or dark brown fibrils or fibrillose scales; surface
of cap not bruising yellow or only sometimes bruising yellow .6
3. Cap white or discoloring grayish to buff or tan; surface usually bruising bright yellow when
rubbed repeatedly, especially near margin .A. xanthodermus, p. 329
4. Cap with pale pinkish-brown to fawn-colored, tan, or reddish-brown fibrils (or at times nearly
white); annulus (ring) on stalk thick and feltlike; stalk 1-3 cm thick at apex, the base enlarged;
cap 6-15 cm or more broad; growing in woods or under trees .A. hondensis, p. 326
4. Not with above features.5
5. Cap 3-9 cm broad, entirely white or whitish with a brown center, or sometimes brownish
throughout; stalk 0.5-1 (1.5) cm thick; found in many habitats on west coast, but especially
common in urban and suburban areas, in grass, under cypress and oak, etc.52
5. Not as above; if found in West then cap (4) 5-25 cm broad, with inky gray to grayish-brown,
brown, or dark brown fibrils or fibrillose scales (never entirely white) and stalk 1 -3 cm thick,
and usually growing in woods or along roads and paths through the woods .6
6. Found in eastern North America; cap typically finely fibrillose.
.A. placomyces& A. pocillator(sQe A. praeclaresquamosus, p. 329)
6. Widespread but most common in West; cap fibrillose to scaly A. praeclaresquamosus, p. 329
7. Some part of fruiting body (especially cap surface) staining yellow when bruised and/or
the flesh smelling sweet when crushed (like almond extract or anise); cap surface typically
yellowing in KOH . 8
7. Not as above (odor rarely slightly almondy, but if so then flesh reddening when cut) .25

*Several of the species depicted or mentioned in this chapter have been named and described by Rick Kerrigan in
an article which, at the time of this writing, is about to be published. The provisional names he has given them are
used here with his permission. They are: A.fuscovelatus, A. arorae, A. sequoiae, A. vinaceovirens, A. blandianus,
A. smithii, A. summensis, A. perobscurus, and A. rubronanus. Another provisional name, A. pinyonensis,
is used here with the permission of Bill Isaacs and Chuck Barrows.
AGARICUS 315

8. Cap white or whitish when young and fresh (but may discolor yellow to buff or amber in age) 9
8. Cap not white (i.e., with distinctly colored cuticle, fibrils, or scales, at least at center) .... 16
9. Growing in grass (lawns, pastures, etc.) or in pinyon-juniper forests . 10
9. Not typically growing in above habitats . 12
10. Cap small (2-5 cm broad), often with a brownish to buff-tinged center; stalk less than 1 cm thick;
spores 5.5 microns long or less .A. comtulus(see A. micromegathus, p. 340)
10. Not as above; usually larger and spores larger . 11
11. Stalk stuffed or sometimes hollow; fruiting body medium-sized, or if large than not particularly
squat; cap surface usually smooth (or with a few fissures); widely distributed, but favoring
lawns over pastures in California; spores 7-8.5 microns long .A. arvensis, p. 332
11. Not as above; either growing in pinyon-juniper forests or if growing in grass, then stalk usually
solid, fruiting body often large (cap 7-50 cm broad), the cap smooth to conspicuously scaly or
warty, stature often robust or squat; especially common in pastures and prairies but also on
lawns; spores either smaller or larger than above .41
12. Fruiting body often robust, quickly staining amber when bruised (especially the cap surface)
and often aging amber overall; cap often fibrillose; odor strongly almondy or anise-like; im¬
mature gills often staining yellow; known only from the West.A. albolutescens, p. 335
12. Not as above ... 13
13. Fruiting body robust (stalk 2 cm thick or more); cap typically with slightly colored (yellowish)
fibrils; rare .A. summensis& A. augustus(white form) (see A. augustus, p. 337)
13. Not as above; common . 14
14. Cap surface typically staining distinctly yellow when rubbed repeatedly, at least at the margin;
common and widespread .A. silvicolagroup, p. 334
14. Not as above; yellow-staining weak or erratic; found in eastern North America or in western
mountains . 15
15. Cap white, 4-8 (15) cm broad; odor pungent or faintly sweet; spores only 4-5 microns long; fairly
common in woods of eastern North America.A. cretacellus
15. Cap white or silvery; gills remaining pink for a long time; found in western mountains .
. A. chionodermus(see A. silvicola group, p. 334)
16. Fruiting body fairly small; cap 1-5 (7) cm broad; stalk usually less than 1 cm thick . 17
16. Fruiting body medium-sized to very large; not as above. 19
17. Growing in grass; odor usually anise-like .A. micromegathus & others, p. 340
17. Growing in woods or under trees; odor mild or aniselike . 18
18. Odor distinctly anise-like; cap fibrils brown to reddish-brown .
.A. semotus(see A. micromegathus, p. 340)
18. Odor faint or absent; cap fibrils pink, purple, purple-gray, or vinaceous when fresh .
.A. diminutivusgroup, p. 340
19. Growing in pastures or grass; fruiting body robust, often squat; cap often with large warts or
scales, but sometimes smooth, whitish becoming yellowish or pale brownish .
. A. crocodilinus (see A. osecanus group, p. 333)
19. Not growing in grass, or if growing in grass then not as above . 20
20. Growing in compost or rich soil; cap with pallid to pale brown to faintly grayish, pale pinkish-
brown, fawn-colored (or sometimes brown or tawny) fibrils, sometimes slightly ruddy or
yellowish in age; stalk not scaly or shaggy(or only obscurely so) below the veil; stalk often with
a swollen base; odor strongly sweet; spores 5.5-7 microns long .A. subrufescens, p. 336
20. Not as above; often but not always growing in woods . 21
21. Cap with pink or purple tints; not common .A. lilaceps, p. 323
21. Not as above . 22
22. Stalk typically rather slender(l -1.5 (2) cm thick at apex), but usually with a bulb at base; fibrils on
cap ochraceous-orange to tawny; found in the coastal forests of northern California and the
Pacific Northwest, especially under Sitka spruce .A. smithii(see A. augustus, p. 337)
22. Not as above . 23
23. Fibrils on cap yellow when young, ochraceous or tawny in age; rare .
. A. summensis{see A. augustus, p. 337)
23. Not as above; fibrils brown to gray to tawny, but not yellow . 24
316 AGARICACEAE

24. Fibrils on cap distinctly gray to grayish-brown to nearly black (at center) when young, but often
paler in age; scales on stalk below the veil often scanty; gills usually passing through a pinkish
phase as they mature; known only from California .A. perobscurus, p. 339
24. Not as above; fibrils on cap brown to tawny; stalk usually conspicuously shaggy or scaly below
the veil, at least when young; gills rarely pink; widely distributed .A. augustus, p. 337
25. Flesh normally staining red to orange or vinaceous when cut or rubbed repeatedly .26
25. Not as above (but cap and flesh may have reddish stains in old age or wet weather) .38
26. Flesh in base of stalk staining orange to yellow-orange when cut; flesh elsewhere reddening at
least somewhat; cap with broad, flattened, chocolate-brown scales, often depressed centrally
at maturity; growing in woods, very rare (reported from the Pacific Northwest) .. A. lanipes
26. Not as above . 27
27. Unbroken veil (i.e., underside of broken veil) soon brown to grayish-brown, purple-brown, or
chocolate-brown to nearly black . 28
27. Not as above . 30
28. Stalk averaging 2.5-4 cm thick at apex, often bulbous A. pattersonae (see A. lilaceps, p. 323)
28. Stalk usually more slender, or if thick then without a bulb at base.29
29. U nderside of unbroken veil soon purple-gray to brown or dark brown; flesh staining only slightly
(often orangish) when cut; stalk typically less than 2 cm thick .A. fuscovelatus, p. 324
29. Not as above . 30
30. Veil forming a sheathlike annulus(ring) on stalk; cap whitish to buff or sometimesdingy brown¬
ish, sometimes cracked into scales; stalk solid; fruiting body very firm or hard; growing in
disturbed soil, mud flats, on lawns near ocean, etc.; coastal in distribution A. bernardii, p. 322
30. Not as above; veil forming a skirtlike to intermediate annulus . 31
31. Growing in compost or manured soil .A. bisporus, p. 319
31. Not as above . 32
32. Fruiting body large and dense (cap 8-25 cm broad when mature, stalk2-7 cm thick; cap brown or
sometimes with pinkish, purplish, and/ or ochre or yellow-orange tones; veil often yellow-
tinged before breaking, forming a skirtlike annulus (ring) on stalk; found mainly under cypress
and other planted trees, but not in manure .A. lilaceps, p. 323
32. Not as above . 33
33. Margin of cap inrolled when young; stature of fruiting body usually rather stout or squat;
annulus (ring) on stalk intermediate or skirtlike . 54
33. Not as above; annulus usually skirtlike. 34
34. Flesh staining distinctly red . 35
34. Flesh staining dingy vinaceous or vinaeous-brown or dingy reddish (usually weakly) .... 38
35. Cap white or whitish, sometimes with faintly grayish or brownish fibrils .
.A. benesi(see A. fuscofibrillosus, p. 325)
35. Not as above; cap darker . 36
36. Common under mountain conifers (especially spruce and fir) in southern Rocky Mountains and
Southwest .A. amicosus (see A. arorae, p.325)
36. Not as above .37
37. Stalk usually less than 1 cm thick; cap 3-7 (10) cm broad; cap surface yellowing in KOH; known
only from California, usually growing in mixed woods and under oak .... A. arorae, p. 325
37. Not as above; stalk usually thicker and! or growing under cypress; cap surface not yellowing
in KOH; widespread .A. fuscofibrillosus & others, p. 325
38. Veil forming a sheathlike or even volva-like annulus on stalk or a large collarlike band (with
both upper and lower edges free from stalk); found in hard-packed or disturbed soil or some¬
times in grass or under cypress; texture very firm; stalk solid . . A. bitorquis & others, p. 321
38. Not as above; veil not sheathlike or bandlike and/ or stalk with a central hollow in age ... 39
39. Cap white or whitish when fresh (but may discolor tan, yellowish, or buff in age) .40
39. Cap colored at least at the center, or with colored fibrils or scales.42
40. Cap medium-sized to very large (10-50 cm broad when mature); either growing in grass or in
pinyon-juniper forests.41
40. Not as above; smaller and/or habitat different .42
AGARICUS 317

41. Found with pinyon and juniper in the Southwest . A. pinyonensis, p. 331
41. Found in grassy places A. osecanusgroup & A. crocodilinus(see A. osecanus group, p. 333)
42. Cap 1 -4 cm broad when expanded; stalk 2-6 mm thick; fruiting body fragile.43
42. Not as above; typically larger (cap 3 cm broad or more; stalk 5 mm thick or more) .44
43. Cap grayish to grayish-brown, granular or powdery; margin usually hung with veil remnants;
gills deep pink to blood-red, then darker; spore print reddish or tinged greenish when moist,
drying darker brown; growing in greenhouses, leaf litter, rich soil, bogs, etc.; widely distributed
but not common .Melanophyllum echinatum
43. Not as above; cap with reddish-brown to pinkish or purplish fibrils, not granular or mealy;
growing in woods .A. diminutivusgroup, p. 340
44. Cap silvery-white or white; fruiting body often rather erect and slender; growing in mountain
forests of the West . A. chionodermus(see A. silvicola group, p. 334)
44. Not as above .45
45. Cap covered with dark brown to purple-brown or wine-colored fibrils or fibrillose scales; stalk
shaggy or sheathed by cottony white scales below the veil, at least when young; common in
woods .A. subrutilescens, p. 326
45. Not as above .46
46. Growing in the open (usually in grass, occasionally in hard-packed soil); veil typically thin and
somewhat cottony, forming only a slight annulus (ring) on stalk or disappearing (annulus
usually intermediate, not skirtlike); gills usually pink or brownish in button stage; cap surface
not yellowing in KOH .47
46. Not as above; habitat different and/or veil typically membranous and forming a distinct
intermediate to skirtlike annulus; gills pinkish, brown, or white in button stage .49
47. Cap white or sometimes with pale brown to grayish fibrils or scattered scales; very common and
widespread .A. campestris& others, p. 318
47. Cap covered with brown to grayish-brown or reddish-brown fibrils, even when young ... 48
48. Cap often ruddy or reddish in age or after handling; growing in lawns or hard-packed soil along
roads; not common .A. rutilescens(see A. cupreobrunneus, p. 319)
48. Not as above; growing in lawns or pastures; common . . A. cupreobrunneus & others, p. 319
49. Stature erect (stalk more than 5 cm long); cap white or pallid (pale buff) .50
49. Not as above; cap darker and/ or stature rather robust and stocky .52
50. Stalk 8-30 cm long; found in redwood and mixed forests of coastal northern California, usually
near rivers or in bottomlands .A. sequoiae(see A. pinyonensis, p. 331)
50. Not as above; stalk generally up to 9 cm long; habitat usually different . 51
51. Veil usually disappearing or leaving scaly zones on mid- or lower stalk rather than forming a
distinct annulus (ring); found under trees or on coastal bluffs; rare .A. altipes
51. Not as above; common . 52
52. Cap white or pallid; veil ample, usually with a cogwheel pattern of patches on underside; stalk
rather stout (3-b cm long); growing in pastures, sometimes in rings; known only from the central
California coast; not common .A. sp. (unidentified)(see A. campestris, p. 318)
52. Not as above; very common . 53
53. Cap covered with purplish to purple-gray or purple-brown fibrils, the margin usually white;
found in woods and under trees in southeastern North America .A. rhoadsii
53. Not as above . 54
54. Gills pinkish to brownish in button stage; fruiting body usually stout, stalk usually 1 -4 cm thick;
found in manure, compost, rich soil, hard-packed ground, and under cypress (or rarely in the
woods); odor never phenolic; cap surface not yellowing in KOH or rarely yellowing slightly
. 55
54. Gills pallid or whitish in button stage (but often pink after veil breaks); stalk0.5-1 (1.5)cm thick;
found almost anywhere (including lawns, gardens, woods, under cypress), but only rarely in
manure or compost piles; odor sometimes phenolic; cap surface yellowing in KOH .
.A. californicus, p. 327
55. Growing in hard-packed ground, often in towns or cities (often developing underground, then
pushing through); veil thick, often forming a double ring; cap surface usually breaking up in
age to form large brown to reddish-brown scales . A. vaporarius
55. Not as above; found in compost, manure, under cypress, or occasionally in other habitats; cap
white or with brown fibrils or fibrillose scales; common. A. bisporus& others, p. 319
318 AGARICACEAE

Agaricus campestris (Meadow Mushroom) Color Plate 71

CAP 4-11(15) cm broad, convex or dome-shaped for a long time, then often becoming
plane; surface dry, smooth or silky-fibrillose, pure white, or sometimes with a few grayish
to brown or cinnamon-buff fibrils or fibrillose scales; margin extending beyond the gills,
often hung with veil remnants. Flesh thick, white, not staining when bruised but sometimes
discoloring brownish or reddish in age or wet weather (especially just above the gills); odor
mild. GILLS close, free at maturity, pale pink in button stage, then bright pink, becoming
purple-brown to chocolate-brown and finally blackish-brown. ST ALK 2-6 (10) cm long, 1 -
2.5 cm thick, usually with a tapered base; firm, white, smooth above the veil, often with a
few fibrils below; stuffed or hollow (but see comments). VEIL thin, somewhat cottony,
white, forming a thin ring on stalk or leaving remnants on cap margin or disappearing
entirely; ring rarely well-formed, intermediate (sometimes flaring) or rarely skirtlike,
median to superior. SPORE PRINT chocolate-brown; spores 6.5-8.5 x 4-5.5 microns,
elliptical, smooth. Cap surface not yellowing in KOH. Basidia mostly 4-spored.
HABITAT: As its name implies (campestre-field), this is a grassland species, occurring
throughout the world from sea level to above timberline. It usually grows in groups or
rings in lawns, pastures and meadows, cemeteries, golf courses, baseball fields, etc., but can
also be solitary. It fruits practically year-round in our area, but is most abundant in the fall
and early winter, when stupendous crops are sometimes produced in our pastures. Thou¬
sands of pounds, in fact, go unpicked every year. Though they can often be seen from the
road, fungophobic Americans drive right by them, some undoubtedly on the way to the
store to buy mushrooms!
EDIBILITY: Edible and excellent, both raw and cooked. It is the most widely picked
mushroom in English-speaking countries (where it is also known as the “pink-bottom”),
and is the popular champignon of France. Those who equate it with the cultivated mush¬
room (A. bisporus) do it an injustice. Its flavor is far superior, though the texture is softer.
When the gills are bright pink and the cap pure white, it is as beautiful as it is delicious.
COMMENTS: The silky white cap, bright pink gills when young, poorly defined or evan¬
escent annulus (ring), tapered stem, absence of a volva, stocky stature, chocolate-brown
spores, and growth in grass are the principal fieldmarks of this universal favorite. It is
the type species of Agaricus as well as the entire order Agaricales. The gills are pinkish
even in the button stage, unlike A. californicus. The cap is white or has scattered brownish
scales, but is not brown and fuzzy in youth as in A. cupreobrunneus. The cap remains
dome-shaped for a long time, and its surface does not bruise yellow, though I have
encountered a solid-stemmed variety in New Mexico whose cap sometimes ages faintly
yellowish. This variety may be A. solidipes, an edible southern species with slightly larger
spores than A. campestris (but identical in most other respects, including habitat).
In most regions the meadow mushroom is a perfectly safe mushroom for beginners.
Unfortunately, in California it is frequently confused with the mildly poisonous phenol¬
smelling Agaricus species. Compare it especially carefully with A. calif ornicus (see com¬
ments under that species), which has a more persistent, membranous veil and whitish
gills in the button stage. A small, anonymous Amanita (p. 275) sometimes grows with A.
campestris in pastures and looks very similar from the top. However, its gills are white to
to yellow-orange, never pink or chocolate-brown. The poisonous destroying angels
(Amanita ocreata, A. virosa, etc.) can also be distinguished by their white gills, plus the
presence of a volva (sack) at the base of the stem. Other species: An unidentified, probably
unnamed Agaricus occurs occasionally in our pastures. It resembles A. campestris in
size, shape, and color, but has an ample veil with a cogwheel pattern of patches on its
underside. Its veil, in fact, is reminiscent of A. arvensis, but the fruiting body is smaller
and does not have a sweet odor. It stains yellow in potassium hydroxide (KOH), but does
not normally yellow naturally, and its edibility has not been determined (it might be a
member of the A. xanthodermus-A. calif ornicus group, and therefore unsavory or toxic).
AGARICUS 319

Agaricus cupreobrunneus (Brown Field Mushroom) Color Plate 73

CAP 2 -7 cm broad, convex becoming broadly convex to plane or slightly uplifted; surface
dry, more or less tomentose (hairy) from a layer of small brown to grayish- or reddish-
brown fibrils or fibrillose scales; margin usually extending beyond gills. Flesh white, rather
soft and fragile, not staining when bruised but often discoloring reddish to brownish in old
age or wet weather, especially just above the gills; odor mild. GILLS free at maturity, close,
pale dingy pinkish in button stage, becoming pinkish-brown to purple-brown, then choco¬
late-brown to blackish-brown. ST ALK 2-4 cm long, 0.7-1.2 (2) cm thick, usually equal but
sometimes thicker at either end; white, smooth above the veil, often somewhat scurfy or
scaly below; hollow or stuffed. VEIL thin, white, somewhat cottony, forming a thin ring
on stalk; ring more or less median, intermediate (sometimes flaring), well-formed or
disappearing, sometimes with a slight second ring below it. SPORE PRINT chocolate-
brown; spores 7-9 * 4-6.5 microns, elliptical, smooth. Cap surface not yellowing in KOH.
Basidia mostly 4-spored.
HABITAT: Solitary to scattered orgregarious(ofteninrings)inpastures, lawns,and other
grassy places, but showing a special affinity for poor soil; very common in California and
probably widespread. In our area it fruits prolifically in the fall and early winter, often
growing with and outnumbering A. campestris.
EDIBILITY: Edible. The flavor is comparable to that of A. campestris, but the texture is
much softer and more fragile. Also, the buttons tend to develop underground, making it
difficult to find specimens with pink gills.
COMMENTS: For many years this species has passed as a small, brown, fuzzy form of A.
campestris. It shares with that species certain fundamental characteristics: e.g., a poorly
defined or even absent annulus(ring), stocky stature, and growth in grass. However, it has a
softer texture than A. campestris, and is smaller and brown-capped from the button stage
on. Other species: A. porphyrocephalus, widely distributed, is very similar but has dull
reddish-brown to purple-brown fibrils or scales on the cap, firmer flesh, and smaller
spores. A. rutilescens is also similar but has a ruddier complexion and a tendency—at least
in some forms—to stain reddish. It grows on lawns or in hard-packed soil along roads, but
is not common. A. argenteus is a grass-loving species with a silky or silvery cap with brown
to grayish-brown fibrils, larger spores, and a better-defined annulus. It is common in
the southern United States. All three of the above species are edible.

Agaricus bisporus (Cultivated Mushroom; Button Mushroom)

CAP 3-16 cm broad, convex when young, often plane or even slightly depressed in age;
surface dry, in one form entirely white, but more often with flattened pale brown to brown
fibrils which in age or dry weather often break up into fibrillose scales; margin inrolled
when young, often extending beyond the gills. Flesh thick, very firm, white, usually (but not
always) discoloring somewhat (brown to reddish or pinkish-orange) when cut and rubbed
repeatedly; odor mild or faintly fruity. GILLS free at maturity, close, pinkish or pale brown
when young, purple-brown to chocolate-brown in age, and finally blackish-brown.
STALK 2 -8 cm long, 1-3 (4) cm thick, usually stout, very firm, equal or enlarged at base;
white or turning dingy brownish with age, smooth or slightly cottony-scaly below ring.
VEIL membranous, cottony, white, two-layered, typically forming a delicate, median to
superior ring on stalk which may collapse in age; ring intermediate or sometimes skirtlike,
its upper surface often striate. SPORE PRINT chocolate-brown; spores 5.5-8.5 * 4-6.5
microns, elliptical, smooth. Cap surface not yellowing in KOH. Basidia mostly 2-spored.
HABITAT: Scattered to densely gregarious or in clumps in compost and manure, rich
soil, along paths and in gardens, and also very common under cypress, but only rarely
found in woods or on lawns; very widely distributed. In our area it “fruits” year-round
in markets. The major fruiting in the wild is typically in the fall or winter.
Agaricus bisporus (=A. brunnescens) looks like the cultivated mushroom, which it is. Specimens at
far right are mature; next to them is a button. Cap usually has brown fibrils.

EDIBILITY: Edible—and if popular demand is any indication—choice (over a half bil¬

lion pounds are cultivated annually in the United States!). However, as with any
mass-produced agricultural product, flavor has been sacrificed for appearance, keeping
quality, yield, and disease resistance. Pesticides are used, of course, and the result is, in
Valentina Wasson’s felicitous phrase, “a sickly simulacrum” of what a mushroom should
be. The “wild” form is slightly better—especially the young, hard buttons.
COMMENTS: Also known as A. brunnescens and A. hortensis (the latter name is
applied only to the white form), the cultivated mushroom mimics the meadow mushroom,
A. campestris, but has 2-spored basidia, a well-developed ring, a browner cap, slightly
reddening flesh, and does not normally grow in grass. If you use mushroom compost (from
a mushroom farm) in your garden or put old cultivated mushrooms in your compost pile,
you are likely to get some sooner or later. The largest fruitings I’ve seen have been around
old compost piles and under cypress, where, curiously, it is one of our most common
mushrooms. Care should be taken not to confuse it with A. californicus, which is also
common in gardens and under cypress. The latter is more slender and usually smells slightly
of phenol, at least when cooked. For a more detailed comparison, consult couplet #54 of
the key to Agaricus. A very similar edible species with more rapidly reddening flesh,
skirtlike ring, and 4-spored basidia, A. blandianus, also occurs quite commonly under
cypress; A. subfloccosus is another similar edible species with 4-spored basidia, but it
has a whitish cap with small, cottony scales (squamules) near the margin, and frequently
has a strong odor as well. It occurs under conifers, including cypress, and is widely
distributed, but not common. See also A. spissicaulis (under A. arorae).

The cultivated or “button” mushroom, Agaricus bisporus. Large specimens from a mushroom farm.
Agaricus bitorquis (-A. rodmani), mature specimens. Note prominent, flaring bandlike annulus. The
flesh is exceptionally firm, especially in the stalk.For a view of it in its natural habitat, see color plate.

Agaricus bitorquis (Banded Agaricus; Urban Agaricus) Color Plate 80

CAP 4-18 cm broad or more, broadly convex becoming plane or with a slight central de¬
pression; surface dry, smooth or occasionally cracking into scales; white or whitish but
often dirty; not bruising yellow but sometimes discoloring sordid yellowish to tan in old
age; margin inrolled when young, often extending beyond gills. Flesh thick, very firm,
white, not staining when bruised (but may discolor slightly); odor mild. GILLS close, free
or nearly free; pallid, soon becoming grayish-pink, then deep reddish-brown to chocolate-
brown, finally blackish-brown. STALK 2-10 (18) cm long, 1-3 (4) cm thick, very firm,
solid, equal or slightly thicker below, but often with a narrowed or pointed base; white,
without scales. VEIL membranous, white, thick, forming a prominent, persistent, more or
less median (or even basal) ring on stalk; ring bandlike (upper edge free or flaring and
lower edge free) or sheathlike (like a volva, with only the upper edge free). SPORE PRINT
chocolate-brown; spores 5-7 x 4-5.5 microns, broadly elliptical, smooth. Cap surface not
yellowing in KOH.
HABITAT: Solitary, scattered, or in groups or rows on road shoulders, in hard-packed
soil, along sidewalks, around playgrounds, and in other disturbed areas; often fruiting
underground (looking for them can be like digging for clams!). Widely distributed but
not common on the coast, where it is largely replaced by A. bernardii. I have heard of
gigantic fruiting bodies (10 inch buttons!) of either this species or A. bernardii buried deep
in alluvial soil in riverbeds in the Livermore-Pleasanton area of California. In colder
regions A. bitorquis often appears along roads where salt is sprayed in the winter. In the
Southwest it is often abundant along highways, but is hard to spot because of its subter¬
ranean habit. However, shaggy manes (Coprinus comatus), which are hard notto spot, are
often an indicator of its presence. In our area I have found it in the fall, winter, and spring.

EDIBILITY: Edible, and in my fickle fungal opinion, the best of all Agaricus species. In
Europe it is cultivated commercially because of its immunity to the virus disease that
plagues A. bisporus. It is larger, meatier, and much firmer than either A. bisporus or A.
campestris. One devotee goes so far as to say: “Life is like an Agaricus bitorquis—all good
except for a few gills.” Life may not be so consistently good, but A. bitorquis certainly is!
COMMENTS: The white cap, bandlike or sheathing ring (see photographs), and un¬
changing flesh plus the solid stem, firm texture, and fondness for hardpacked soil distin¬
guish this hardy, handsome fungus from other species of Agaricus. There is no anise or
phenol odor and it does not stain yellow or red. W hen it fruits underground you must search

321
Agaricus bitorquis is common inland in disturbed or hard-packed soil. Note how the large bandlike
ring can mimic a volva (specimens at top and far right). A. bernardii(not illustrated) is a similar species
that is common along the coast. It has reddening flesh and often has a scaly cap.

very carefully for the telltale cracks in the soil that mark its presence (see color plate!).
When the annulus (ring) is sheathlike it sometimes resembles a volva, but the chocolate-
brown spores and gills prevent confusion with Amanita. For years it has been called
Agaricus rodmani; A. edulis is another name for it. A. chlamydopus is a similar species
with a cottony white cap, sheathing veil, stout stature, and larger spores. It is particularly
common in the southern U nited States on lawns and along roads, and is edible. ^4. vinaceo-
virens is a cypress-loving species with a repulsive briny odor and a slightly scaly or scurfy
whitish to light brown cap. It often develops vinaceous tints in age and sometimes stains
greenish as well. Its edibility is unknown.

Agaricus bernardii (Salt-Loving Agaricus)

CAP 5-15 cm or more broad, convex to plane or somewhat depressed centrally; surface
dry, smooth or often breaking up to form scales or warts; white or buff, but may discolor
dingy brownish in age; margin inrolled when young. Flesh thick, very firm, white, staining
reddish-orange to reddish, vinaceous, or reddish-brown when cut (sometimes slowly);
odor mild to pungent or briny. GILLS free at maturity, close, soon grayish-pink or pinkish,
then reddish-brown and finally deep chocolate-brown or blackish. STALK 4-10 (13)
cm long, 2-4 (8) cm thick, solid, very firm, equal or tapered below (rarely thicker
below), smooth, white. VEIL membranous, thick, somewhat rubbery, white, forming a
more or less median, sheathlike (or occasionally bandlike) ring on stalk (the upper edge
often flaring). SPORE PRINT chocolate-brown; spores 6-7.5 * 5-6 microns, broadly
elliptical, smooth. Cap surface not yellowing in KOH.

HABIT AT: Solitary or scattered to densely gregarious in sand, sandy soil, disturbed areas,
on lawns, and in various saline habitats; occurring along the Atlantic and Pacific coasts
and in Europe. In our area it is quite common in thefalland winter, usually appearingafter
the first soaking rains. In Santa Monica, California, I have seen it in the summer on lawns.
It apparently has an even greater tolerance (or liking) for salt than does A. bitorquis,
and like that species, sometimes fruits underground.

EDIBILITY: Edible and choice—almost the equal of A. bitorquis, but a little chewier and
sometimes with a slightly salty or briny taste.
COMMENTS: Also known as A. halophilus and A. maritimus, this is a characteristic
coastal species. It has the general appearance of its close relative, A. bitorquis, (i.e., hard
flesh, sheathing or even bandlike veil, and whitish color), but differs in staining reddish
when cut and is more apt to have a warty or scaly cap. In dry weather, however, the staining
may be slow and/or slight.
Agaricus lilaceps is one of the largest and firmest of its tribe. Note thick stalk, skirtlike annulus (ring),
and tendency of the flesh to darken (redden) when cut. The veil is often tinged yellow before it breaks.

Agaricus lilaceps (Giant Cypress Agaricus)

CAP 7-25 cm broad or more, broadly convex when young, usually plane in age; surface
dry, smooth or covered with flattened brown to pale brown or cinnamon-brown fibrils
which only rarely break up into scales, but sometimes developing strong pink, lilac,
purplish, ochre, or tawny-orange tones (see comments). Flesh very thick and firm, white,
usually staining vinaceous or dark reddish (often slowly) when cut; odor mild to faintly
sweet or fruity. GILLS free at maturity, closev pallid or pale pinkish, soon becoming
reddish- or purplish-brown, then chocolate-brown to blackish-brown. STALK 5-22
cm long, (2) 3-6 (9) cm thick, equal or enlarged below, very firm and thick, white or often
brownish-stained (but sometimes developing pink to purplish hues as on the cap), usually
buried deeply in the humus; smooth or fibrillose (especially below), base often with
yellow or ochre stains. VEIL membranous, fairly thick, underside sometimes white but
very often yellow at first, with white to brownish patches that soon wear away; rupturing
to form a skirtlike or sometimes flaring, superior ring on stalk. SPORE PRINT chocolate-
brown; spores 5-7 * 4-5 microns, broadly elliptical, smooth. Cap surface negative in KOH
or staining very faintly yellow.
HABITAT: Scattered to densely gregarious or clustered on ground under old cypresses
(or occasionally other trees), or where cypresses have been cut down; known only from
California. It is locally common in the fall and winter or whenever it is damp enough; I
have seen large fruitings in Monterey and San Mateo counties.
EDIBILITY: Edible and choice! It is one of the meatiest of all edible mushrooms, but
like A. bitorquis and A. bernardii, it requires thorough cooking to render it tender.
COMMENTS: This massive Agaricus has a rather interesting history. It was originally
described as a lilac- or pinkish-hued species on the basis of specimens collected in Pacific
Grove, California, in the 1930’s. However, careful study by Rick Kerrigan suggests that
it is typically a brown-capped species with reddening flesh, and only develops the brighter
colors in certain localities (e.g., the Monterey Peninsula) and/ or under certain conditions
(perhaps cold weather or refrigeration). In any event, it is easily recognized by its impres¬
sive size, thick hard stem, slowly reddening flesh, and frequently yellow-tinged veil (when
unbroken). It is our heaviest Agaricus—not as broad as A. crocodilinus nor as tall as A.
augustus, but much denser. As in many other species of Agaricus, soggy or weathered
specimens will not necessarily display the “correct” staining reactions, but are usually

323
324 AGARICACEAE

recognizable by their size, stature, and color. The common brown-capped form was provi¬
sionally called A. “luteovelatus” by Rick Kerrigan until it became evident that it was
actually A. lilaceps. Another large edible cypress-lover with reddening to barely reddening
flesh, A. pattersonae, has a scalier, darker brown cap. Its veil is brownish on the underside
rather than yellow, and frequently separates into two distinct rings. It is slightly smaller
than A. lilaceps (stalk 2.5-4 cm thick) and sometimes grows with other trees (e.g., redwood).

Agaricusfuscovelatus (Purple-Veiled Agaricus)

CAP 3 -9 cm broad, obtusely bell-shaped to convex when young, plane in age or with a
slightly uplifted margin; surface dry, covered with brown to cinnamon-brown or dark
reddish-brown fibrils or scales on a white to dingy, gray, or violet-gray background. Flesh
thick, firm, white, staining slightly or slowly yellowish-orange to pinkish-orange or reddish
when cut (especially in stalk); odor mild or faintly fruity. GILLS pallid or grayish-pink
becoming pale brown, then chocolate-brown or darker; close, free or nearly free. ST ALK
4-12 cm long, 0.7-2 (2.5) cm thick, equal or enlarged slightly below, smooth or slightly
scaly near base; white, discoloring somewhat in age; stuffed or hollow. VEIL thin, mem¬
branous; underside at first with a layer of white felty patches, soon becoming purplish-
gray to purple-brown or chocolate-brown to nearly black and scaly; veil often not
breaking free from the cap until the cap has completely expanded, then forming an apical
or superior, fragile, skirtlike, striate ring on stalk. SPORE PRINT chocolate-brown;
spores 6.5-8 x 5-6.5 microns, broadly elliptical, smooth. Cap surface not yellowing in KOH.

HABITAT: Scattered to densely gregarious or clustered under cypress or sometimes other

trees such as cedar and Cryptomeria (an exotic bushy conifer); known only from coastal
California. It is sometimes common in the fall and winter, less so in the spring, but does not
seem to fruit every year. Under a single cypress tree in a cemetery I have seen more than
300 fruiting bodies mingled with dozens of A. bisporus!
EDIBILITY: Edible, but rather tough (especially the stem).
COMMENTS: The most peculiar feature of this most peculiar Agaricus is the chocolate-
brown to purplish-gray or “fuscous” color of the unbroken veil. Its overall appearance
is also distinctive, but hard to describe: the stalk tends to be straight and cylindrical and
the veil, which is close to the top of the stem, remains intact for an unusually long time
before rupturing (it sometimes stretches over the gills like a piece of tissue paper even after
the cap has fully expanded and spores are being produced!). The staining reaction is best
seen by cutting the stalk in half crosswise. A. lilaceps and A. pattersonae (see comments
under A. lilaceps) are somewhat similar but much larger.

Agaricus fuscovelatus. Note the dark color of the unbroken veil and the more or less equal (cylin¬
drical) stalk. The flesh turns orange or reddish when cut.
AGARICUS 325

Agaricusfuscofibrillosus (Bleeding Agaricus) Color Plate 74

CAP 5-13 cm broad, convex to plane; surface dry, smooth, with flattened brown to reddish-
brown fibrils which only rarely break up into scales. Flesh thick, firm, white, quickly
staining red when cut or bruised; odor mild or faintly fruity. GILLS close, free at maturity,
pinkish becoming reddish-brown, finally chocolate-brown or darker. ST ALK 4-10(14) cm
long, 0.8-2 (3) cm thick, equal or thicker below, hollow or stuffed; smooth and white above
the ring, whitish below or with brownish to vinaceous fibrils or scales, or in one variant
sheathed by a “boot” of brownish to vinaceous fibrils; staining red quickly when bruised.
VEIL membranous, white or becoming brownish in age; forming a superior, skirtlike ring
on stalk. SPORE PRINT chocolate-brown; spores 5-6 (7) * 4-4.5 microns, elliptical,
smooth. Cap surface not yellowing in KOH.
HABITAT: Solitary to scattered or gregarious on ground under cypress or rarely other
trees; sporadically common in our area in the late fall and winter, but probably more
widely distributed (in habitats other than cypress).
EDIBILITY: Edible and “rich” according to Shalom Compost, who put it in a soup.
Reddish-staining Inocybes (e.g., /. pudica) are poisonous but smaller, with paler spores.
COMMENTS: This Agaricus is easily recognized by its “bleeding” flesh and brown to
reddish-brown fibrillose cap. It is one of several “bleeders” that have traditionally been
lumped under the names A. haemorrhoidarius and A. silvaticus. (The “true” A. haemor-
rhoidarius has distinct fibrillose scales on the cap and is said to be fairly common in the
forests of eastern North America, but has not yet been found in California; A. silvaticus
has been reported from the Pacific Northwest.) A. fuscofibrillosus, on the other hand, has
an innately fibrillose cap that does not normally become scaly, and grows almost exclu¬
sively with cypress ( in our area). There are several other red-staining species, including:
A. benesi(-A. albosanguineus), cap pure white or tinged grayish to pale cinnamon, widely
distributed but rare except in our area, where it is fairly frequent under cypress; A.patter-
sonae, a larger, thicker-stemmed species (see comments under A. lilaceps);A. rubronanus,
a very small species (cap 2-3 cm broad, stalk 3-6 mm thick); and a large, stately, delicious
but unidentified species with dark reddish-brown fibrils or fibrillose scales and minute
spores (4-5 microns long). I have found the latter several times in mixed woods, but it is not
common. A. arorae also “bleeds,” but usually grows with oak and stains yellow in KOH.

Agaricus arorae
CAP 3-7 (10) cm broad, convex to plane; surface dry, with brownish to reddish-brown
fibrils or fibrillose scales, at least at the center, on a white to reddish background; staining
reddish when rubbed. Flesh white, reddening when cut (usually quickly, but sometimes
slowly); odor mild or faintly fruity. GILLS free at maturity, close, pinkish becoming
purple-brown, then chocolate-brown or darker. STALK 5-14 cm long, 0.5-1.5 (2) cm
thick, equal or enlarged at base, hollow in age; white, sometimes with scaly zones, staining
reddish when bruised and often aging reddish-brown or vinaceous. VEIL membranous,
thin, white (or dingy in age), forming a median to superior, skirtlike, fragile ring on stalk.
SPORE PRINT chocolate-brown; spores 4-5.5 * 3-4 microns, broadly elliptical, smooth.
Cap surface staining yellow in KOH.
HABITAT: Solitary to widely scattered or in small groups on ground in mixed woods and
under oaks, fruiting mainly in October and November, sometimes common. It is known
only from Santa Cruz County, California, but probably has a wider distribution.
EDIBILITY: Unknown.
COMMENTS: This odd species appears to be a compromise or “bridge” between the red-
stainingand yellow-staining (rufescent and lutescent) sections of A garicus. The flesh when
326 AGARICACEAE

fresh bruises red quite dramatically, but the surface of the cap turns yellow in potassium
hydroxide! In the field it can be separated from other“bleeding” species such as A.fusco-
fibrillosus by its slimmer build (see photo on p. 311) and fondness for oak. A. amicosus
also reddens when cut and yellows in KOH; it is common under mountain conifers in the
Southwest and southern Rockies, and is edible. A. spissicaulis has a stocky stature like A.
bisporus (and is sometimes mistaken for that species) and slowly reddening flesh, yellows
somewhat in KOH, has a slight almond odor, and is widely distributed but rare.

Agaricus subrutilescens (Wine-Colored Agaricus) Color Plate 75

CAP 5-15 (20) cm broad, convex or with a somewhat flattened top, becoming broadly
umbonate to plane or with an uplifted margin; surface dry, covered with brown to
purple-brown or wine-colored fibrils or fibrillose scales(sometimes only at the center) ona
whitish to dingy background. Flesh white, firm, not staining when bruised; odor mild or
slightly fruity. GILLS close, free at maturity; at first whitish, then pinkish, then darkening
slowly to pinkish-brown and finally chocolate- or blackish-brown; when pinkish usually
bruising brighter rosy-pink when cut. STALK 5-20 cm long, 0.5-1.5 (4)) cm thick, equal or
thicker below, smooth and white or reddish above the ring, sheathed with soft cottony
white scales below which break up into fibrillose patches in age or wear away; stuffed or
hollow, often fragile in age. VEIL membranous, white, forming a thin, usually superior,
skirtlike ring on stalk. SPORE PRINT chocolate-brown; spores 4.5-6 x 3-4 microns,
elliptical, smooth. Cap surface usuallystaininggreenish-oliveinKOH(sometimesslowly).
HABITAT: Solitary, scattered, or in small groups in woods, usually under conifers;
common in the fall and winter along the west coast, also reported from Japan. In our area it
favors redwood, but occurs in Oregon under alder, in northern California under Sitka
spruce, in southern California under oak, and in the Sierra foothills in mixed woods.
EDIBILITY: Edible and choice, but has been known to cause rather severe gastric upsets
in some people. It is not as meaty as many Agaricus species, but it is delicious.
COMMENTS: The dark purple-brown cap and shaggy white stem distinguish this hand¬
some mushroom from other Agaricus species. Along with A. hondensis, it is our most
common Agaricus of deep, undisturbed woods. A. hondensis, however, has a paler cap
and smooth (not shaggy) stalk. Because of the greenish KOH reaction and a number of
other chemical peculiarities, A. subrutilescens is something of an anomaly. It does not
fit well into any of the existing “sections” of Agaricus, but the rosy-staining gills suggest
a possible relationship to the red-staining species (section Sanguinolenti). It is usually
a rather tall and slender mushroom, but robust individuals can be found, particularly
under pine. A small Texan species, A. vinaceo-umbrinus, also stains greenish in KOH.

Agaricus hondensis (Felt-Ringed Agaricus)

CAP 6-15 (20) cm broad, convex becoming plane; surface dry, smooth, whitish or with pale
pinkish-brown to pinkish-gray to fawn-colored flattened fibrils or fine fibrillose scales
(at least at center), the fibrils often darkening in age to brown, reddish-brown, or reddish-
gray, but in one northern form darker brown from the beginning. Flesh thick, white,
unchanging or staining pale yellowish when bruised, then often slowly discoloring
pinkish; odor of crushed flesh mild or faintly phenolic, but usually distinctly phenolic in
base of stalk. GILLS pale pinkish to pinkish-gray becomingbrown, thenchocolate-brown
or darker; free at maturity, close. ST ALK 7-20 cm long, 1 -2.5 cm thick but with a thicker or
bulbous base; firm, smooth, without scales, white or discoloringdingy pinkish or brownish
in age or after handling; flesh in extreme base usually staining pale yellowish when bruised.
VEIL membranous, white, forming a thick, feltlike, superior ring on stalk; ring skirtlike
Agaricus hondensis. A common woodland species with a large, thick, felty ring, smooth stem, and
pale brown to reddish-brown fibrils on the cap. Tempting, but not edible!

but often flaring outward instead of collapsing against stalk. SPORE PRINT
chocolate-brown; spores 4.5-6 * 3-4 microns, elliptical, smooth. Cap surface staining
yellow in KOH.
HABITAT: Solitary or in groups, troops, or rings in woods, particularly where there are
thick accumulations of fallen twigs and other debris. Very common in our area in the late
fall and winter under both hardwoods and conifers; like A. subrutilescens, which grows
in similar habitats, it is apparently restricted to the west coast. The main fruiting typically
follows close on the heels of A. praeclaresquamosus. It was originally described from
La Honda, California.
EDIBILITY: Poisonous to many people, causing stomach distress, vomiting, etc. It’s
difficult to imagine a more delicious-looking mushroom, but it has an unpleasant, astrin¬
gent-metallic taste even when cooked.
COMMENTS: This handsome, alluring woodland Agaricus is distinguished by the pale
cap fibrils which often darken in age, the thick felty annulus(ring), smooth naked stalk, and
chocolate-brown spores. The phenol odor is often absent in the cap but usually quite
pronounced when cooked or if the base of the stalk is broken open and crushed. It is
frequently mistaken for edible species, particularly A. subrutilescens and A. augustus,
both of which have cottony scales on the stem when young. It has also been confused
with A. silvaticus, a species with reddening flesh (see A. fuscofibrillosus).

Agaricus californicus (California Agaricus) Color Plate 72

CAP 3-9 (12) cm broad, at first convex or marshmallow-shaped, the margin inrolled
and often somewhat lobed, then expanding to broadly convex or plane or broadly umbo-
nate; surface dry, smooth or with fibrils or small scales; color variable: most often white to
silvery gray with a brown center, but sometimes entirely white and at other times with
brown to grayish-brown fibrils or scales throughout; usually darker in age and often with a
metallic luster; often pinkish- or reddish-stained in wet weather; margin often extending
beyond gills. Flesh thick, white, unchanging or staining slightly yellowish when crushed,
then eventually sordid reddish or brown; odor of crushed flesh phenolic but often faint.
GILLS close, free at maturity, usually pallid until the veil breaks, then bright pink to
pinkish-brown, finally chocolate-brown to blackish-brown. STALK 3-8 (12) cm long,
0.5-1 (1.5) cm thick, equal or slightly enlarged at base, stuffed or hollow, smooth, without
scales; white, but often discoloring pinkish to dingy brown in age or after handling. VEIL
membranous, thick, white, often with felty patches on underside; forming a persistent,

327
Agaricus californicus is often confused with A. campestris and A. bisporus. Note how the gills are
whitish in the button stage, and how the veil forms a thick persistent annulus (ring) on the stalk. Also
compare the shape of the young caps to that of A. campestris and A. bisporus.

superior to median ring on stalk or occasionally clinging to cap margin; ring skirtlike or
intermediate. SPORE PRINT chocolate-brown; spores 5-7 * 4-5 microns, elliptical,
smooth. Cap surface staining yellow in KOH.
HABITAT: Solitary to scattered or densely gregarious (but not often in rings) onground,
growing anywhere and everywhere, but especially abundant in suburbia—on lawns,
in gardens, parks, under trees (eucalyptus, acacia, oak, cypress, etc.), also in the woods,
but only rarely in pastures (except under trees). Found year-round in our area, but most
abundant in the fall, when it often mingles with Lepiota naucina and other Agaricus
species. It is known only from the west coast, but may very well have a wider distribution.
EDIBILITY: Mildly poisonous to some people (causing stomach upsets), and frequently
mistaken for the common meadow mushom (A. campestris), with which it often grows.
COMMENTS: This ubiquitous mushroom gives my students more trouble than any
other, not only because it closely mimics edible species, but also because of the endless
variation it exhibits. “Typical” forms—if they can be said to exist—do not stain bright
yellow like A. xanthodermus, but yellow slightly in cooking, in addition to giving off a
phenol odor. The persistent membranous ring, modest size, rather long stalk, slight
phenol odor when fresh (often not evident to those unfamiliar with the odor) and whitish
(not pinkish!) gills in the button stage (i.e., before the veil breaks) help distinguish it from
A. campestris, A. cupreobrunneus, and A. bisporus. The cap varies tremendously in color
and scaliness, but is usually brownish at least at the center, and often somewhat shiny. As
a rule, specimens growing in sheltered situations (such as under oak) are paler or even pure
white, while those growing in the open can be quite dark. Specimens on lawns are usually
rather firmly rooted in the ground and often have to be dug up to avoid breaking the stem.
For a comparison with A. campestris, see that species and the key to Agaricus. Since it is
the most ubiquitous Agaricus in coastal California (one quickly tires of finding it), the
name A. californicus is certainly appropriate. It has also been called the “Fool’s Agaric,”
but I don’t care for this name because fools aren’t the only ones to be fooled by it!

328
AGARICUS 329

A garicus praeclaresquamosus (Flat-Top Agaricus) Color Plate 76

CAP 5-25 cm broad, at first convex or somewhat marshmallow-shaped, then broadly
convex or plane; surface dry, covered with flattened inky-gray to grayish-brown or brown
fibrils or fibrillose scales (at least at center) on a whitish background, but often developing
reddish or pinkish stains in wet weather; in one form bruising yellow. Flesh thick, white,
unchanging or staining slightly yellow when bruised and then slowly discoloring brownish
or vinaceous; odor of crushed flesh phenolic (especially in base of stalk). GILLS close, free
at maturity, at first pallid, then grayish or light pink, becoming reddish-brown to
chocolate-brown and finally blackish-brown. ST ALK 7-18 cm long, 1 -3 (4) cm thick, equal
or enlarged below or sometimes tapering to a point if growing in clusters; stuffed or hollow,
smooth, without scales; white, but often discoloring reddish-brown to dingy brown in age
or upon handling; flesh in extreme base usually (but not always) staining bright yellow
when cut. VEIL membranous, white, thick, feltlike, somewhat rubbery, often splitting at
the margin; rupturing to form a persistent, superior, skirtlike or intermediate ring on stalk.
SPORE PRINT chocolate-brown; spores 4-6.5 * 3-4.5 microns, elliptical, smooth. Cap
surface staining yellow in KOH.
HABIT AT: S olitary or in groups or clusters in woods or under trees, especially along roads
and paths; widely distributed, but especially common along the west coast. It is frequent
in our area in the fall and winter, and occasional in the early spring. The main crop is usually
in November or December, but I have seen gorgeous fruitings of the large form (see
comments) under redwood in Humboldt County, California, in September.
EDIBILITY: Poisonous to many, causing vomiting and diarrhea, but some people eat it
with impunity. Like A. hondensis, it is tempting, but has an unpleasant metallic taste.
COMMENTS: Better known as A. meleagris, this beautiful omnipresent Agaricus can
be told by the inky to grayish-brown fibrils on the cap, plus the thick veil, smooth stem,
and phenolic odor of the crushed flesh. In addition, the base of the stalk often bruises bright
yellow when nicked, as in A. xanthodermus. The above description embraces two forms:
one medium-sized (but generally larger than A. californicus) and sometimes clustered, the
other very large and strikingly beautiful (about the size of A. augustus) and often gre¬
garious but rarely clustered. The latter is especially tempting for the table(see color plate!),
but can be distinguished from A. augustus by the smooth stem, grayer cap, and different
odor. The larger form is usually found under redwood, whereas the medium-sized form
grows under various trees, including redwood and oak. Other species: A. placomyces, a
common woodland mushroom in eastern North America, is very similar to the medium¬
sized form of A. praeclaresquamosus, but has a more fibrillose (less scaly) cap, a less
rubbery veil, and often shows yellowish to brown droplets on the underside of the veil or
on the stalk. Another eastern species, A. pocillator, closely resembles A. placomyces,
but lacks veil droplets and often has a double ring, is usually smaller and slimmer, and
sometimes has a rimmed (cuplike) basal bulb. It is especially common in the South. Both
of these species are eaten by some people, but are best avoided.

A garicus xanthodermus (Yello w-S taining Agaricus)

CAP 6-15 (20) cm broad, round to somewhat marshmallow-shaped or convex, then
expanding to broadly convex or plane; surface dry, smooth, pure white to gray or grayish-
buff, or often whitish at the margin and buff to tan toward the center; often discoloring
brownish in old age and sometimes breaking up into scales; typically staining bright yellow
quickly when rubbed repeatedly, especially on the margin, but then slowly discoloring
brownish or vinaceous; margin inrolled somewhat when young and often lobed. Flesh
thick, firm, white, turning yellowish when crushed; odor phenolic (unpleasant). GILLS
Agaricus xanthodermus growing with Alyssum (the flowers) along a road. Note how the sliced speci¬
men at far right is stained yellow at base of stalk. Cap color ranges from white to grayish to buff.

close, free at maturity, at first white, then pinkish or grayish-pink, finally chocolate-
brown to blackish-brown. STALK 5-12 (18) cm long, 1-2 (3) cm thick, equal or with an
enlarged base, smooth, stuffed or hollow, without scales; white, usually bruising yellow,
then brownish; in age often discolored brownish; flesh in very base turning bright yellow
when cut. VEIL membranous, white or yellow-stained, usually with patches on underside;
forming a large, thick, feltlike, median to superior ring on stalk; ring skirtlike or inter¬
mediate, often flaring at first. SPORE PRINT chocolate-brown; spores 4.5-6 * 3^1.5
microns, broadly elliptical, smooth. Cap surface staining yellow in KOH.
HABITAT: Scattered to densely gregarious under trees and hedges, in yards, on lawns,
along roads and paths, also in woods, pastures, and under cypress; widely distributed.
Abundant in our area from fall through spring and even showing up in the summer. I have
seen stupendous fruitings in an old olive orchard, under acacia and eucalyptus, and in
numerous cypress plantations as well as with oak.
EDIBILITY: Poisonous to many people—causing headaches, nausea, vomiting, and
diarrhea. It is a very tempting, meaty mushroom but the unpleasant odor becomes
unbearably obnoxious when the mushroom is cooked, and the taste is awful as well.
Yet there is no accounting for taste. I know one person who gathers it—and nothing else!
COMMENTS: The tendency of all parts to stain bright yellow, the phenolic odor, and the
white to grayish or tan cap plus the presence of a veil and chocolate-brown spores are the
trademarks of this cosmopolitan and variable species. The yellow-staining is usually most
dramatic on the margin of the cap, but is most reliable in the extreme base of the stalk, which
turns bright chrome-yellow when nicked. The latter is perhaps the best fieldmark, since

Agaricus xanthodermus. Note yellow stains on cap and the prominent annulus in mature specimens.
These handsome white examples of Agaricus xanthodermus might easily be mistaken for Agaricus
arvensis or another edible species. Their phenolic odor, however, is usually quite pronounced and the
very base of the stalk turns bright yellow when cut.

only the equally poisonous A. praeclaresquamosus stains so brilliantly in the base of the
stalk. Several edible Agaricus species, such as A. augustus and A. albolutescens, stain
yellow on the cap, but do not subsequently discolor brownish. The edible horse mushroom
(A. arvensis) and giant horse mushroom (A. osecanus groupj superficially resemble A.
xanthodermus, but do not stain yellow at the base of the stalk and do not smell like phenol.

Agaricus pinyonensis* (Pinyon Agaricus)

CAP 6-20 cm broad, convex or shaped like an inverted bowl, expanding slightly in age but
not normally becoming plane; surface smooth or occasionally breaking up to form a few
small scales; entirely white or with a tan- to buff-tinged center, often discoloring buff to very
pale tan overall in old age; sometimes yellowing when bruised. Flesh very thick (at least 2
cm) and firm, white, typically not staining; odor mild or merely mushroomy. GILLS
pinkish when young, then darkening to purple- or chocolate-brown and finally blackish-
brown; close, free at maturity; narrow in relation to the flesh. ST ALK 5-20 cm long, 2-5 cm
thick or more, often swollen below (just above the base); white and smooth or nearly so,
sometimes discoloring like the cap in old age; firm; solid, stuffed, or with a hollow center.
VEIL thick, membranous, white, usually with patches on underside when young; typically
forming a median to superior skirtlike ring on stalk. SPORE PRINT chocolate-brown;
spores 6.5-8 * 4-5 (?) microns, broadly elliptical, smooth. Cap surface yellowing in KOH.
HABITAT: Solitary or in groups on ground under pinyon and juniper; known only from
New Mexico, but probably occurring throughout the Southwest in the appropriate
habitat. It fruits mainly in August and September, but does not appear every year.
EDIBILITY: Delectably delicious—a favorite of fungus-fanciers in the Southwest. Its
discoverers and namers, Bill Isaacs and Chuck Barrows of Santa Fe, New Mexico, rate it
among the best in the genus (“firm like A. bitorquis and milder than A. campestris”).
COMMENTS: This magnificent white mushroom is remarkable for its thick flesh, firm
texture, and fine flavor. It is somewhat reminiscent of A. bitorquis, but its veil is quite
different and it apparently grows only with pinyon and juniper. Its name (which has yet to
be officially published and may actually turn out to be A. “barrowsii”) is therefore apt,
but its affinities within the genus Agaricus are rather unclear. It may be related to A. arven-
*Provisional name by Isaacs & Barrows

331
Left: Agaricus pinyonensis is a robust whitish species of the Southwest. Note how thick the flesh is!
Right: Agaricus arvensis is a sweet edible species with a distinct anise odor and a tendency to discolor
yellow in age. For close-up of unbroken veil in this species, see photo on p. 312.

sis, but does not normally smell sweet and does not necessarily stain yellow when bruised.
Other species: A medium-sized to fairly large (cap 4-14 cm) whitish species, A. sequoiae,
grows solitary to gregarious or in large clusters in the redwood and mixed forests of
coastal northern California, usually near rivers or at least in bottomlands. It has a longer
(8-30 cm) and more slender stem that is usually equal, and it does not stain yellow when
bruised or in KOH, but sometimes has a yellow-tinged veil. It has a mild odor and is edible
according to Rick Kerrigan, who named it.

Agaricus arvensis (Horse Mushroom)

CAP (4) 7-20 cm broad, oval or convex becoming broadly convex or plane; surface dry,
smooth or sometimes cracking into small scales, especially at the center; white to creamy,
buff, or yellowish (especially toward center), usually bruising yellow if rubbed (especially
when young); margin sometimes hung with veil remnants. Flesh thick, firm, white,
unchanging or yellowing slightly when crushed; odor sweet (like anise or almond extract)
when young, often somewhat musty in age. GILLS close, free at maturity, pallid becoming
grayish (rarely pinkish), then chocolate-brown or darker. ST ALK 5-12 (17) cm long, 1 -3 cm
thick, equal or slightly enlarged below, stuffed or hollow, smooth or with small cottony
scales below the ring; white, sometimes bruising or aging yellowish, but extreme base not
bruising yellow when cut. VEIL membranous, white or tinged yellow, with cottony patches
on underside that often split to form a starlike or cogwheel pattern; forming a fragile,
superior, skirtlike ring on stalk. SPORE PRINT chocolate-brown; spores 7-8.5 * 5-6
microns, elliptical, smooth. Cap surface staining yellow in KOH.
HABITAT: Solitary, scattered, or in groups in grassy areas—lawns, pastures, etc.;
common and widely distributed. It is frequently encountered in our area on lawns and in
cemeteries in the spring, summer, and fall, but seems to be replaced in our pastures by the
giant horse mushroom (A. osecanus groupj.
EDIBILITY: Edible (for most people) and excellent. Buttons have a slightly sweetish
flavor intermediate between that of A campestris and A. augustus. Be sure not to confuse
it with the deadly white Amanitas, which have white spores, white gills, and a volva.
COMMENTS: The white to pale yellowish cap, sweet odor when young, well-developed
membranous veil which often has a cogwheel pattern of patches on its underside (see photo
on p. 312), chocolate-brown spores, and growth in grass typify this beautiful, cosmopoli¬
tan mushroom. The cap will often age or stain yellowish—more so than that of A. croco-

332
AGARICUS 333

dilinus and A. osecanus. However, the base of the stalk does not stain bright yellow when
cut—an easy way to distinguish it from the poisonous, yellow-staining A. xanthodermus.
The stature of A. arvensis is generally more robust than that of A. silvicola, its smaller-
spored sylvan look-alike, but it is not nearly as ponderous as that of the giant horse mush¬
room (A. osecanus groupj or A. crocodilinus. The growth in grass is quite characteristic,
though a swamp-inhabiting form (var. palustris) has been described. There are numerous
variants around A. arvensis that have not been critically evaluated, including a small,
slender form (cap only 3-9 cm broad) that occasionally occurs in our area on lawns.
A. fissuratus is a very similar coastal species with a frequently cracked (fissured), white
to yellowish cap and slightly larger spores. It is common in the Puget Sound area of
Washington, but probably has a wider distribution, and is edible.

Agaricus osecanus group Color Plates 78,79,81

(Giant Horse Mushroom)
CAP 7-40 cm broad, rounded or convex, then broadly convex to plane; surface dry, often
very smooth (like kid leather), but frequently with minutely cottony scales in youth and
sometimes cracking into scales or warts in age or dry weather; white, or discoloring buff
or yellowish in age (especially toward center); surface bruising slightly yellowish when
young and fresh (but usually not staining in age). Flesh thick, very firm when young but
quite soft in age, not usually bruising yellow; odor slightly sweet (like almond extract or
or anise) when young, but often unpleasant (musty or like wet straw) at maturity. GILLS
close, free in age, white becoming grayish or grayish-pink, then reddish-brown, finally
chocolate-brown to blackish-brown. STALK 4-16 cm long, (1.5) 2.5-5 (8) cm thick, often
spindle-shaped (swollen at or below the middle) but sometimes equal; firm, usually solid;
smooth above the ring, often with small pointed but easily-obliterated scales below; white,
sometimes aging yellowish or rusty-yellow; flesh in base not staining bright yellow when
nicked. VEIL membranous, white, with patches on underside which form a cogwheel
pattern or adhere to stalk as downward-pointing scales; rupturing to form a median to
superior, skirtlike or collapsed ring on stalk. SPORE PRINT chocolate-brown; spores
(5)6-7 x 4.5-5.5 microns, elliptical, smooth. Cap surface yellowing in KOH.
HABITAT: Solitary to scattered or in groups or rings in pastures, lawns, and other grassy
areas; distribution uncertain, but extremely abundant at times in our coastal pastures
(when they are “popping” it is not unusual to pick 100 pounds from a single pasture—along
with some giant puffballs!) There is usually one crop in the fall and another, larger one in
the spring, but it is also to be expected during warm spells in the winter. Its range extends at
least into northern California and Oregon, where the very similar A. crocodilinus (see
comments) is also common.
EDIBILT Y: Edible (for most people), and very popular with my group of acquaintances.
The young, hard buttons are best—mature specimens have very soft flesh and a musty odor
and do better in the compost pile than the frying pan.
COMMENTS: This fine and forthright fungus can be spotted from afar, shining like a light
bulb when its spotless white skin catches the sun. It is essentially a larger, squatter, robuster
version of the horse mushroom (A. arvensis), but differs in having a solid, often swollen
stem and smaller spores. Also, the sweet odor and yellow bruising reaction of young spe¬
cimens are weaker than in typical A. arvensis (and may actually be non-existent), while
the musty odor that develops in age is stronger. The appearance of the cap—whether warty
or smooth—seems to depend largely on age, exposure, and weather conditions (when
growing in the open or exposed to the wind it is more likely to be warty or cracked). Be
careful not to confuse it with the poisonous A. xanthodermus, which may also be large
and white, but has a phenol odor, yellows more rapidly when bruised, and stains bright
yellow in the base of the stem.
 *
*

s- *
•••• •%

;
■ - ,. - / -T > ** ^^ v
m*w

W^sSmSSSBBmSBMS^m^

Agaricus crocodilinus, mature specimens. It is common for the caps to be warty, but they can also
be smooth. Note the squat stature. Each of these caps is about one foot across. (Bill Everson)

yellow in the base of the stem. An equally edible but larger-spored species, A. crocodilinus
(“Crocodile Agaricus”—see photo above) is common in parts of northern California
and the Pacific Northwest, and has also been reported from the prairies of eastern New
Mexico and Colorado. It is just as large or even larger than the giant horse mushroom, and
looks virtually the same. True, it is more apt to be warty or scaly and is often slightly
browner in age, but it can also be perfectly smooth and pure white, so that in regions where
both occur the two species can only be separated with certainty by measuring the spores
(in A. crocodilinus they are 8-11 (14) microns long). Our giant horse mushroom,
incidentally, may very well be an unnamed, endemic species. It belongs to a group or
“complex” that is very confused taxonomically, and is listed here as a member of the
A. osecanus group largely because of its spore size. However, it differs somewhat from the
“true” A. osecanus of Europe, and could just as well be called A. nivescens—another
European species with a solid stem and a shade smaller spores. Until a critical study of this
group is completed, it is perhaps best to call it “giant horse mushroom,” because its
common name cannot be subsequently invalidated or proved “incorrect!” Since all of
these species are equally and unequivocally edible, the exact (or inexact) differences
between them needn’t concern you. At least, they don’t concern me! Other species:
A. macrosporus (-A. villalicus) of Europe is a large-spored species very similar to, if not
identical with, A. crocodilinus; the suggestive moniker/l. urinescens has been given to still
another large-spored variant—a fitting tribute to the indiscreet odor that frequently
develops in old age.

Agaricus silvicola group (Woodland Agaricus)

CAP 5-12 (18) cm broad, convex becoming plane; surface dry, smooth or silky-fibrillose,
sometimes obscurely fibrillose-scaly in age; white, usually aging yellowish, especially at the
center, and staining at least slightly yellow when bruised, particularly on margin. Flesh
firm, white, unchanging or yellowing slightly when crushed; odor sweet (like anise or
almond extract), at least when young. GILLS close, free at maturity, white becoming gray
or pinkish-gray, then brown and finally chocolate-brown or darker. STALK 5-14 (20) cm
long, 1-2 (2.5) cm thick, usually enlarged below, stuffed or hollow, smooth or with small
cottony scales below ring; white or pinkish at apex, white below, but often aging or bruising
yellowish; base not staining bright yellow when cut. VEIL membranous, white or stained

334
Left: This stately member of the Agaricus silvicola group is fairly common in our area under tanoak.
Cap is white and stains yellow. Right: Agaricus albolutescens, also common, resembles A. silvicola
but is usually more robust, has a stronger odor, and stains more dramatically.

yellow, with patches on underside that sometimes form a cogwheel pattern; forming a
prominent, superior, skirtlike ring on stalk. SPORE PRINT chocolate-brown; spores
5-6.5 x 3.5-4.5 microns, elliptical, smooth. Cap surface staining yellow in KOH.
HABITAT: Solitary, scattered, or in small groups in woods; widely distributed and
common, but rarely fruiting in large numbers. In our area it is fairly common in the fall
and winter under oak, tanoak, and conifers; at higher elevations it fruits under conifers.
EDIBILITY: Edible (for most people) and choice, with caution. Make sure there is no
volva and that the mature gills are not white—I have seen it mix company with deadly
Amanitas that were very similar in size, shape, and color!
COMMENTS: Also spelled A. sylvicola, this species or species “complex” is recognized
by its white cap, tendency to stain or age yellow, anise odor (sometimes faint, but usually
detectable when the young flesh is crushed), skirtlike annulus (ring), chocolate-brown
spores, and woodland milieu. It rather closely resembles A. arvensis, but is usually more
erect and less robust, has smaller spores, and does not grow in grass. There are several
variants in need of critical study. The “typical” A. silvicola grows mainly under conifers,
at least in the West, but a large, very stately and striking variety (see photo) is fairly common
in our area under tanoak, while a form with an abruptly bulbous stem^zl. abruptibulbus)
occurs in California as well as in eastern North America. None of these forms stain as
dramatically as A. albolutescens, and they do not smell as strong nor are they as robust as
that species. Other species: A. chionodermus looks similar, but has bright pink to reddish
gills for a long time, a mild odor, and a white to silvery-white, often fibrillose cap. It grows
mainly under conifers and is edible and apparently widely distributed—I have seen it in
New Mexico. See also A. sequoiae (under A. pinyonensis), A. summensis (under A.
augustus), and A. albolutescens.

Agaricus albolutescens (Amber-Staining Agaricus)

CAP 7-18 cm broad, convex to broadly convex or plane; surface usually dry, smooth or
fibrillose, at first white but quickly staining amber to yellow-orange when bruised and
often entirely yellowish to yellow-orange to amber or ochraceous in age; margin often hung
with veil remnants. Flesh thick, white, usually bruising yellowish if crushed; odor strongly
sweet and aniselike or almondy. GILLS free at maturity, close, pallid becoming grayish
or grayish-pink, then eventually chocolate-brown or darker; often bruising yellow when
immature. STALK 5-14 cm long, 1.5-3 cm thick, usually enlarged below (up to 5 cm

335
This squat fragrant Agaricus is common in the Sierra Nevada during the spring and early summer. It
appears to be a form of A. albolutescens.

thick), firm, white or discoloring yellowish, smooth above the ring, smooth or slightly
cottony-scaly below; flesh in base not usually bruising bright yellow, but exterior of base
may. VEIL membranous, white or yellow-stained, with patches on underside that some¬
times form a cogwheel pattern; rupturing to form an ample, superior, skirtlike ring on
stalk. SPORE PRINT chocolate-brown; spores 5-7 x 3.5-4.5 microns, elliptical, smooth.
Cap surface staining yellow in KOH.
HABITAT: Solitary, scattered, or gregarious on ground in woods; known only from the
West. In our area it is quite common from late fall through early spring under oak(often
in the company of A. hondensis). In the Sierra Nevada a vernal variant occurs.
EDIBILITY: Delectably delicious, with a strong sweet flavor. As with most Agaricus
species, some people are adversely affected by it.
COMMENTS: This woodland species is reminiscent of A. silvicola, but is easily distin¬
guished by its squatter or more robust stature, much stronger odor, and amber-staining
cap. It also tends to occur in larger numbers than A. silvicola, at least in my experience.
(I have seen fairy rings containing nearly 100 specimens!) The rapid yellow-staining of the
cap can lead to confusion with the poisonous A. xanthodermus, which eventually discolors
brownish after staining yellow, smells like phenol, and stains bright yellow in the base of
the stalk. A. arvensis is also somewhat similar but usually grows in open, grassy places. In
the Sierra Nevada and other mountain ranges, what appears to be a variant of A. albo¬
lutescens is often common (especially in the spring). It has a fibrillose cap and its stem is
tougher and more cylindrical (less bulbous) than the coastal version, but it is otherwise
quite similar (see photograph above).

Agaricus subrufescens (Almond Mushroom) Color Plate 82

CAP (6) 8-25 cm broad, round or marshmallow-shaped becoming convex to plane; surface
dry, smooth but with fine flattened fibrils which may break up into minute scales except at
the center(but in dry weather sometimes cracking into large warts); fibrils pallid to buff to
pale brown, pale pinkish-brown, or fawn-colored, often becoming browner or ruddier with
age; background white to pinkish-buff, often becoming yellow in age or when bruised.
Flesh thick, firm, white, not bruising yellow or only very slightly; odor strongly sweet (like
almond extract), especially when young. GILLS free at maturity, close, whitish becoming
grayish or pinkish, then reddish-brown and finally chocolate-brown or darker. ST ALK 6-
15 cm long, 1.5^4 cm thick, equal or with an enlarged base, smooth or with a few fibrils or
scales below the ring; white, but often staining or aging yellow; base often staining yellow to
yellow-orange but flesh in base not staining bright yellow when cut; base sometimes with
white mycelial threads attached. VEIL thick, membranous, white, usually with patches
(often obscure) on underside; forming a superior, skirtlike ring on stalk. SPORE PRINT
chocolate-brown; spores 5.5-7 x 4-5 microns, elliptical, smooth. Cap surface staining
yellow in KOH.

336
Agaricus subrufescens is a rare but remarkable species. The odor and flavor are strongly almondy, but
the stalk is not nearly as shaggy as that of A. augustus, and the cap is paler.

HABITAT: Scattered to densely gregarious or clustered in compost, manure, and rich

soil; originally described from eastern North America and apparently widely distributed,
but rare. I have seen it fruit by the hundreds in manure-filled trenches on a berry farm near
Watsonville, California, in the late spring, summer, and early fall (sometimes accom¬
panied by Lepiota lutea and L. americana). It is easily grown at home in a vegetable garden
(if you use compost) and mycelium from the just-described “patch” is now being marketed
for this purpose. It has the advantage of liking it hot, meaning you can enjoy it at the same
time as your other summer vegetables (A. bisporus is also grown easily in gardens, but
likes it much cooler).
EDIBILITY: Edible (for most people) and delicious! The almondy odor and taste is much
stronger than that of A. augustus—so strong, in fact, that some people don’t care for it. I
put several specimens in a cream-of-mushroom soup and those who tasted it swore I had
dumped in a bottle of almond extract! It was apparently cultivated around the turn of
the century, but lost out to A. bisporus—which raises an interesting prospect: if it had
become the “cultivated mushroom” instead of A. bisporus, would the word“mushroomy”
have a completely different (sweet and almondy) connotation?
COMMENTS: The strong almond extract odor and taste plus the robust stature, presence
of an annulus, chocolate-brown spores, and light brownish cap make this a fairly easy
mushroom to recognize. The cap is quite similar in color to that of A. hondensis, but the
habitat and odor are completely different, while the paler color and smoother stem distin¬
guish it from A. augustus. Other species: In eastern North America, an edible Agaricus with
tawny scales on the cap (like A. augustus), a shaggy stalk, and weakeralmond odorhasalso
gone under the name A. subrufescens, apparently because of its similarly-sized spores.
Whether it is a distinct species or merely an extreme form of A. subrufescens is for licensed
Agaricus-experts to decide.

Agaricus augustus (The Prince) Front Cover, Color Plate 77

CAP 7-30 (40) cm broad, usually marshmallow-shaped but sometimes convex, slowly
expanding to plane or with an uplifted margin; surface dry, covered with numerous dark
brown to warm brown or tawny-brown fibrils or fibrillose scales on a white background
that usually becomes yellowish, buff, or ochre in age, giving an overall golden tone to many
mature specimens; center often darker; surface sometimes breaking up into warts in dry
weather and bruising yellow when rubbed, at least when young. Flesh thick, white, firm;
odor sweet (like almond extract), especially when young. GILLS close, free at maturity,
remaining pallid for a long time, finally turning grayish-brown (or very briefly pinkish),
eventually chocolate-brown to blackish-brown. STALK 8-35 cm long, 1-4 (6) cm thick,
equal or enlarged slightly below, the base usually buried deep in ground; white, but often
aging or bruising yellowish; smooth above the ring, sheathed with white or brown-tipped

337
Agaricus augustus. Note the marshmallow-shaped caps of these unexpanded buttons. In this stage
they are delicious! See also Color Plate 79, photo on front cover, and photo on p. 311.

scales below (but these often wearing away in age); rather tough and fibrous. VEIL
membranous, with white to brown cottony patches on underside (but these sometimes
disappearing); forming a large, ample, superior, skirtlike ring on stalk. SPORE PRINT
chocolate-brown; spores 7.5-10 x 5-6 microns, elliptical, smooth. Cap surface staining
yellow in KOH.
HABITAT: Solitary or in groups or clumps on ground in the woods (especially under
redwood), but usually near roads and paths, in clearings, and other places where the soil
has been disturbed; sometimes also in flower beds, composted areas, under trees in towns,
in arboretums, etc.; widely distributed, but frequent only along the Pacific Coast and
perhaps more common in our area than anywhere else. It shows a definite preference for
warm weather, fruiting in the spring, summer, and fall—or in the winter if it’s mild enough.
Several crops are produced each year, so visit your patches regularly. It is curious that such
a large mushroom requires so little moisture to fruit—a little fogdrip is all it needs!
EDIBILITY: Edible (for most people) and one of the very best! It’s especially significant
because it fruits in the spring and summer, when edible fungi are scarce in our area. It’s like
getting two mushrooms in one: delectably sweet and almondy when young, strong and
mushroomy at maturity. Unfortunately maggots, slugs, sowbugs, and centipedes are fond
of it too!
COMMENTS: This prince of a mushroom is distinguished by its large size—caps of “LP”
dimensions (one foot) are not uncommon—almond extract odor, yellow-staining cap with
brown fibrils or scales, prominent annulus (ring), chocolate-brown spores, and shaggy
stem (when young) which is usually buried in the ground. The veil is large and exquisitely
constructed, and the marshmallow shape of the young caps (i.e., somewhat flattened on
top) is also characteristic, though several other mushrooms (e.g., A. praeclaresquamosus,
Lepiota rachodes) hae the same shape. The gills remain whitish for a long time—falsely
suggesting that the spores are white—but eventually they will darken to brown and finally
chocolate-brown. The poisonous A. praeclaresquamosus is often common in the same
habitats, but has grayer cap fibrils, a smoother stem, and smells like phenol. A. hondensis
also has a smooth stem, while the edible A. perobscurus has darker cap fibrils when young, a
smoother stem, and subtler odor. Other species: A. smithii (also called A. perrarus) is a

338
AGARICUS 339

similar but somewhat smaller, more slender version with an ochraceous to ochraceous-
brown cap and almondy odor. It is common in northern California and the Pacific
Northwest under coastal conifers, particularly Sitka spruce. A pale form of A. augustus
also occurs, but is rare. It resembles the typical form but has pale yellow cap fibrils which
give the cap an overall creamy-white appearance. Somewhat similar to this pale form is A.
summensis—a large, robust species whose cap is whitish with yellowish to ochraceous-
ta wny fibrils. It has a thick, often bulbous stem(2.5 cm thick or more) and is rare. I have seen
it in our area under forest trees along roads. Finally, A. augustus has a smaller-spored look-
alike in eastern North America (see comments under A. subrufescens). All of the above
species are edible and delicious.

Agaricusperobscurus (The Princess)

CAP 7-20 cm broad, convex or marshmallow-shaped becoming broadly convex to plane
or with margin slightly uplifted; surface dry, at first covered with dark (grayish-brown
to dark brown or blackish) fibrils which tend to break up in age to form scattered, sparse
grayish-brown to dingy olive-brown fibrillose scales on a whitish to ochre-buff background
(but center usually remaining dark); bruising yellow only slightly if at all. Flesh thick, white,
not usually bruising yellow; odor typically of almond extract but often faint or even absent.
GILLS close, free at maturity, pallid becoming pinkish or pinkish-brown, then brown and
finally chocolate-brown or blackish-brown. STALK 7-15 cm long, 1-3 cm thick, equal or
more often enlarged at base; white or discoloring slightly in age, smooth above the ring,
slightly scaly or fibrillose below, but often appearing smooth, especially in age. VEIL
membranous, with cottony patches on underside (but these often disappearing), forming
an ample, superior, skirtlike ring on stalk. SPORE PRINT chocolate-brown; spores 6.5-
7.5 * 4-5 microns, elliptical, smooth. Cap surface staining yellow in KOH.
HABITAT: Solitary or in groups under or near trees (usually planted) or shrubs, some¬
times also at the edges of woods; known only from coastal California. It is fairly frequent
in the San Francisco Bay region from late fall through spring or even summer, but uncom¬
mon to rare elsewhere (not nearly as numerous as A. augustus). I usually find it under pine,
cypress, and acacia, but have never seen it in the redwood forests where A. augustus thrives.
EDIBILITY: Edible and choice—about as good as A. augustus.
COMMENTS: This “sister” to A. augustus is likely to be mistaken for that species, but has
a subtler almond odor and less pronounced tendency to stain or age yellow, a smoother

Left: Agaricus perobscurus is a smoother-stalked version of A. augustus. Note how cap is quite dark in
these young specimens. Right: Agaricus semotus (see comments under A. micromegathus on p. 340)
is a small woodland species with a distinct anise odor and pinkish to reddish to brownish cap fibrils.
340 AGARICACEAE

(less shaggy) stalk, and consistently smaller spores. In addition, the gills pass through a
more marked pinkish phase before turning brown, and the fibrils on the cap are much
darker when young(the fibrils may be quite sparse in age, however, giving the cap an overall
paler appearance). It is also likely to be mistaken for the poisonous A. praeclaresquamosus,
but that species has a totally smooth stalk, a phenol odor, often stains bright yellow in
the base of the stem, and is more common in habitats characteristic of A. augustus than
those favored by A. perobscurus.

Agaricus micromegathus (Anise Agaricus)

CAP 2-5 (8) cm broad, convex to plane; surface dry, with silky fibrils or fibrillose scales
(sometimes very sparse toward margin); fibrils at first pinkish in one form, yellow-brown
in another, but in both forms becoming brownish to grayish-brown in age on a whitish to
buff background; yellow to orange stains often developing in age or upon handling. Flesh
thin, fragile, white, unchanging or bruising yellowish; odor distinctly sweet (aniselike or
almondy). GILLS free at maturity, close, pallid or grayish, then pinkish, finally chocolate-
brown or darker. STALK 2-6 cm long, 3-10 mm thick, equal or slightly enlarged at base,
white or staining yellow to orange upon handling (especially below); stuffed or hollow,
fragile at maturity. VEIL membranous, thin; forming a fragile, superior to median ring on
stalk, or disppearing entirely; ring skirtlike or intermediate. SPORE PRINT chocolate-
brown; spores 4.5-5.5 * 3.5-4 microns, broadly elliptical, smooth. Cap surface staining
yellow in KOH.
HABITAT: Solitary to scattered or gregarious in lawns, pastures, fields, and other grassy
or open places; widely distributed. Fairly common in our area in mild weather, especially in
the fall, often in the company of A. campestris and A. cupreobrunneus.
EDIBILITY: Edible, but thin-fleshed and fragile. Its fragrance suggests that the flavor is
good, but it doesn’t often occur in enough quantity to invite collecting. Chuck Barrows
of Santa Fe, New Mexico, attained a small degree of notoriety by smothering an ice cream
cone with raw A. semotus (a very similar species—see comments).
COMMENTS: This is one ofseveralpuny,poorly-knownspecieswithananise odor, slight
annulus (ring), and fragile flesh. They have a tendency to become yellow or rusty-yellow
in age, and can be distinguished from most other Agaricus species by their size alone. Simi¬
lar species with a distinct anise odor include: A. semotus (see photo on p. 339), with brown
to reddish fibrils on the cap and slightly longer stalk, found in woods and under trees (espe¬
cially oak); and A. comtulus (also spelled A. comptulus), with a white to yellowish cap
that is usually tinged brown or buff at the center, found in grass. Both of these are edible
and common, but rarely collected because of their small size. Also see A. diminutivus.

Agaricus diminutivus group (Diminutive Agaricus)

CAP 1 -4 cm broad, oval or convex becoming plane or slightly umbonate; surface dry, with
flattened pink to purplish-pink to amethyst-gray to reddish-brown fibrils, at least at the
center; margin often paler. Flesh thin, white, not staining; odor mild or faintly fragrant
(like anise). GILLS free at maturity, close, pallid or pink becoming reddish-brown, then
chocolate-brown or darker. STALK 2-7 cm long, 2-6 mm thick, equal or witha small basal
bulb, white or pallid, but in age often stained yellowish or orange below the ring; stuffed
or hollow, fragile. VEIL membranous, thin, white; forming a fragile superior to median,
skirtlike ring on stalk which often disappears in age. SPORE PRINT chocolate-brown;
spores 4.5-6 * 3.5-4.5 microns, broadly elliptical, smooth. Cap surface yellowing in KOH.
HABITAT: Solitary, widely scattered, or in small groups in humus in woods, widely
distributed. The most common variant in our area fruits in the late falland winter in mixed
woods and under live oak, but farther north this group is common under conifers.
 >

Agaricus diminutivus group. Note small size, slender stature, and fragile veil that may or may not
form a distinct ring on the stalk.

EDIBILITY: Presumably edible, but too small to be of value and sometimes confused
species of Inocybe (e.g., I. pudica), most of which are poisonous.
COMMENTS: Members of this species “complex” are easily distinguished from other
Agaricus species by their petite size, reddish-pink to purplish cap color, and woodland
habitat. The form illustrated is the common one in our area. It has a beautiful amethyst-
tinged cap and seems to fit descriptions of A. purpurellus (a member of the A. diminuti¬
vus group) quite well. However, there are probably more names (e.g., A. dulcidulus, A.
amethystina) in this group than there are species, so a critical study is necessary before
any positive identifications can be made.

COPRINACEAE
THIS is a large family of fragile mushrooms with deep brown to black spores and a carti¬
laginous or fragile stem. The gills are not decurrentasin Gomphidiusand Chroogomphus,
and those species with dark brown spores do not normally have free gills and an annulus
(ring) on the stalk, as in Agaricus. The cap cuticle is cellular(composed of round or pear-
shaped cells) as opposed to filamentous as in the Strophariaceae, but since this distinction
is microscopic, it is better to learn the characteristics of each genus than attempt to distin¬
guish the two families in toto (for instance, Coprinus has gills that digest themselves, and
Panaeolus usually grows on dung or manure). As a rule, however, the fragile white to
brown species with a dry or only slightly tacky cap belong to the Coprinaceae, while those
with a viscid and/or brightly colored cap belong to the Strophariaceae. Both families
should be checked when in doubt. There are three common genera in the Coprinaceae,
keyed below.

Key to the Coprinaceae

1. Mature gills (and often the cap) digesting themselves, i.e., either turning into an inky black
fluid or withering away . Coprinus, p. 342
1. Gills not digesting themselves ... 2
2. Spore print black; sides (faces) of gills often mottled; growing in grass, dung, or manure . .. .
.Panaeolus, p. 353
2. Not as above; growing in humus, wood chips, on wood, etc., or if in grass then spore print dark
brown or purple-brown; not normally growing in dung . Psathyrella, p. 361

341
342 COPRINACEAE

COPRINUS (Inky Caps)

Minute to medium-sized or sometimes large, ephemeral, saprophytic mushrooms. CAP usually
oval, cylindrical, or conical when young, usually striate at maturity in the smaller species and often
translucent or deliquescing. GILLS free or attached, gray to black and autodigesting at maturity.
STALK usually white and hollow, thin and fragile, or if thick, then cartilaginous. VEIL present
or absent, sometimes forming an annulus (ring) on stalk. VOLVA typically absent (but a rudimen¬
tary one present in some species). SPORE PRINT typically black {but deep brown in a few species).
Spores mostly elliptical, smooth or roughened, with a germ pore; discoloring in concentrated
sulfuric acid. Cap cuticle usually cellular.

MEMBERS of this genus are called inky caps because the gills and often the cap digest
themselves at maturity, turning into an inky black fluid that drips to the ground. The auto¬
digestion or deliquescing of the gills plus the black spore print are the main diagnostic
features of Coprinus. In Bolbitius the gills sometimes liquefy but the spore print is rusty-
brown, while in Psathyrella and Panaeolus the spore print may be black but the gills do
not deliquesce.
The autodigestion process is a unique method of spore dispersal that should not be
confused with the normal process of decay that occurs in most mushrooms. Rather than
maturing at an even rate, the spores near the margin of the cap ripen first. Enzymes are
simultaneously released which dissolve the surrounding tissue, causing the edge of the cap
to spread out and curl back. This pulls the gills apart, enabling the spores to be discharged
into the air. If you look at the gills of a shaggy mane (Coprinus comatus), you’ll see that
they are crowded together like the pages of a book. If autodigestion did not occur, the
spores would be discharged onto adjoining gills and their dispersal would be greatly
impeded. Of course, many spores are trapped in the inky liquid that drips to the ground,
but millions more are successfully discharged into the air.
In the larger inky caps, both the cap and gills dissolve, leaving nothing but a few rags of
tissue stuck to the top of the stalk. This phenomenon was undoubtedly the inspiration for
Shelley’s memorable lines:
Their mass rotted off them flake by flake
Til the thick stalk stuck like a murderer’s stake,
Where rags of loose flesh yet tremble on high
Infecting the winds that wander by.

In the smaller species, such as C. plicatilis, an inky fluid is not necessarily formed. There
is so little substance to the gills that they wither instead of liquefying, and the cap may be so
thin that it is translucent-striate (the gills can be seen through it as radiating lines).
Inky caps frequent areas inhabited by livestock and human beings. They fruit whenever
it is moist and mild enough, and are among our most common urban and suburban
mushrooms. The smaller types are especially prevalent on dung and manure, while the
larger ones crop up in gardens, cellars, along roads, on or around stumps, in disturbed
soil, and humus. A few grow in the woods. As we alter the environment and create new
“niches,” we can expect new species to evolve. One winter, following three weeks of rain,
an unidentified Coprinus sprouted from the orlon carpet of my leaky ’67 Rolls Canardly
(it rolls down one hill and canardly make it up the next). My friends naturally assumed it
was the result of carrying around so many mushrooms in my car, yet it was a species I had
never seen before!
Several of the larger inky caps are good edibles, and the shaggy mane, C. comatus, is a
popular and unmistakable favorite. Most species, however, are much too thin and insub¬
stantial to warrant collecting. A few are said to be poisonous, and the common inky cap,
C. atramentarius, contains a compound which reacts with alcohol in the body to produce a
very peculiar set of symptoms (see p. 896). Coprinus species should be eaten before they
Autodigestion in Coprinus comatus. Left: The gills have just begun to blacken and liquefy at the
cap margin. Right: Dry weather has caused the autodigestion process to cease with a few rags of tissue
left. In wet weather it continues until only the stalk remains. (Rick Kerrigan)

deliquesce, and consequently should not be kept overnight. They are among the most easily
digestible of all fungi, but their high water content makes them unsuitable for some dishes.
As they frequently occur in large numbers, you may wish to preserve them for later use.
This is best effected by marinating, drying, or sauteeing and then freezing. The “ink” from
the fleshier species, incidentally, makes a very passable ink if diluted with a little water.
Coprinus is a large genus, with over 200 species known. It is a difficult group to study
because the fruiting bodies are so ephemeral (sometimes lasting only a few hours) and
because many species fruit where you don’t normally look for mushrooms. Consequently,
the North American species are not well known. As their favored habitats (dung, asphalt,
etc.) are ubiquitous, most inky caps are widely distributed and likely to occur in California.
The smaller ones are especially difficult to identify—even with the aid of a microscope. A
few distinctive and/ or common types are presented here.

Key to Coprinus
1. Growing on dung, manure, straw, or compost. 2
1. Growing on ground, wood chips, wood, or indoors . 9
2. Partial veil typically forming a distinct annulus (ring) on stalk . 3
2. Annulus absent or rudimentary . 5
3. Cap minute (usually less than 1 cm broad); annulus disclike or saucerlike, sometimes as large
as the cap; common on horse dung . C. ephemeroides, p.352
3. Not as above; fruiting body larger . 4
4. Cap 2-5 cm high when unexpanded, at first entirely white or white with brownish scales.
. C. sterquilinus & others (see C. comatus, p. 345)
4. Not as above . 9
5. Cap at first covered with white to buff or gray universal veil remnants (hairs, fibrils, or powdery
or flaky scales) . 6
5. Universal veil absent; cap hairless or with only a few minute hairs . 8
6. Cap white or pallid with white to pale gray powdery or mealy scales .
.C. niveus & others (see C. radiatus group, p. 351)
6. Not as above (but cap may be covered at first with white fibrils or hairs) . 7

343
344 COPRINACEAE

7. Cap minute (usually less than 8 mm high before expansion)' .C. radiatus group, p. 351
7. Cap usually larger than above .C.fimetarius & others (see C. lagopus group, p. 350)
8. Cap minute (1-5 mm high before expansion) .C. miser (see C. plicatilis, p. 352)
8. Cap small, but typically larger than above .C. ephemerus (see C. plicatilis, p. 352)
9. Cap cylindrical before expansion and 4-25 cm or more tall, usually at least somewhat shaggy,
entirely white or with a brown center and/ or brownish scales; partial veil present when young,
often forming a movable annulus (ring) that may drop off; cosmopolitan C. comatus, p. 345
9. Not as above . 10
10. Stalk tough and woody.(see Podaxales & Allies, p. 724)
10. Not as above . 11
11. Growing in deserts or arid regions, typically terrestrial (not on dung); cap usually with one or
more thick, feltlike, persistent patches of universal veil tissue; stalk sometimes with a volva
at the base . 12
11. Not as above . 14
12. Volval patch (veil tissue) on cap usually single and stellate (with starlike arms or points) . 13
12. Volval patch or patches irregular, not stellate; base of stalk bulbous with a volva-like rim . . .
.C. xerophilus
13. Stalk typically with a volva-like basal bulb . C. asterophora
13. Basal bulb and volva lacking or rudimentary, at least at maturity.C. asterophoroides
14. Gills with yellow edges; stalk with yellow hairs . C. sulphureus
14. Not as above . 15
15. Base of stalk or immediate vicinity with cinnamon to yellow-brown or yellow-orange mycelial
threads or a woolly mycelial mat (oozonium); growing on wood (often indoors) .
. C. radians (see C. micaceus, p. 348)
15. Not as above . 16
16. Cap typically 5 cm broad or more when expanded, dark brown (or becoming black) beneath a
layer of whitish universal veil tissue which breaks up into discrete patches or flakes or “spots”;
stalk fairly thick (usually 0.5-1.5 cm); terrestrial . C. picaceus, p. 346
16. Not as above (cap may have a coating of whitish hairs, but if so, then differently colored or stalk
thinner). 17
17. Cap at first clothed with white to grayish hairs (see Color Plate 86), gray to dark brown or black
beneath the hairs; stalk typically long, slender, fragile . . . C. lagopus group & others, p. 350
17. Not as above . 18
18. Cap with large flaky patches or flat scales which may wear off in age; growing in clusters (occa¬
sionally scattered) on dead hardwoods (but wood may be buried) in eastern North America
. C. quadrifidus, C. variegatus, & C. americanus (not good edibles—some variants are bitter)
18. Not as above . 19
19. Cap whitish except at center; partial veil usually forming a membranous ring on stalk; growing
on rotting wood . C. alnivorus (see C. comatus, p. 345)
19. Not as above . 20
20. Cap at first with a dense coating of white universal veil fibrils, whitish to pale tawny beneath the
fibrils or if darker then growing in ashes; stalk 3-10 mm thick C. domesticus & others, p. 349
20. Not as above; cap bald or With whitish particles, or if with whitish fibrils then usually darker
or brighter than above (beneath the fibrils) and not found in ashes; stalk thin or thick . . 21
21. Stalk typically 6 mm thick or more; cap 2-8 cm broad and/ or high, brown to grayish .
.C. atramentarius, p. 347
21. Stalk typically 1-6 mm thick; cap up to 5 cm broad or high, but if larger than 3 cm then typically
buff to yellow-brown or reddish-brown when fresh . 22
22. Cap conspicuously pleated in age (see Color Plate 87) and often expanding to plane; usually
growing scattered or at least not in clumps. C. plicatilis, p. 352
22. Not as above; often in clumps or clusters . 23
23. Stalk typically 3-8 cm long and 2-6 mm thick .C. micaceus & others, p. 348
23. Stalk typically 1-4 cm long and 1-2 mm thick . 24
24. Gills crowded .C. impatiens (see C. disseminatus, p. 352)
24. Gills well-spaced or fairly close but not crowded .C. disseminatus, p. 352
Shaggy manes in egg batter and bread crumbs, anyone? This beautiful cluster of Coprinus comatus
is at the perfect stage for eating. How do they taste? See photo on p. 890 for the answer.

Coprinus comatus (Shaggy Mane) Color Plate 85

CAP 4-15 (25) cm tall, cylindrical or columnar, expanding somewhat as margin curls up
until it is more or less bell-shaped, then deliquescing from the bottom up; surface not vis¬
cid, white with a brown to pale cinnamon-brown or buff center, soon breaking up into
shaggy white to brown scales (universal veil remnants) which often recurve in age; margin
striate in age and often tattered. Flesh soft, white. GILLS very crowded (like pages in a
book), free or nearly so, at first white, then passing through delicate shades of pink, pinkish-
red, or vinaceous; finally black and inky (deliquescing). STALK 5-20 (40) cm long, 1-2 cm
thick, tapering upward or with an enlarged, more or less pointed base; smooth, white,
hollow or stuffed with a pith, cleanly separable from cap. PARTIAL VEIL membranous,
forming a small, white, movable, inferior ring which often drops to the base of the stalk or
falls off. SPORE PRINT black; spores 10-16 (18) x 7-9 microns, elliptical, smooth, with
a germ pore.
HABITAT: Sometimes solitary but more often scattered to densely gregarious or in loose
clumps on hard ground and grassy areas, rich or disturbed soil, etc. Common throughout
the northern hemisphere, especially along roads and trails. In our area the major crop is
in the fall or early winter, but it can be found most any time. In the Southwest I have seen
troops of them crowding highway shoulders for miles—both in the spring and fall. For
some reason “shags” seem to gravitate toward asphalt and often burst up through it.
They’ve been known to ruin tennis courts, and one is reported to have lifted a 10 pound slab
of concrete in a heroic attempt to proliferate its species. Alexander Smith describes a
fruiting of over 1000 specimens on a baseball field, “extending in a line from first base
to short left field.”
EDIBILITY: Edible and delicious—one of the best known and safest of all wild mush¬
rooms. The flavor is very delicate but the texture is marvelous—not slimy as in okra, but

345
346 COPRINACEAE

succulent as in octopus. For a delicious snack, slice them in half and dip them in egg batter
and bread crumbs (or flour), then saute them briefly and serve hot! Use only young caps—
darkened ones are mostly water—and don’t pick more than you can eat in two meals unless
you plan to preserve them, for they will deliquesce quickly. In rare instances they may
react with alcohol in the body to produce effects similar to those of C. atramentarius.
However, I have cooked them in wine many times with no ill effects.
COMMENTS: Shaggy manes are the soldiers among mushrooms, the sentinels of the
roads. Their tall, shaggy cylindrical heads are distinctive even in silhouette, and when inky
individuals are found in the vicinity of young ones, there can be no doubt as to their identity.
The poisonous Chlorophyllum molybdites can be similar when young, but expands in age
and doesn’t deliquesce. Podaxispistillaris is quite similar in shape and color, but lacks gills.
“Shags” are as pleasing to the eye as to the palate, especially in the delicate reddish tints the
gills assume as they mature. “Shags” also possess a special spontaneity—seemingly
popping up overnight after a rain, while most mushrooms fruit several days after. Extreme
aridity may arrest development so that the spores never mature and the gills remain white,
or the gills blacken and then shrivel up rather than deliquescing. Several forms and varieties
of the shaggy mane have been described, including a small, oval one only 5-6 cm high. The
height of the larger forms seems to depend partially on the depth of the humus they must
transcend—I have found specimens in redwood duff by a road that were two feet tall—but
the “normal” height is 6-20 cm. Other species: C. colosseus is a giant version of the shaggy
mane with stalk 35-50 cm long and spores 17-20 microns long; it has been described from
Washington by Fred Van De Bogart, but is rare. Several small (cap less than 5 cm high)
versions of the shaggy mane also occur, including: C. palmer anus, terrestrial; C. alnivorus,
wood-inhabiting; C. sterquilinus and C. umbrinus, on dung or compost piles, the former
with large spores and the latter with a brownish stem base; and C. spadiceisporus, on the
dung of wild animals (rabbit, deer, etc.). All of these have the ring (annulus) characteristic
of the shaggy mane and are too small and rare to be worth eating.

Coprinuspicaceus (Magpie Mushroom)

CAP (3)5-8 cm broad, oval or cylindrical when young, broadly bell-shaped in age; surface
at first covered with whitish universal veil material which soon breaks up into discrete
patches, flakes, or “spots” on a dark brown to eventually black background; finely striate
nearly to the center in age, and deliquescing. Flesh thin, fragile, soft. GILLS white, soon
becoming reddish- or vinaceous-tinted, finally black and inky (deliquescing); crowded,
more or less free. STALK 7-15 (25) cm long, 0.5-1.5 cm thick, base usually enlarged and
with woolly hairs; hollow or stuffed, white, smooth, fairly fragile. PARTIAL VEIL
absent. SPORE PRINT black; spores (13) 14-20 * 10-13 microns, elliptical, smooth.
HABITAT: Solitary or in small groups on ground in woods and along paths, etc.; widely
distributed, but not common. I have found it only once in our area, but it is not uncommon
in southern California when it is damp enough. In Europe it grows under beech and is said
to be partial to alkaline soil.
EDIBILITY: Poisonous, at least to some people.
COMMENTS: The mottled brown-and-white or black-and-white cap makes this one of
the easiest of all inky caps to recognize. Large specimens are reminiscent of the shaggy
mane in shape—but the resemblance ends there. Greg Wright of Los Angeles, who has
had more experience with this species than I, says the odor is also distinctive—like that of
“mothballs, a hot grass pile, or burnt hair, or rank.” Other species: In my own pathetic
excuse for a garden I have found a somewhat smaller, mottled (black-and-white), unidenti¬
fied Coprinus with a distinct annulus (ring) on the stalk.
Coprinus atramentarius, the common inky cap. Note hollow stalk in the beautiful slice at bottom.

Coprinus atramentarius (Inky Cap; Tippler’s Bane)

cap 2 -8 cm high and/ or broad, round or oval and often somewhat lobed when young,
becoming conical to bell-shaped, or in old age convex; deliquescing from the margin
toward the center; surface dry, smooth or with silky whitish fibrils (universal veil tissue)
when young, grayish-brown to grayish-tan when young (the center often browner or with
small brown scales and the margin sometimes pallid), becoming lead-gray to inky-gray in
age; margin striate or grooved, usually tattered or splitting in age. Flesh thin, soft, pallid or
grayish. GILLS free or nearly free, crowded, at first white, soon gray or with pinkish to
vinaceous tints, finally black and deliquescing(becominginky). STALK4-15(20)cmlong,
0.6-1.5 cm thick, equal or with a narrowed or enlarged base; white or with grayish to brown¬
ish fibrils below; hollow. PARTIAL VEIL fibrillose and evanescent or leaving a median
to basal ring or ridged zone on stalk, or sometimes even a rudimentary volva near base.
SPORE PRINT black; spores 7-12 * 4-6 microns, elliptical, smooth.

Coprinus atramentarius usually grows in groups, tufts, or clusters. Left: These specimens have
rounded smooth caps. Right: A large scaly-capped cluster.
348 COPRINACEAE

HABITAT: Scattered to densely gregarious or in massive clumps in cultivated areas,

lawns, gardens, roadsides, around or on old stumps, etc., sometimes also in the woods;
widely distributed and very common, fruiting in our area practically year-round. It used to
be an unwanted intruder in cultivated mushroom beds, and is one of our characteristic
“suburban” mushrooms. The largest fruiting I’ve seen in the wild was under aspen in New
Mexico. Several crops are generally produced each year.
EDIBILITY: Edible when young and fairly good—but sometimes reacting with alcohol
in the body to produce a peculiar type of poisoning (see p. 896 for details). The grayish-
brown caps look rather unappetizing—and precisely for this reason were the very first wild
mushroom I ventured to eat. Inky cap-and-salami sandwhiches became a staple item in
my teen-age diet until I discovered finer and more flavorful fungal foods, such as
“Sparassis Sole” and “Agaricus Elegante.”

COMMENTS: The lead-gray to brownish, bell-shaped caps of this species are a familiar
sight in vacant lots and gardens. Several varieties occur, including one with a pointed or
umbonate cap (var. acuminatus) and one with a copious universal veil. The typical variety,
however, has a non-umbonate cap and only slight universal veil remnants on the cap (if
any). The young unexpanded buttons provide an unexpected visual treat when sliced open
lengthwise, but true to their name, they turn into an unsightly inky black mess as they
mature, suitable for writing but not biting. The inky cap sometimes rivals the shaggy mane
in size and abundance and may even mix company. I have seen gigantic 10-lb. clusters
fruiting from an old cut stump in a field, but it does not necessarily grow in clumps. Other
species. C. insignis (-C. alopecia) of eastern North America is a similar species with rough
spores. It grows on hardwood stumps, especially maple, and is poisonous to some people.

Coprinus micaceus (Mica Cap; Glistening Inky Cap)

CAP 2-4 cm high when young, 1.5-5 cm broad when expanded; at first oval, soon bell¬
shaped, then expanding to convex; surface sprinkled at first with minute glistening whitish
particles (universal veil remnants) which often disappear in age, otherwise tan to yellow-
brown to ochre, buff, fulvous, or cinnamon-brown (margin often paler), becoming grayer
in age, especially toward margin; striate at least half way to center, the margin usually
tattered or split at maturity. Flesh thin, pallid or white, soft. GILLS crowded, adnate to
adnexed or free, palllid soon becoming gray or brownish, finally black; deliquescing
(sometimes partially, sometimes completely). STALK 3-8 (12) cm long, 2-6 mm thick,
more or less equal, fragile, hollow, smooth, white or discoloring buff. PARTIAL VEIL
absent or rudimentary, but stalk sometimes with a slight basal ring (presumably formed
by the universal veil). SPORE PRINT dark brown to black; spores 7-11 * 4-6 microns,
elliptical but often compressed (flattened) somewhat, smooth.
Coprinus micaceus. Note white stalk, striate cap, and tendency to grow in clusters. Tattered specimens
at right have autodigested.
A clump of Coprinus micaceus glistening in the rain. Note how the cap is striate nearly to the center.
The fine mica-like particles (veil remnants) often present on the cap have been washed off by the rain.

HABITAT: Gregarious, usually in clusters, on wood or woody debris, around stumps,

or on roots or buried wood (thus appearing terrestrial); widely distributed and very
common. It fruits practically year-round in our area. Like C. atramentarius, it is a common
feature of suburbia but also occurs in the woods (on cottonwood, oak, alder, etc.).
EDIBILITY: Edible, but thin-fleshed and watery. The flavor is said to be good, however,
and since it often occurs in large numbers, it is easy to gather enough for a meal. It does
not always digest itself completely, another point in its favor.
COMMENTS: The cap color of this ubiquitous Coprinus varies considerably, but typi¬
cally falls somewhere between tan and rusty-yellow. It is striate, but not translucent as in
the much smaller C. disseminatus. It might be mistaken for a Psathyrella if the gills do not
liquefy, but old caps become very tattered as the margin gets eaten away. The mica-like
particles responsible for its name are not always evident because they wear away quickly.
Other species: C. silvaticus is a very similar species that lacks the particles when young
and has larger spores. C. radians is also similar, but can usually be distinguished from
C. micaceus by its deliquescing gills and the presence of a cinnamon-brown to yellow-
brown or yellow-orange woolly mycelial mat or network of mycelial strands (oozonium)
in the surrounding substrate. It often grows indoors (on wet wood in basements, bath¬
rooms, etc.), and bountiful crops sprout periodically from the woodwork of a popular
cafe in Santa Cruz, California. Its employees are divided on how to respond: whether to
advertise its presence as a unique organic addition to the existing decor, or whether to
hush it up by organizing daily “search and squish” patrols, for fear of alienating the more
squeamish members of their clientele.

Coprinus domesticus (Domestic Inky Cap)

CAP 1-5 cm high, oval to bell-shaped, expanding slightly in age; surface buff, pallid, or
yellowish with a tawny to ochre or fulvous center, at first covered with silky white to buff
fibrils or patches (universal veil remnants) which often disperse or disappear in age; conspi¬
cuously striate nearly to the center; margin often splitting in age. Flesh thin, pallid. GILLS
crowded, free or adnexed, white becoming gray to reddish-gray, finally black and inky
(deliquescing). STALK 3-8 cm long, 3-10 mm thick, equal or with swollen base, white.

349
Left: Coprinus domesticus, immature specimens. Cap is grayish-buff to whitish and conspicuously
striate. Right: An unidentified charcoal-loving Coprinus (see comments under C. domesticus).
Note striations and prominent universal veil remnants on cap.

hollow. PARTIAL VEIL evanescent or sometimesforminga ragged ring or ridged zone on

stalk, or even a rudimentary volva at its base. SPORE PRINT black; spores 7-10 * 3.5-4.5
microns, elliptical or bean-shaped, smooth.
HABITAT: Gregarious in small, scattered clumps on wood chips, compost, and vegetable
debris in a large garden in Saratoga, California; probably widely distributed. Crops were
produced in the above locality more or less continuously except in very cold weather.
EDIBILITY:“Of very good flavor when cooked,” says Greg Wright of Los Angeles—
but I don’t recommend it because of possible confusion with other species.
COMMENTS: The name C. domesticus has apparently been applied to more than one
species, so its use here must be regarded as tentative. However, our specimens compare
favorably to the original (European) description of the species. It resembles C. micaceus,
but has a somewhat paler cap with silky white fibrils (instead of particles) when young,
and deliquesces to a greater degree. A rather similar but unidentified species with a darker
(ochre to tawny) cap and a white fibrillose universal veil (see photo above) occurs in our
area on ashes.

Coprinus lagopus group (Woolly Inky Cap) Color Plate 86

CAP 1.5-5 (7) cm high when unexpanded, 2-6 cm broad when expanded; at first oval or
cylindrical to acorn-shaped or conical, then expanding to nearly plane with the margin
splitting and often recurving (curling up and back); deliquescing in wet weather; surface
dry, grayish to brownish-gray becoming blackish, but at first clothed with delicate white
to grayish hairs or fibrils (universal veil tissue) which break up into patches and often wear
away in age; striate nearly to the center beneath the hairs, at least at maturity. Flesh very
thin, soft. GILLS more or less free, narrow, fairly close, pallid soon becoming grayish,
then black and inky (but in dry weather merely withering). STALK 4-15 (20) cm long, 2-5
mm thick, equal or tapering upward, very fragile, hollow, white, at first clothed with
minute white hairs (like cap); base with long woolly white hairs. PARTIAL VEIL absent
or evanescent. SPORE PRINT blackish; spores 10-13 x 6-7 microns, elliptical, smooth.
HABITAT: Solitary, scattered, or in dense troops in leaf litter, woody debris, burned
areas, etc. (usually in the woods); widely distributed. Along with closely related species
(see comments), it is fairly common in our area in the fall, less so in the winter and spring.
The fragile fruiting bodies last only a few hours before consuming themselves.

350
COPRINUS 351

EDIBILITY: Probably harmless—but also fleshless and flavorless.

COMMENTS: The above description encompasses a “complex” of closely related inky
caps that are one big headache for compulsive Coprinus-categorizers. In addition to the
“true” C. lagopus, there is the very similar C. lagopides, which has smaller spores (7-10
microns long) and appears to be fairly common in our area. There are also several look-
alikes that are somewhat smaller and grow on dung, manure, or wet straw. These include:
C. fimetarius and C. macrorhizus (the latter with a rooting stem base); C. macrocephalus,
with large spores (11-16 microns long); and C. tectisporus, which grows in greenhouse soil
and only partially deliquesces. To make matters more confusing, the name C. cinereus is
often used, sometimes for a woodland species like C. lagopides but with a taller cap, and
sometimes as a synonym for C. fimetarius and C. macrorhizus. You needn’t know the
exact names of these inky caps, however, to appreciate their beauty (see color plate!), and
they are easily recognized as a group by their slender, fragile, white stalk and striate
grayish cap that is clothed exquisitely with white hairs when young. The dung- and straw-
inhabiting forms, incidentally, are often abundant in our area and are extremely ephemeral
—the caps deliquesce within a few hours, leaving behind only the long, slender, fragile,
bean-sprout-like stalks. (For a miniature dung-inhabiting version, see the C. radiatus
group.) Other species: C. arenatus is also similar to C. lagopus, but grows in sand; C.
narcoticus grows in manure and rich soil and has a strong, unpleasant odor. Finally,
there is C. “chevicolai, ” which fruited one season on the floor of my ramshackle ’67 Chevy
Nova Supersport. It superficially resembled C. lagopus, but was smaller and differed in
several other respects, not the least of which was its choice of abode!

Coprinus radiatus group (Miniature Woolly Inky Cap)

CAP minute, 2-8 mm high, 1-4 mm broad before expanding, at first oval or cylindrical,
then expanding to plane or nearly plane (with the margin usually splitting and curling up),
then deliquescing; surface gray or with a brownish center, but at first covered with long
white to grayish hairs and/or scales (universal veil tissue); striate nearly to the center
beneath hairs. Flesh very thin, soft. GILLS narrow, well-spaced in age, free or nearly free,
pallid soon becoming gray to black, deliquescing with the cap. ST ALK1 -5 cm long, about 1
mm thick, more or less equal, hollow, very fragile, white, with rooting, hairy base.
PARTIAL VEIL absent. SPORE PRINT black; spores 11-14 * 6-8 microns, elliptical,
smooth.
HABITAT: Solitary or in groups on dung, compost, and manure; widely distributed and
fruiting whenever it is damp enough. It is one of our commonest dung fungi, especially on
piles of horse manure mixed with straw.
EDIBILITY: Who knows? Who cares?
COMMENTS: The small size and dense white to grayish hairs on the cap distinguish this
diminutive dung-lover. It is essentially a miniature version of the equally common C.
cinereus-C. fimetarius-C. macrocephalus complex (see comments under the C. lagopus
group), and is itself a “complex” of difficult-to-distinguish forms. Because of its small size
and brief life span it is best observed in a “dung garden” (dung or manure placed in a moist
chamber). Other dung lovers include: C. pseudoradiatus, very similar but with smaller
spores and a non-rooting stalk base; C. niveus, larger, with a mealy snow-white cap (at
least when fresh); C. semdanatus, like C. niveus, but with a slightly darker cap (beneath
the veil remnants); and a host of small species which cannot be reliably identified without
a microscope. Also see C. ephemeroides, which has a large annulus (ring), C. ephemerus
and C. miser (under C. plicatilis), which have a bald or nearly bald cap, and comments
under the C. lagopus group.
352 COPRINACEAE

Coprinus ephemeroides (Ringed Dung Inky Cap)

CAP minute, 1-5 mm broad, convex or in age nearly plane; surface with minute granules or
particles (but may appear smooth), grayish or tinged yellowish to ochre at the center;
practically transparent; striate. Flesh very thin (almost non-existent). GILLS usually
adnexed or free, becoming grayish or black, narrow, soon shrivelling or deliquescing.
STALK 0. 5-6 cm long, about 1 mm thick or less, equal or with a small bulb at base, very
fragile, whitish, smooth; base sometimes with a rudimentary volva (in one form).
PARTIAL VEIL membranous, forming a large, persistent, more or less median ring on
stalk; ring disclike or saucerlike. SPORE PRINT black; spores 6-9 x 3-6 microns, angular
(appearing almost triangular), smooth.
HABITAT: Solitary or in groups on dung and manure (especially of horses) and straw
piles; widely distributed and common. Fruiting in our area whenever it is moist enough.
EDIBILITY: Academic—a tremendous number would be needed for one mouthful!
COMMENTS: One of our tiniest gilled mushrooms, this inky cap is easily recognized by
its growth on dung and the large saucerlike annulus (ring) on the stalk. The ring is some¬
times as broad as the cap, giving the illusion of a stalk with two caps! The name C. ephemer¬
oides should not be confused with C. ephemerus, a slightly larger dung-lover that lacks an
annulus (see comments under C. plicatili's). C. bulbilosus is a synonym.

Coprinusplicatilis (Pleated Inky Cap) Color Plate 87

CAP 0.5-1.5 cm high when young and 1-3 cm broad when expanded; oval or cylindrical
to conical when young, broadly convex or plane in age; surface buff to yellow-brown,
usually with a darker (cinnamon-brown or fulvous) center, in age becoming grayish except
for center; deeply grooved (pleated) nearly to center; margin sometimes recurved in age.
Flesh very thin, fragile. GILLS free (but attached to a collar around stalk apex), well¬
spaced, narrow, soon gray and eventually black, but tending to wither rather than liquefy.
STALK 3-7.5 cm long, 1-3 mm thick, more or less equal, very fragile, thin, smooth, white
or buff, hollow. PARTIAL VEIL absent. SPORE PRINT black; spores 10-13 x 6.5-10
microns, broadly elliptical, smooth.
HABITAT: Solitary, scattered, or in small groups in lawns and other grassy areas, under
trees, in woods, along paths and roads, etc; widely distributed and fairly common in our
area, especially in the fall and early winter. I have seen it fruit prolifically with the similarly
named Marasmius plicatulus in grass under eucalyptus.
EDIBILITY: Unequivocally inconsequential due to its thin flesh.
COMMENTS: The exquisitely pleated cap, small size, and terrestrial habit distinguish
this petite, parasol-like inky cap (see color plate!). The free collar to which the gills are
attached is also distinctive, but is not evident in one form (perhaps a distinct species). It is
larger than C. disseminatus and occurs in smaller numbers, and its cap lacks the white
patches or hairs (universal veil remnants) typical of C. lagopus and most other inky caps.
Other species: C. ephemerus also lacks hairs on its cap, but is slightly smaller and grows in
dung, straw, and compost; C. miser is a minute species (cap 1-5 mm high, reddish-brown
to orange-brown when fresh, but fading) that it also common on dung.

Coprinus disseminatus (Little Helmet; Fairy Bonnet)

CAP 5-10 mm broad, oval soon becoming bell-shaped, then sometimes convex; surface
minutely scurfy when young, then smooth; pallid or buff with a cinnamon-brown to honey-
brown center, becoming grayish toward the margin in age; deeply striate or pleated to the
Coprinus (-Pseudocoprinus) disseminatus. This minute attractive mushroom usually grows in
swarms and does not deliquesce. Note the bell-shaped striate cap and widely spaced gills.

center and translucent at maturity. Flesh very thin, soft. GILLS adnate to adnexed or
appearing free, fairly well-spaced, at first white but soon gray, finally black or slightly
paler; not deliquescing. ST ALK 1.5-4 cm long, 1 -2 mm thick, equal, white or buff, hollow,
fragile, smooth, often curved. PARTIAL VEIL absent. SPORE PRINT dark brown to
black; spores 7-10 * 4-5 microns, elliptical, smooth, with a large apical germ pore.
HABITAT: Densely gregarious (sometimes hundreds) on or near decayed wood and
debris, or on buried wood; usually found in woods or grassy areas, widely distributed.
It fruits throughout the mushroom season in our area, but is not particularly common.
In eastern North America it is often abundant.
EDIBILITY: Too small to be of any value.
COMMENTS: This dainty little fungus grows in troops that do indeed resemble “little
helmets” or “fairy bonnets.” Because the gills do not deliquesce, the “splitters” have
rewarded it with the name Pseudocoprinus disseminatus. It has the general aspect of C.
micaceus, but is much smaller. The translucent, pleated cap separates it from Psathyrella,
(to which it has also been assigned); it is also reminiscent of Mycena, but the spore print is
black or nearly black, not white. Other species: C. impatiens is a similar but slightly larger
species with crowded, at least somewhat deliquescent gills; it usually occurs in smaller
groups of up to one dozen individuals.

PANAEOLUS
Small to medium-sized, fragile, dung- or grass-inhabiting mushrooms. CAP usually bell-shaped or
conical when young (but sometimes convex). GILLS typically attached, gray to deep brown to black
when mature; sides (faces) often mottled. STALK typically thin, brittle or fragile. VEIL present or
absent, but not usually forming an annulus (ring). VOLVA absent. SPORE PRINT typically
black (but dark brown in P. foenisecii). Spores mostly elliptical, with a germ pore; retaining their
color in concentrated sulfuric acid. Cap cuticle cellular.

THIS is a small genus of little brown mushrooms (“LBM’s”) with a bell-shaped to conical
cap and thin, brittle stalk. The sides (faces) of the gills often have a mottled appearance
(see photograph on p. 356) due to uneven maturation of the spores, but they do not deli¬
quesce as in Coprinus. Psathyrellas are similar but do not typically grow in dung, and those
that grow in grass tend to have a convex cap and/or dark brown spores. Psilocybe and
Conocybe are common in dung, but do not have black spores.

353
354 COPRINACEAE

Panaeolus is abundant in pastures, lawns, dung, and manure heaps, fruiting whenever
it’s moist. It often mixes company with other nondescript“LBM’s”(Co/7ocy6e, Agrocybe,
Stropharia, etc.), and would rapidly be relegated to the ranks of fungal forgetability were
it not for the fact that some of its members contain traces of psilocybin and other pupil-
dilating (the term “mind-expanding” being open to debate) compounds. However, in their
search for a more promising and less painful reality, people will stoop to anything, even if
it grows on cow patties and is only two inches tall. Thus, after every rain, our pastures are
marred by hordes of “magic mushroom” hunters, inevitable plastic bags in hand.
Actually, there is considerable confusion as to which species are pupil-dilating. Traces
of psilocybin have been isolated in virtually every species, but its presence and concentra¬
tion is contingent upon a number of genetic, geographic, and environmental factors. My
own experience with the P. campanulatus-P. sphinctrinus complex indicates that a sizable
amount may produce mild hilarity, or even a transitory state of pseudoerotic effervescence.
More often, however, it will produce a queasy stomach and if gulped down too eagerly,
hiccups. It hardly seems worth the effort to harvest and digest the necessary number of
fungal fructifications (30-50), but some people will do anything to “alter” reality, and it can
be argued, I suppose, that doing anything is better than doing nothing.
Indiscriminate sampling of “LBM’s” is foolish, of course, since some are poisonous, so
care must be taken to identify your Panaeolus correctly. Six species are described here.

Key to Panaeolus
1. Cap andlor stalk staining blue to blue-green when bruised or in age . 2
1. Not as above . 3
2. Fruiting body staining blue to blue-green only at base of stalk (or mycelium) and only faintly;
common in temperate regions . P. subbalteatus group, p. 358
2. Fruiting body staining blue to blue-green in most parts (cap, stalk, flesh); largely tropical and
subtropical .P. cyanescens & others, p. 358
3. Cap striate or pleated and/ or very tiny (less than 5 mm broad or high) . (see Coprinus, p. 342)
3. Not as above . 4
4. Partial veil present (covering the gills when young), either forming an annulus (ring) or fibrillose
zone on stalk or leaving toothlike remnants on cap margin . 5
4. Veil absent (check several specimens in different stages if possible) . 7
5. Veil usually forming a distinct annulus on stalk; cap pale (white to buff or pale tan), often viscid
when moist .P. semiovatus, p. 355
5. Veil typically leaving remnants on cap margin (but stalk sometimes with a black ring of spore
dust); cap usually darker than above, not viscid . 6
6. Cap reticulate (netted) or coarsely wrinkled (see photo at bottom of p. 357) .
.P. retirugis (see P. campanulatus group, p. 356)
6. Not as above .P. campanulatus group, p. 356
7. Mature gills and spore print dark brown; growing in grass but not on dung P.foenisecii, p. 360
7. Mature gills and spore print black (gills may be brown when young); in dung, grass, etc. . . 8
8. Cap whitish to buff or dingy yellowish when fresh, 4-10 cm broad; stalk solid, usually at least
4 mm thick . P. solidipes, p. 355
8. Not as above (but cap may fade to whitish as it dries out) . 9
9. Fruiting body very small; stalk whitish or nearly translucent; on dung (seePsathyrella, p. 361)
9. Not as above; common . 10
10. Cap typically bell-shaped or conical, even at maturity .
. P. acuminatus & others (see P. campanulatus group, p. 356)
10. Cap typically convex to plane, at least in age . 11
11. Gills brown before becoming black; cap often with a darker marginal band as it begins to dry
out; often but not always growing in small clusters; common P. subbalteatus group, p. 358
11. Not as above; not common .P.fimicola (see P. campanulatus group, p. 356)
PANAEOLUS 355

Panaeolus solidipes (Solid-Stemmed Panaeolus) Color Plate 83

CAP 4-10 cm broad, rounded to convex or broadly bell-shaped; surface smooth or
wrinkled, not viscid or only slightly so, buff to whitish, or dingy yellowish in age (or gray
from spore dust); sometimes breaking up into small scales as it matures. Flesh fairly thick,
whitish. GILLS adnate to adnexed, close; edges whitish; faces pallid becoming mottled
with gray and black, and finally entirely black. STALK (4) 8-20 cm long, (3) 5-15 mm thick,
equal or thicker below, solid, rather tough, smooth or longitudinally twisted-striate; often
beaded with droplets when young and moist, especially at apex; white to buff or tinged
gray from spores. VEIL absent. SPORE PRINT black; spores 14-22 * 9-14 microns,
elliptical, smooth. Chrysocystidia present on gills.
HABITAT: Scattered to gregarious on dung (especially of horses), manure, and straw;
widely distributed. In my experience it is not particularly common, but often prolific when
it fruits. I have seen large numbers near Saratoga, California, in January, and near
Flagstaff, Arizona, in September. O.K. Miller says it is common during the summer in
Alaska and the Yukon.
EDIBILITY: Edible. I haven’t tried it, but Mcllvaine says, “it is one of the best of the
toadstools.” The relatively large size makes it the only non-hallucinogenic Panaeolus
worth eating.
COMMENTS: The large size and solid stem are remarkable for a Panaeolus. Along with
the black spores, pale cap, absence of a veil, and growth on dung, these features make it
easy to recognize. P. phalaenarum and P. sepulchralis are synonyms, and it has also been
placed in the genus Anellaria (along with P. semiovatus, to which it is closely related). The
latter species, however, has a veil which usually forms a ring (annulus) on the stalk.

Panaeolus semiovatus (Ringed Panaeolus)

CAP 2-6 (9) cm broad and 2-6 cm high, oval at first, then bluntly conical or parabolic;
surface viscid when moist, often shiny when dry, smooth or slightly wrinkled, pale tan
becoming buff or whitish in age, or tinged gray from spores; margin sometimes hung with
veil remnants. Flesh soft, pallid. GILLS adnate to adnexed or seceding; edges whitish,
faces pallid becoming brown or grayish, then mottled with black, and finally entirely
black. STALK (5) 8-15 (18) cm long, (3) 5-10 (12) mm thick, equal or with enlarged base,
stuffed or hollow, whitish to buff, apex often striate. VEIL membranous, usually forming
a superior to median ring on stalk which is blackened by falling spores, but sometimes
merely leaving a fibrillose zone. SPORE PRINT black; spores 15-22 * 8-12 microns,
elliptical to somewhat pip-shaped, smooth. Chrysocystidia present on gills.
HABITAT: Solitary or in groups on dung and manure, especially of horses; widely
distributed and fairly common in mild, wet weather. It is not uncommon in our area, but
the largest fruitings I’ve seen were in New Mexico, in the summer.
EDIBILITY: Edible, according to most sources. However, there is one dubious report
of psilocybin-containing specimens from Colorado.
COMMENTS: Also known as Panaeolus separatus and Anellaria separata, this species
is easily recognized by its white to buff cap, presence of an annulus (ring), and growth in
horse dung (see photograph at bottom of p. 357). Since the cap is slightly viscid, it might
be mistaken for a Psilocybe or Stropharia. The mottled gills, however, signify Panaeolus.
Specimens without a distinct annulus (ring) resemble P. solidipes, but that species has a
thicker solid stem and lacks a veil even in the button stage. The paler cap color helps dis¬
tinguish P. semiovatus from the P. campanulatus group, which can have a “ring” of black
spore dust on the stem.
Panaeolus campanulatus group. This close-up clearly shows the mottled gills and tendency of the
veil to break up into toothlike remnants that cling to the margin of the cap.

Panaeolus campanulatus group (Bell-Shaped Panaeolus)

cap l -4 cm broad and/or high, bluntly conical or bell-shaped, scarcely expanding in
age; surface not viscid, often shiny when dry, smooth or finely wrinkled or often cracking
to form scales (especially in sunlight); some shade of brown, gray, or olive-gray when
fresh (or in one form reddish-brown), paler (olive, tan, or buff) when faded, or dusted
black by spores; margin hung with small, white, toothlike veil remnants, at least when
young. Flesh thin, fragile. GILLS adnate or adnexed but often seceding, fairly close; edges
whitish, faces gray becoming mottled with black; entirely black in age. STALK 6-15 cm
long, 1-3 (5) mm thick, equal or thicker at apex, often quite long and thin, very brittle or
fragile, brown to grayish, minutely powdered. VEIL evanescent, usually seen in young
specimens as flaps of sterile tissue on margin of cap, and sometimes in older specimens as a
thin band of black spore dust on the stalk. SPORE PRINT black; spores 13-18 x 7.5-12
microns, elliptical, smooth.
HABITAT: Solitary or in “families” on or near dung or in grass where cattle have grazed,
sometimes also in compost; widely distributed and very common. It is often abundant in
our coastal pastures in the fall, winter, and spring.
EDIBILITY: Some strains of this species “complex” (see comments) have mild hallucino¬
genic effects when eaten raw in large quantities (see p. 354). More often, however, there
is no effect. Traces of psilocybin as well as serotonin have been isolated.

COMMENTS: The gray to brown, more or less bell-shaped cap, long thin brittle stalk,
mottled gray to black gills, and toothlike veil remnants on the margin of the cap when
young are characteristic of a closely-knit group of dung-lovers: P. campanulatus,

356
Panaeolus campanulatus group. Conical to bell-shaped cap, black gills, long brittle stalk, and growth
on or near dung typify this common species.

P. sphinctrinus, and P. papilionaceus. Since even the experts cannot agree on the exact
differences (if any) between them, it would be presumptuous to attempt to differentiate
them here. At any rate, they are among our most common cow patty (“meadow muffin”)
mushrooms, along with Psilocybe coprophila and Stropharia semiglobata. The stalk is so
fragile that it is difficult to bring home a specimen in one piece. Similar species include: P.
retirugis, a smaller species with an often conspicuously veined or reticulate cap (see photo¬
graph),.also found in our area, but not nearly as common and seemingly partial to horse
dung; the P. acuminatus-P. rickenii group, with a conical to bell-shaped cap and no veil
(check young specimens!); and P. fimicola, which also lacks a veil, but has a more convex
or hemispherical (domed) cap. None of these are worth eating.

Two common inhabitants of horse dung. Left: Panaeolus retirugis (see comments above) is easily
distinguished by its wrinkled or netted (reticulate) cap. Right: Panaeolussemiovatus (-P. separatus)
has a smooth, pale cap and a veil that usually forms a distinct annulus (ring) on the stalk (see de¬
scription on p. 355).
358 COPRINACEAE

Panaeolus cyanescens (Blue-Staining Panaeolus)

CAP 1.5^1 cm broad, bell-shaped to convex; surface smooth or sometimes cracked, not
viscid, brown when moist, fadingtograyishorwhitishasitdries; margin often wavy orsplit
in age. Flesh thin, bruising blue or bluish-green. GILLS adnate to adnexed or seceding,
gray to black, the faces usually mottled. ST ALK 6-12 cm long, 2-4 mm thick, equal or with
a slight bulb at base, usually long and slender, smooth, pallid to yellowish, grayish, or
pinkish, the base brownish or tinged flesh-color; bruising bluish at least somewhat when
handled. VEIL absent. SPORE PRINT black; spores 12-14 * 8.5-11 microns, elliptical,
smooth.
HABITAT: Solitary to widely scattered or in groups on or near dung in pastures;
widely distributed in the tropics and subtropics. It is fairly common along the Gulf Coast
of the United States and also occurs in Mexico and Hawaii. Two similar species (see
comments) have been reported from California.

EDIBILITY: Hallucinogenic—as might be expected of a blue-staining Panaeolus.

P. tropicalis (see comments) is also potent, and both species are gathered by “magic
mushroom” hunters in Hawaii.
COMMENTS: Blue-staining Panaeolus species are largely tropical and subtropical in
distribution. They can be distinguished from the blue-staining Psilocybes by their black
spore print, non-viscid cap (unless very wet), and growth on dung. Other blue-staining
species include: P. tropicalis, with smaller spores (10-12 microns long) and no pinkish
tinge to the stalk; and P. cambodginensis, with an olive to ochre-brown cap when moist,
and smaller spores. All of these are placed by some in a separate genus, Copelandia. The
latter two have been found in southern California on well-fertilized lawns.

Panaeolus subbalteatus group (Belted Panaeolus)

CAP 2-6 cm broad, convex or bluntly conical becoming broadly convex to broadlv
umbonate to plane or with uplifted margin; surface smooth or wrinkled, in age some¬
times breaking into scales(fissured), not viscid; color variable: brown to reddish-brown or
cinnamon-brown when moist, fading as it dries to tan, buff, or even whitish (or grayish
from spores), often with a darker (reddish-brown to brown or dark gray) marginal zone
when partially dry. Flesh thin, brownish. GILLS adnate to adnexed or seceding, close,

Panaeolus subbalteatus group growing on a lawn fertilized with horse manure. The cap is convex
to plane or wavy rather than conical, and usually develops a dark marginal band (as shown at left)
when it begins to lose moisture.
Panaeolus subbalteatus group. Note clustered growth habit, relatively thick stem (for a Panaeolus),
and convex cap. These were growing in a garden fertilized with compost from a mushroom farm.

broad, at first pale watery brown or reddish-brown, darkening gradually to black; edges
whitish, faces usually mottled in age. STALK 4-10 cm long, (1) 3-6(10) mm thick, equal or
tapered at either end, hollow but not fragile, brown to reddish-brown, but often appearing
whitish from a fine powder, or dusted gray by spores; apex often paler; usually longitudi¬
nally striate throughout; base (and mycelium) occasionally staining faintly bluish when
bruised. VEIL absent. SPORE PRINT black; spores 10-14 * 7-9 microns, elliptical,
smooth.

HABITAT: Scattered to densely gregarious—often in small clumps—in manure, com¬

post, and fertilized lawns; widely distributed. In our area it fruits practically year-round
but is most common in warm weather. I’ve seen several hundred specimens in a garden
overgrown with vetch and mulched with compost from a mushroom farm. (It is one of the
major mushroom “weeds” in cultivated mushroom beds.)
EDIBILITY: Hallucinogenic—the psilocybin content varies from moderate to low,
perhaps due to differences in the nitrogen concentration of the substrate. A middle-aged
Santa Cruz woman who mistook them for cultivated mushrooms (Agaricus bisporus) was
hospitalized with hallucinations (a frightening experience if you’re unprepared for them!),
but recovered shortly. It is one of the more popular“recreational” species on the west coast,
and is easily cultivated.
COMMENTS: The above description encompasses what may be a single variable species
or a complex of closely related forms. The key features are the convex to broadly umbonate
to plane cap (not bell-shaped!), brown gills when young, rather firm stalk, absence of a
veil, black spore print, and frequent presence of a darker marginal band on the cap as it
dries out. The tendency to grow in small clusters is also unusual fora Panaeolus. The black
spores and non-viscid cap separate it from Psilocybe, and the larger size and black gills in
age distinguish it from the common, lawn-inhabiting Panaeolus foenisecii, which may
also have a zoned cap. P. subbalteatus is the most common “psilocybin” mushroom in
California, but does not normally stain blue. However, the base of the stalk may occasion¬
ally stain blue very slowly, and it is sometimes coated with cottony, bluish-tinged mycel¬
ium. One variant (also hallucinogenic) that is common on fertilized lawns in our area has
a slimmer stalk (1-3 mm thick) and smaller cap. Whether it is a “new” species or merely a
diminutive form of P. subbalteatus is for the Panaeolus-yun&its to decide.
Panaeolus foenisecii often grows on lawns with Marasmius oreades and Conocybe lactea. Cap is
dark brown when moist, paler when dry, and often features a dark marginal band (shown at left) in
intermediate stages. Gills and spores are brown, never black, and there is no veil.

Panaeolus foenisecii (Haymaker’s Panaeolus)

CAP 1-3 (4) cm broad, bluntly conical to bell-shaped, expanding to convex, broadly
umbonate, or nearly plane; surface smooth or cracking into scales in dry weather, hygro-
phanous but not viscid; chestnut-brown to dark brown or cinnamon-brown when moist,
fading as it dries to dingy buff or tan, often with a darker marginal band when partially
dry. Flesh thin, fragile. GILLS adnate to adnexed or seceding, fairly close, brown be¬
coming deep brown, deep grayish-brown, or chocolate-brown, the faces often somewhat
mottled and the edges paler or whitish. STALK 4-8 cm long, 1.5-4 mm thick, equal or with
an enlarged base, fragile, more or less smooth, white to dingy brownish (often becoming
brown from the base upward). VEIL absent. SPORE PRINT deep brown or purple-
brown; spores 12-17 * 6-9 microns, elliptical, roughened.
HABITAT: Scattered or in groups on lawns and other grassy places; very common in
warm wet weather (or on watered lawns) throughout the northern hemisphere. It is one
of our characteristic spring, summer, and early fall lawn mushrooms, often mingling
with Conocybe lactea, Marasmius oreades, Psathyrella candolleana, and the Agrocybe
pediades group. In contrast to most Panaeolus species, it doesn’t grow on dung, and in
our area it doesn’t even seem to grow in pastures.
EDIBILITY: Harmless in small quantities, but potentially poisonous to toddlers in
the “grazing” stage, since chemical analysis has revealed traces of psilocybin in some
collections. However, western material is apparently “inactive.”
COMMENTS: This pixieish Panaeolus is one of our very common lawn-inhabiting
“LBM’s.” Its small size, thin fragile stem, absence of a veil, growth in grass (not on dung!),
and dark brown spores and gills distinguish it. Because the gills and spore print are not
black as in other Panaeolus species and the spores are not smooth, the “splitters” (see p. 10)
have erected a special genus for it, Panaeolina. Alexander Smith, on the other hand,
dumps it into Psathyrella in his voluminous testament to that genus (his arguments for
placing it in that genus are actually quite persuasive, but he has the weight of tradition
against him in this case). Other species: P. castaneifolius is a similar diminutive lawn
lover with a slightly thicker (3-6 mm) stem, purple-black gills, and purple-black, slightly
roughened spores; I have not seen it in our area but it may occur. P. subbalteatus is also
similar, but has black gills at maturity and black spores and is usually larger. P. acuminatus
(see comments under P. campanulatus group) also grows on lawns, but has smooth black
spores and lacks the marginal band on the cap.

360
 361

PSATHYRELLA
Small to medium-sized mushrooms found mostly on wood or in humus. CAP conical to convex or
plane, often hygrophanous, not usually viscid, typically some shade of brown, buff, or gray. Flesh
markedly fragile. GILLS typically dark brown to black at maturity, usually attached but not de¬
current. STALK usually slender, fragile, and white or pallid. VEIL absent or present, usually not
forming an annulus (ring) on stalk. VOL VA absent. SPORE PRINT deep brown to purple-brown
or blackish, or rarely reddish. Spores smooth or rough, with a germ pore; discoloring in concen¬
trated sulfuric acid. Cap cuticle cellular.

FEW fleshy fungi have less to offer the average mushroom hunter—not to mention the
average human being—than the Psathyrellas. They constitute an immense, monotonous,
and metagrobolizing multitude of dull whitish, buff, grayish, or brownish mushrooms with
a fragile stem, fragile flesh, and purple-brown to blackish spores. Psathyrellas are so
nondescript and unassuming that it’s much easier to define what they are not than what they
are: their gills d o not deliquesce as in Coprinus, their flesh never bruises blue as in Psilocybe,
their cap is not colorful as in Stropharia and Naematoloma, and only rarely is it viscid; and
they don’t grow in dung like Panaeolus. Some Psathyrellas are quite attractive, however,
and all have their indispensable “roles” to fulfill.
Psathyrellas are largely wood inhabitors, but often appear terrestrial because they feed
on wood in the final stages of decay, after all the other wood-lovers have had their fill.
Some species, such as P. candolleana, are ubiquitous, but most favor damp, shady situa¬
tions, especially along trails or streambeds. In California, a very good place to look for
them is in stands of willow and alder, and a very good time to look for them is in the late fall
or early winter, but I can’t think of a very good reason to look for them.
Psathyrellas are listed in older books under several different genera, including Psathyra
and Hypholoma. Several are edible, but most have not been tested and few are fleshy or
distinctive enough to warrant collecting. Alexander Smith has authored an abstruse
monograph in which he describes more than 400 North American species. Most of them
can only be identified if one has a microscope plus a special fondness for esoteric under¬
takings. My advice is to leave the Psathyrellas to professional psathyrellologists, who
are paid to wrestle with such matters. A mere seven species are described here.
Key to Psathyrella
l. Growing either in sand dunes or on burnt ground or burnt debris or on other mushrooms . 2
1. Not as above; growing in grass, humus, wood chips, on wood, dung, etc.5
2. Growing on shaggy manes (Coprinus comatusf northern in distribution .P. epimyces
2. Not as above . 3
3. Growing in sand dunes or sand, often barely poking above ground . P. ammophila & others
3. Growing on burnt ground or debris . 4
4. Cap with whitish fibrils when young (but these soon wearing away) . . . P. carbonicola, p. 366
4. Not as above . 5
5. Stalk usually with a membranous, well-defined ring (annulus) after veil breaks .
.P. longistriata & others, p. 362
5. Not as above; ring absent or fibrillose and evanescent . 6
6. Cap (2) 5-10 cm broad and distinctly fibrillose or fibrillose-scaly, at least when young, yellow-
brown to rusty-brown or sometimes dark brown to blackish-brown; stalk typically at least
5 mm thick .P. velutina & others, p. 366
6. Not as above . 7
7. Cap distinctly striate or pleated nearly to the center; usually growing in groups or clusters . . .
.(see Coprinus, p. 342)
7. Not as above; if growing in clusters then not striate or striate only near margin when moist 8
8. Clustered on ground or wood chips, each cluster arising from a buried “tap root”; stalks usually
at least twice as long as widths of caps .P. multipedata (see P. hydrophila, p. 364)
8. Not as above . 9
362 COPRINACEAE

9. Typically growing in clusters on or near wood, stumps, etc. 10

9. Not as above . 13
10. Spore print distinctly reddish-tinted P. sublateritia& P. conissans (see P. hydrophila, p. 364)
10. Spore print some shade of gray, brown, or black . 11
11. Veil present when young, but often disappearing in age . 12
11. Veil absent .P. spadicea (see P. hydrophila, p. 364)
12. Cap yellowish to honey-colored when moist, fading to buff or whitish as it dries; especially
common on lawns and in towns.P. candolleana & others, p. 363
12. Cap rusty-brown to cinnamon-brown when fresh, fading to tan as it dries; found mainly in the
woods . P. hydrophila, p. 364
13. Stalk 0.8-1.5 cm thick, white; cap 4-10 cm broad, rounded becomingconvex or even plane, at first
pallid from silky fibrils, but dark gray to grayish-brown in age; found under aspen in the Rocky
Mountains, oak in California (see photo at bottom of p. 363) .P. uliginicola
13. Not as above; stalk generally less than 1 cm thick . 14
14. Growing in grass, dung, or manure . 15
14. Not as above (may occasionally grow in grassy places, but not associated with grass) .... 17
15. Growing in dung; not common .P. stercoraria & others
15. Growing in grass; common . 16
16. Cap yellowish or honey-colored when moist, fading to buff or whitish as it dries; veil present
when young but often disappearing in age .P. candolleana & others, p. 363
16. Not as above .(see Panaeolus, p. 353)
17. Odor typically fruity or pungent (like root beer, cat urine, etc.); cap often radially wrinkled
and striate; cap and/ or stalk often vinaceous- or purple-tinged.P. bipellis
17. Not as above . 18
18. Veil present when young (covering the gills when very young), usually leaving remnants on
or near the cap margin . 19
18. Veil absent or rudimentary and quickly disappearing . 20
19. Cap soon becoming broadly convex to plane, honey-colored or paler when moist, fading to
buff or whitish as it dries out .P. candolleana & others, p. 363
19. Not as above; cap remaining broadly conical to bell-shaped or becoming convex only in old
age and/ or differently colored .P. longipes group & others, p. 364
20. Cap white or whitish .P. sp. (unidentified) (see P. carbonicola, p. 366)
20. Cap not white; stalk usually at least twice as long as cap width P. gracilis group & others, p. 365

Psathyrella longistriata (Ringed Psathyrella)

CAP 2.5-8 (10) cm broad, conical to convex becoming broadly convex to plane or
umbonate in age; surface smooth or slightly wrinkled, with whitish veil fibrils when very
young, not viscid; some shade of brown, but sometimes fading in age. Flesh thin, fragile.
GILLS adnate to adnexed or seceding, close, pallid becoming dark brown or purple-
brown. STALK 4-10 cm long, 0.4-1 cm thick, equal, pallid or white, fragile, hollow, with
scattered whitish scales (veil remnants) below ring. VEIL membranous, white, forming a
persistent, superior, usually striate ring on stalk which is eventually darkened by spores.
SPORE PRINT deep brown to nearly black; spores 7-9 * 4-5 microns, elliptical, smooth.
HABITAT: Solitary to scattered or in small groups in woods, particularly under conifers;
fairly common in the Pacific Northwest, less so in California—I have found it but a few
times in our area, in the winter.
EDIBILITY: Unknown.
COMMENTS: The persistent ring on the stalk is unusual for a Psathyrella, and helps to
distinguish this western species. Panaeolus semiovatus and Stropharia semiglobata are
somewhat similar but grow in dung or grass, not in the woods. Another Psathyrella with
a prominent ring, P. kauffmanii, occurs under hardwoods in eastern North America, and
is quite common in the aspen forests of the Rocky Mountains and Southwest.
Psathyrella candolleana is a common suburban species with a convex to plane or wavy cap and very
fragile flesh. The cap is usually honey-brown to yellowish when fresh but fades as it dries. Note the
dark spore dust on cap at upper right and the slight veil remnants on margins of caps at left.

Psathyrella candolleana (Suburban Psathyrella)

CAP 2-7 (10) cm broad, bluntly conical or convex when young, becoming broadly convex
to plane or broadly umbonate in age; surface smooth or with a few scattered patches of
whitish fibrils (veil remnants) when young; hygrophanous but not viscid; brown, honey-
colored, or yellowish when moist, fading quickly as it dries to buff or whitish (the
center often remaining darker); margin often hung with veil remnants, at least when
young. Flesh very thin, fragile. GILLS close, adnate but sometimes seceding, at first
whitish, soon grayish or grayish-purple, finally dark brown. STALK 4-10 cm long, 2-7
(10) mm thick, equal, hollow, fragile, white or whitish, often silky or scurfy. VEIL white,
usually disappearing, but sometimes forming a fibrillose ring on stalk. SPORE PRINT
deep brown; spores 7-10 x 4-5 microns, elliptical, smooth.
HABITAT: Scattered to gregarious or tufted in Jawns, gardens, on or about old hardwood
stumps, on buried roots or debris, etc.; widely distributed and very common in urban
and suburban settings, sometimes also in the woods. It fruits year-round in our area but
is most common in the late spring, summer, and early fall.
EDIBILITY: Edible. Some authors describe it as delicious, but it is very fragile, insub¬
stantial, and not particularly easy to identify. Its main asset is its easy availability.
COMMENTS: The fragile, usually convex cap that is more or less honey-colored when
fresh and pallid in age, plus the fragile white stalk, deep brown spores, and fondness for
suburbia mark this ubiquitous but boring mushroom. The cap color is so variable that it is
more of a hindrance to identification than a help. The situation is complicated by the
presence of some very similar suburbanites, including P. hymenocephala, with a more
cinnamon-colored cap when young, and P. incerta (formerly Hypholoma incertum and
often listed as a synonym for P. candolleana), with smaller spores and a pale yellowish cap.
All of these are so fragile that getting them home in one piece is difficult!

Left: Psathyrella uliginicola or a similar species (see couplet #13 on p. 362) occurs under oak in Cali¬
fornia; it is unusually robust for a Psathyrella, with a convex to plane cap that is striate in old age.
Right: A cluster of Psathyrella candolleana with clearly visible universal veil remnants. (Greg Wright)
Psathyrella hydrophila commonly grows in clusters on wood. Left: A large clump growing from an
old (buried) stump. Right: Close-up showing the white stalk, dark gills, and fragile veil remnants
on margin of cap.

Psathyrella hydrophila (Clustered Psathyrella)

CAP 2-5 (7) cm broad, bluntly conical or more often convex when young, expanding to
nearly plane in age; surface smooth, hygrophanous but not viscid, dark reddish-brown
to orange-brown or rusty-brown when moist, fading as it dries to pale tan or sometimes
grayish; margin often darker brown and/or hung with veil remnants. Flesh thin, fragile.
GILLS crowded, buff to pale brown, becoming chocolate-brown or dark brown in age;
adnate to adnexed. STALK 3-7 (10) cm long, 2-6 (10) mm thick, equal, hollow, fragile,
smooth, white to faintly grayish or sometimes brownish in old age, especially below. VEIL
fibrillose, evanescent, leaving remnants on cap margin and occasionally an obscure zone
of hairs on stalk. SPORE PRINT deep brown; spores 4-6 x 3-4 microns, elliptical, smooth.
HABITAT: Gregarious, usually in tufts or large, dense clusters on hardwood stumps, logs,
and buried wood; widely distributed. Common in our area from fall through early spring.
EDIBILITY: Not recommended. Reportedly harmless, butsomecollectionsarebitterand
closely related species haven’t been tested.
COMMENTS: This is one of several Psathyrellas that grow in attractive, often large
clumps on decaying wood. The principal fieldmarks are the white or pallid stalk, dark
brown spores, fragile texture, and smooth, hygrophanous, usually convex cap. It is likely
to be mistaken for a Naematoloma, but is much more fragile and not as brightly colored,
or for a Galerina or Pholiota, which have paler brown spores. Other clustered Psathy¬
rellas include: P. spadicea, similar, but with a thicker (4-10 mm) stalk and no veil, growing
mainly on Populus (poplar, cottonwood, aspen); P. fuscofolia, also lacking a veil, but
with stalk only 2-4 mm thick, on decaying wood; P. circellatipes, also slender-stemmed,
with spores 12-15 microns long, favoring aspen; P. multipedata, with clusters originating
from a common, deeply rooted base or “pseudorhiza”; P. sublateritia, with a reddish-
hued spore print (rather unusual for a Psathyrella); and P. conissans, with a pinkish-red
spore print (highly unusual for a Psathyrella).

Psathyrella longipes group

CAP 2-7 cm broad, broadly conical or bell-shaped, expanding slightly in age (rarely plane);
surface smooth, hygrophanous, brown to cinnamon-brown or yellow-brown when moist,
fading to tan, buff, or whitish as it dries; margin often adorned with widely spaced, tooth¬
like veil remnants. Flesh thin, fragile. GILLS close, adnate but often seceding, at first
whitish, soon darkening to brown, finally dark brown to blackish. STALK 6-16 cm long,
2-8 mm thick, equal or enlarged slightly at base, fragile, hollow, white. VEIL white,

364
Left: Psathyrella longipes group, mature specimens. Note relatively large size (fora Psathyrella) and
widely spaced veil fragments on margin of cap. Right: Psathyrella gracilis, or one of its numerous
look-alikes. Note hygrophanous bell-shaped cap and long, slender, fragile stem.

somewhat membranous, but soon disappearing except for remnants on margin of cap.
SPORE PRINT deep brown to blackish; spores 10-15 x 6.5-9 microns, elliptical, smooth.
HABITAT: Solitary, scattered, or in small groups on ground or debris in woods; found
mostly in the West. Fairly common in our area from fall through early spring, under both
hardwoods and conifers.
EDIBILITY: Unknown.
COMMENTS: The broadly conical to bell-shaped cap, long white fragile stalk, small veil
remnants on the cap margin, relatively large size (for a Psathyrella), and dark brown to
almost black spores are characteristic, but there is a multiplicity of Psathyrella look-alikes
that are best identified with a microscope. In the “true” P. longipes, the spores are slightly
flattened in end view; in the very similar but more common P. elwhaensis, they are not.
Other species include: P. atrofolia, widely distributed and common, cap honey-brown
fading to buff, with veil remnants on the margin (but not widely spaced or toothlike) and
spores 8-10 microns long; P. subnuda, slightly larger, with a more cinnamon-colored
convex cap when moist and practically no veil; andP. conopilea, with a reddish-brown to
orange-brown conical cap that fades to buff, and spores 14-18 microns long, especially
common in southern California under oaks and in gardens, but widely distributed. All of
the above species tend to grow scattered to gregarious on the ground or in lignin-rich
humus, rather than in clusters on wood. They do not grow in dung or manure like
Panaeolus. See also P. gracilis, which lacks a veil.

Psathyrella gracilis group (Graceful Psathyrella)

CAP 1.5 A (5) cm broad, conical to bell-shaped becoming convex or nearly plane; surface
smooth, not viscid, brown to dull yellowish-brown when moist, paler (or with a pinkish
tinge) as it dries, and sometimes grayish from spore dust; translucent-striate when moist.
Flesh very thin, fragile. GILLS close, broad, adnate but often seceding, pallid becoming
grayish to brown and then dark brown. ST ALK 6-12 cm long, 1 -3 mm thick, more or less
equal, straight, fragile, thin, white or stained darker by spores. VEIL absent or rudimen¬
tary (and quickly disappearing). SPORE PRINT dark purple-brown to nearly black;
spores 10-15 x 6-8 microns, elliptical, smooth.
HABITAT: Scattered or in groups or troops onground, debris, and wood chips in woods,
parks, under trees, etc.; common and widely distributed. I have seen colossal fruitings on
wood chip mulch in a park at regular intervals throughout the mushroom season.
366 COPRINACEAE

EDIBILITY: Too thin and fragile to be worthwhile.

COMMENTS: This species lacks the veil remnants on the cap margin characteristic of
the P. longipes group. The long, thin, straight, whitish stalk, brown to grayish cap that is
so thin as to be translucent-striate when moist, absence of a veil, dark spores, and terrestrial
growth habit are the main fieldmarks. There are many similar Psathyrellas which can only
be differentiated microscopically.

Psathyrella carbonicola (Charcoal Psathyrella)

CAP 1.5-6 cm broad, bluntly conical to convex; surface dry, at first covered with a dense
coating of whitish fibrils, these eventually wearing away to reveal the chocolate-brown to
brown background; fading in age. Flesh rather thin. GILLS close, adnate but sometimes
seceding, pale brown becoming dark brown in age. STALK 3-7 cm long, 2-6 mm thick,
more or less equal, rather fragile, sheathed with white fibrils or scales below the veil, at
least when young; base often brownish in age. VEIL white, evanescent or forming a slight
fibrillose superior ring on stalk. SPORE PRINT dark brown; spores 6-8 * 3-4 microns,
elliptical, smooth.
HABITAT: Scattered to densely gregarious or clustered on charred soil and wood,
common after forest fires in northern and western North America. In our area I have seen
large fruitings in the winters following controlled burns. It is often accompanied by Pho-
liota highlandensis, P. brunnescens, and Myxomphalia maura.
EDIBILITY: Unknown.
COMMENTS: The whitish fibrillose coating on the cap and stalk when young plus the
growth on burnt ground make this one of the easiest of all Psathyrellas to identify. Other
species: P. canoceps is one of several small species with a coating of silky white fibrils on
the cap, but it does not grow in burned areas; a small, unidentified, whitish-capped
species with black spores is common in our area in woods and under trees.

Psathyrella velutina
CAP (2) 5-10 cm broad, obtuse to convex or broadly umbonate, becoming nearly plane;
surface dry, densely fibrillose or fibrillose-scaly but sometimes nearly smooth in age; dull
yellow-brown to tawny to rusty-brown or sometimes darker brown; margin often paler,
splitting in age, and hung with veil remnants. Flesh rather thick, brownish to ochre. GILLS
crowded, adnate to notched or seceding, pale yellowish becoming light brown to rusty-
brown, finally deep brown as spores mature; faces usually mottled in age, edges white and
sometimes beaded with droplets. STALK 5-15 cm long, (0.3) 0.5-1.5 (2) cm thick, equal or
swollen at base, fibrillose or scaly, dry, whitish above, light brown to dingy tawny or ochre
below. VEIL fibrillose-cottony, usually forming an obscure superior hairy or cottony ring
or zone on stalk which is darkened by spores. SPORE PRINT blackish-brown; spores
8-12 * 5-8 microns, elliptical, minutely roughened, with a prominent snoutlike germ pore.
HABITAT: Solitary or in groups or small clusters in grassy places, roadsides, around
sawdust and compost piles, on gravelly ground, or sometimes in the woods; widely dis¬
tributed, but infrequent in our area. I have found it in the fall and spring.
EDIBILITY: Edible, but not recommended.
COMMENTS: Also known as Lacrymaria velutina, this species is unusually large and
sturdy for a Psathyrella, and is likely to be looked for in another genus. The fibrillose to
fibrillose-scaly cap and stalk, obscure hairy annulus (ring), and blackish-brown spores
are distinctive. P. lacrymabunda is often listed as a synonym, but has smooth spores
according to Smith. Both species are sometimes placed in a separate genus, Lacrymaria.
Other species: P. maculata is somewhat smaller, has blackish- to grayish-brown fibrillose
patches on the cap, and grows in clumps on alder in the Pacific Northwest. Also see
P. uliginicola (couplet #13 of the key) and P. carbonicola.
 367

oj
STROPHARIACEAE
THIS is a fairly common family of saprophytic mushrooms with brown to purple-brown
to purple-black spores and attached gills. A veil is usually present, but does not necessarily
form an annulus (ring) on the stalk. The gills are not normally decurrent as in Gomphidius
and Chroogomphus, nor are they usually free as in Agaricus, nor do they deliquesce as in
Coprinus. The small, fragile species resemble Psathyrella and Panaeolus, but tend to have a
viscid and/ or brightly colored cap. (For a comparison of the brown-spored species with
genera in the Cortinariaceae, see comments under the genus Pholiota.) The mushrooms in
this family also share several anatomical (microscopic) characteristics: the cap cuticle is
usually filamentous rather than cellular (see p. 19), the spores are smooth and often have a
germ pore, and the gills frequently feature special sterile cells (chrysocystidia) which have a
highly refractive golden content when mounted in potassium hydroxide (KOH).
Four genera are recognized here, all of which intergrade to some extent: Pholiota has dull
brown to rusty-brown spores and is consequently placed in the Cortinariaceae by some
mycologists; Stropharia, Psilocybe, and Naematoloma have deep brown to purplish
or black spores, and are sometimes lumped together (by the “lumpers,” of course) in a single
giant genus, Psilocybe. The latter three genera are differentiated largely on microscopic
characteristics such as the presence or absence of chrysocystidia, but can usually be told in
the field by the combination of characteristics outlined in the key.
This is not an important family from a gastronomic standpoint. However, it is the most
significant group for “magic mushroom” hunters, because Psilocybe is the principal genus
of hallucinogenic or “pupil-dilating” mushrooms. The active principles are psilocybin
and psilocin (see p. 895 for details, and also read comments on pp. 31-32).

Key to the Strophariaceae

1. Spore print dull brown to cinnamon-brown or rusty-brown .Pholiota, p. 384
1. Spore print purple-brown to purple-gray, purple-black, or black . 2
2. Lower portion of stalk and /or other parts of fruiting body staining blue or green when handled
(sometimes slowly) .Psilocybe, p. 368
2. Not as above (but cap may be blue or blue-green to begin with) . 3
3. Growing on dung or manure . 4
3. Not as above (but may grow in grass) . 6
4. Cap white to yellow, yellow-brown, or pale tan . 5
4. Cap darker (orange-brown to reddish-brown, grayish-brown, dark brown) Psilocybe, p. 368
5. Spore print black; cap white to buff or very pale tan.(see Panaeolus, p. 353)
5. Spore print purple-brown to purple-black; cap usually yellowish or darker Stropharia, p. 374
6. Veil membranous or cottony-membranous, usually forming a distinct ring (annulus) on stalk
.Stropharia, p. 374
6. Veil absent, or if present then fibrillose and disappearing or merely forming a fibrillose zone
on stalk which may subsequently be darkened by falling spores . 7
7. Cap small (usually less than 4 cm broad), viscid when moist, some shade of brown, gray, dull
olive, buff, or if whitish then usually narrowly conical or bell-shaped .... Psilocybe, p. 368
7. Not as above; cap white or brightly colored (yellow, red, green, etc.), or if dull-colored then
not viscid, even when wet; cap usually 2 cm broad or more . 8
8. Cap viscid or slimy when moist; veil present at least when young; growing solitary to scattered
or gregarious, but not usually clustered .Stropharia, p. 374
8. Cap usually not viscid; veil absent or present; often (but not always!) growing in tufts or clusters
.Naematoloma, p. 381
368 STROPHARIACEAE

PSILOCYBE
Small to medium-sized, saprophytic mushrooms found in a variety of habitats. CAP smooth, with
a viscid (when moist), often separable pellicle (skin); usually some shade of brown, gray,
yellow-brown, or buff GILLS typically attached, dark at maturity. STALK usually slender, often
turning blue or green when handled. VEIL often present, but not usually forming a distinct annulus
(ring) on stalk (except P. cubensis). VOLVA absent. SPORE PRINT purple-gray to purple-brown
to nearly black. Spores mostly elliptical, smooth, with a germ pore. Chrysocystidia typically absent
on the gills. Cap cuticle filamentous.

PSILOCYBE enjoys a notoriety grossly disproportionate to its visibility, for it embraces

some of the most exalted and sought-after of all mushrooms—as well as some of the most
mundane. The exalted ones are the hallucinogenic (“pupil-dilating”) species popularly
known as “magic mushrooms.” They contain psilocybin and/or psilocin, and cause
startling changes in one’s perceptions and sensations if consumed in sufficient quantity.
The changes are similar to those provoked by LSD (see p. 895). The mundane ones are
those that don’t contain psilocybin or psilocin—and since most Psilocybe species don’t
contain those compounds, it follows that most Psilocybes are mundane.
The Psilocybes as a group are difficult to characterize: the majority are listless little
brown mushroom (“LBM’s”) with a viscid cap (when moist) and dark (purplish to
nearly black) spores. The hallucinogenic species usually turn blue or greenish when
bruised, especially on the stem, but almost any “LBM” can be mistaken carelessly for a
Psilocybe—with potentially disastrous results! A good spore print is crucial, as it will
eliminate the brown-spored genera (Galerina, Inocybe, Conocybe, etcj, which contain
many poisonous species. Among the dark-spored genera, Coprinus has deliquescing gills,
Psathyrella typically has a non-viscid cap and never stains blue; Panaeolus species with
a viscid cap grow on dung and have black spores; and Naematoloma and Stropharia
species are usually brightly colored, while the cap color in Psilocybe (with the notable ex¬
ception of P. cubensis) is typically some shade of brown, gray, or buff.
Contrary to popular belief, Psilocybes do not grow exclusively on that brown stuff that
sounds like a bell. Rather, they occur in a wide variety of habitats: in grass, on wood chips
and mulch in landscaped areas, on decaying wood, and in humus or beds of moss in forests
and bogs. The hallucinogenic species are particularly abundant in two disparate locales:
the Pacific Northwest and southern Mexico. In our area, alas, they are like solar eclipses—
seemingly rare, though actually more common than any one person’s experience would
indicate. In other words, they are not something you can really look for. It is more a matter
of geography—being in the right place at the right time. (This situation may change, how¬
ever, as introduced species like P. cyanescens spread.)
Since it was discovered that Native Americans near Oaxaca, Mexico, used certain
mushrooms to induce altered states of consciousness, Psilocybes have received an inor¬
dinate amount of attention in the North American press. Underground newspapers and
magazines are full of frivolous articles on “getting off,” there is a glut of “magic mush¬
room” field guides and cultivation manuals available, and, as so often happens, it has be¬
come difficult to sort fact from fiction. Those wishing to pursue the subject should read the
comments on pp. 31 -32 and then invest in a responsible book on “psilocybin” mushrooms
(e.g., Psilocybe Mushrooms and Their Allies by Paul Stamets), and use it in conjunction
with a general field guide.
Whatever you do, don’t rely on shortcuts and don’t sample Psilocybes indiscriminately!
True, most of the hallucinogenic species exhibit a blueing reaction when bruised, but the
“optimum” dosage varies greatly from species to species, and as with any hallucinogenic
drug, there is no way to predict the specific effects it will have on you. Furthermore, I have
PSILOCYBE 369

seen people mistake the staining reactions of other mushrooms (for instance, the black¬
ening of the flesh in Hygrocybe conica) for the blueing reaction in Psilocybe. All the more
reason to develop a systematic knowledge of mushrooms’ habits and characteristics
before venturing into the realm of the “LBM’s.”
Psilocybe is a fairly large and difficult genus. Only a few species are “pupil-dilating,”
and those that aren’t are too small or too rare to be of food value. Five species are
described here and several others are keyed out.

Key to Psilocybe
1. Some part of fruiting body aging or bruising blue to green (check cap, veil, stalk base, flesh) 2
1. Not aging or bruising blue or green . 8
2. Fruiting in warm, muggy weather; mainly tropical and subtropical . 3
2. Fruiting in cool or cold weather; mainly temperate. 4
3. Growing on or near dung or manure; cap whitish to yellowish or yellow-brown .
.P. cubensis, p. 373
3. Not as above .P. caerulescens & others (see P. cubensis, p. 373)
4. Cap narrowly conical to bell-shaped and not expanding much, usually less than 2.5 cm broad;
veil absent or rudimentary . 5
4. Not as above; cap usually expanding, at least in age; veil present, at least when young .... 6
5. Growing mostly in grass .P. semilanceata, p. 370
5. Growing mostly under conifers .P. pelliculosa & others (see P. semilanceata, p. 370)
6. Growing on hardwood logs and debris in eastern North America; blueing only very slowly when
bruised .P. caerulipes {see P. stuntzii, p. 372)
6. Not as above; found mainly on west coast . 7
7. Most parts of fruiting body staining distinctly blue to blue-green when bruised; veil usually
disappearing or forming only a very slight ring on stalk .P. cyanescens & others, p. 371
7. Fruiting body blueing only slightly if at all; veil often forming a distinct (but small and fragile)
ring on stalk that is sometimes greenish- or bluish-tinged . P. stuntzii, p. 372
8. Growing on dung or manure . 9
8. Not as above (but may grow in grass) . 10
9. Veil absent or rudimentary.P. coprophila & others, p. 370
9. Veil present, often forming a slight fibrillose ring on stalk P. merdaria(see P. coprophila, p. 370)
10. Cap convex to plane, 1.5-4 cm broad, dark reddish-brown to brown (but often paler toward
margin or fading overall in age to pale tan); stalk 2-4 mm thick, usually with white mycelium
at base; typically growing in dense groups or clusters on lawns or in other disturbed areas . .
.P. castanella (-P. californica)
10. Not as above; cap differently shaped and/or habitat or color different . 11
11. Veil present, usually forming a small annulus (ring) on stalk . P. stuntzii, p. 372
11. Not as above; veil absent or if present, usually disappearing . 12
12. Cap typically conical or bell-shaped and scarcely expanding; usually growing in grassy areas
.P. semilanceata, p. 370
12. Cap differently shaped and/or growing in moss or wet places . 13
13. Growing in woods or grass in eastern North America (especially the South); cap dark brown to
rusty-brown, fading to ochre or yellow-brown, bell-shaped to convex .
.(set Naematoloma ericaeum under N. dispersum, p. 384)
13. Not as above . 14
14. Cap tawny to dull orange-brown; base of stalk with hairs; found in wet areas, especially common
in the Pacific Northwest .P. (-Galerina) corneipes
14. Not as above . 15
15. Growing in swamps or bogs; stalk very long (5 cm or more) and thin; cap reddish-brown to
blackish-brown .P. atrobrunnea
15. Growing in moss; stalk usually less than 6 cm long; cap dark reddish-brown to tawny or yellow-
brown .P. montana
Psilocybe coprophila is the most common and widespread member of its genus. Note small size and
growth on dung. Cap is brown to red- or orange-brown and viscid when moist.

Psilocybe coprophila (Meadow Muffin Mushroom)

CAP 0.5-2 (3) cm broad, hemispherical to convex or broadly bell-shaped, or at times nearly
plane in age; surface viscid when moist, then dry, smooth or with minute white particles or
patches at margin (when young), dark reddish-brown to cinnamon-brown, brown, or
orange-brown, fading to tan or sometimes grayish-brown as it dries (or darker from spore
dust); margin striate when moist. Flesh thin, brownish. GILLS adnate to slightly de¬
current, fairly well-spaced, grayish-brown becoming deep purple-brown or black. STALK
1-4 cm long, 1-4 mm thick, more or less equal, fibrillose, pallid to yellowish or brown,
darkening in age but not bruising blue. VEIL absent or rudimentary and evanescent.
SPORE PRINT purplish-brown to nearly black; spores 11-14 * 6.5-8.5 microns,elliptical,
smooth. Chrysocystidia absent on gills.
HABITAT: Solitary or in small colonies on dung and manure (especially cow patties);
widely distributed. It is one of our most common dung fungi, fruiting whenever it is damp
enough. It can be “raised” by bringing home a “meadow muffin” and keeping it moist.
EDIBILITY: Generally regarded as harmless, but some strains apparently contain enough
psilocybin to be rewarded with the euphemistic label “active.” A large number would be
needed to produce any noticeable effects, however.
COMMENTS: The small size, viscid brownish cap, purple-brown to dark brown spore
print, and absence of an annulus (ring) distinguish this diminutive dung addict. Also found
on dung is P. (-Stropharia) merdaria, which is similar, but has a dingy yellow to orange-
brown or cinnamon-brown cap, and a veil which often forms a fibrillose annulus (ring)
on the stalk. It is common also, and widely distributed. Other species: P. angustispora is
a minute conical brown inhabitant of the Pacific Northwest; it grows on the dung of elk,
sheep, etc.

Psilocybe semilanceata (Liberty Cap)

CAP 0.5-2.5 cm broad and high, narrowly conical to bell-shaped with a pointed umbo,
scarcely expanding in age; surface smooth, chesnut-brown to brown or olive-brown and
at least slightly viscid when moist, fading to tan, olive-buff, or even yellowish as it dries;
margin sometimes with bluish or olive stains. Flesh very thin, pallid. GILLS adnate to
adnexed or seceding, pallid, soon becoming gray, then finally dark purple-brown or
chocolate-brown; edges whitish. STALK 3-10 cm long, 1-2 (3) mm thick, equal, often

370
PSILOCYBE 371

curved or sinuous, pliant, whitish or with brownish base, sometimes with a bluish or blue-
green tinge in age, especially at base. VEIL absent or rudimentary. SPORE PRINT purple-
brown; spores 11-14 * 7-9 microns, elliptical, smooth. Chrysocystidia absent on gills.
HABITAT: Widely scattered to gregarious in pastures, tall grass, etc., but not on dung;
widely distributed. It is especially common west of the Cascades from northern California
to British Columbia. It fruits from late summer through early winter or sometimes in the
spring. I have not found it in our area.

EDIBILITY: Hallucinogenic (see p. 895), and often gathered for recreational use despite
its small size. It is not as potent as P. cyanescens, but is much stronger than P. pelliculosa.
COMMENTS: The liberty cap is one of the most distinctive “pupil-dilating” Psilocybes.
The sharply conical or “peaked” cap, dark spore print, small size, and growth in grass
(often tall) make it distinct. The flesh and stalk bruise blue only slightly, if at all, but may age
olive or slightly bluish. Another “liberty cap,” P. pelliculosa, is often mistaken for P. semi-
lanceata. It is common under conifers in the Pacific Northwest and northern California.
In addition to its different habitat, it is not quite as conical and has a more pronounced
tendency to bruise or age blue-green. It is only mildly hallucinogenic, with 20-40 caps
constituting an “average” dose. A third species, P. silvatica, has spores less than 10 microns
long; it also grows under conifers and occurs across the northern half of the continent.
There are also a number of small to minute, more or less conical, brown orgray Psilocybes
(e.g., P. montana) that do not contain psilocybin. They grow mostly in bogs or in beds of
moss and are very difficult to distinguish (see key to Psilocybe). All of the species discussed
above, including the liberty cap, resemble Panaeolus species, but have a viscid pellicle
(skin) that peels easily from the cap, slightly paler spores, and do not grow in dung.

Psilocybe cyanescens (Potent Psilocybe) Color Plate 88

CAP 1.5-4 (5) cm broad, soon convex to broadly convex, then plane or with an uplifted,
often wavy margin; surface smooth, viscid when moist, dark brown or reddish-brown
becoming caramel-brown, then fading as it dries to tan, yellowish-brown, or paler;
sometimes with blue or blue-green stains, especially near margin. Flesh thin, bruising blue
or blue-green. GILLS typically adnate but sometimes seceding, fairly close, brown or
cinnamon-brown becoming dark smoky-brown or sometimes bluish-stained; edges
whitish. STALK 3-8 cm long, 2-6 (8) mm thick, equal or with an enlarged base, sometimes
curved; dry, whitish, but staining blue to bluish-green when handled or bruised. VEIL
fibrillose or cobwebby, copious but disappearing or at most forming a very slight ring
or hairy zone on stalk. SPORE PRINT purple-brown to purple-gray or purple-black;
spores 9-12 * 5-9 microns, elliptical, smooth. Chrysocystidia absent on gills.

HABITAT: Widely scattered to densely gregarious on wood chips, sawdust, mulch, and
humus, and on lawns rich in lignin; partial to coniferous debris, but also fond of alder
and eucalyptus. It is fairly common in the San Francisco Bay area in cold weather
(December-February), especially in landscaped areas and mulched flower beds, and is
also fairly common in Oregon, Washington, and British Columbia. It is easily cultivated,
and its aggressive mycelium responds readily to transplanting—given the proper
conditions.
EDIBILITY: Hallucinogenic (see p. 895) and extremely potent, especially raw. Along with
P. baeocystis and P. strictipes (see comments), it is the most powerful known hallucino¬
genic mushroom in the north temperate zone. Only one or two caps are needed to induce
marked changes in perception and sensation. Psilocin is primarily responsible, with
psilocybin and possibly other compounds contributing to the effects. Together they are
said to constitute 0.6% of the mushroom on a dry weight basis—substantially more than the
372 STROPHARIACEAE

better known P. cubensis. A six-year-old Washington boy died after ingesting an unknown
quantity of P. baeocystis along with other unidentified mushrooms. However, effects on
adults other than those typical of psilocybin- and psilocin-ingestion have not been
reported.
COMMENTS: The dark spores, viscid caramel-brown cap that fades as it dries, evanescent
veil, and blueing of the stalk and flesh are the fallible fieldmarks of this potent Psilocybe.
There are two very similar, closely-related, equally potent species on the west coast: P.
baeocystis, with a less copious veil and more conical cap that is usually olive-brown when
young; and P. strictipes, with a long, slender stalk (10-13 cm long, 2-3 mm thick). They may
well occur in California, but earlier reports of P. baeocystis from San Francisco were
apparently based on P. cyanescens. Spore prints should be taken to distinguish all three of
these species from deadly Galerinas and other “LBM’s.”

Psilocybe stuntzii (Stuntz’s Blue Legs)

CAP 1-4 (5) cm broad, bluntly conical becoming convex to broadly umbonate, plane,
or with an uplifted margin in age; surface smooth, viscid when moist, color variable: deep
olive-brown to chestnut-brown when young, but often fading as it ages or dries to dingy
yellow-brown or yellowish-buff; margin striate when moist and often tinged greenish.
Flesh thin, pallid to brownish. GILLS adnate or adnexed, pallid soon becoming grayish
or brownish; close or fairly well-spaced. STALK 2-6 cm long, 1.5-4 mm thick, equal or
thicker at either end, often curved, yellowish to brown or sometimes with darker or bluish
stains, especially below; not viscid, often with mycelial threads at base. VEIL membranous
but thin; forming a fragile ring or fibrillose zone on stalk which is often blue or bluish-
green but eventually may be darkened by falling spores or may disappear. SPORE PRINT
dark purple-brown; spores 8-12 * 6-8 microns, elliptical, smooth. Chrysocystidia absent
on gills.
HABITAT: Scattered to densely gregarious or clustered on wood chips, mulch, etc., in
lawns, gardens, and landscaped areas; also under conifers and in fields. It is known only
from the west coast and is especially common in the Puget Sound region of Washington in
the fall, early winter, and spring. (The appearance of large numbers in wood chip mulch on

Psilocybe stuntzii growing in a flower pot mulched with fir bark. Note the annulus (ring) on stalk
and the viscid cap (at least when moist). The white globules are fertilizer pellets.
PSILOCYBE 373

the University of Washington campus in Seattle more than a decade ago helped spur the
“magic mushroom” craze that subsequently swept the Pacific Northwest.) In our area I
have found it in mulched flower pots.
EDIBILITY: Weakly hallucinogenic (see p. 895), but popular with “magic mushroom”
hunters because it often fruits in large numbers. Be sure not to confuse it with deadly
Galerina species, which can look quite similar, but have rusty-brown spores—they will
even grow intermixed with P. stuntzii!
COMMENTS: Like the liberty cap (P. semilanceata), this “LBM” contains psilocybin,
but bruises blue only weakly if at all. However, the dark spore print and ring on the stalk
plus the viscid cap when moist are distinctive. It is named after the late, great Dr. Daniel
Stuntz of the University of Washington, who was the first person to collect it—or more
precisely, the first to collect specimens for the herbarium rather than for consumption!
The common name, which was coined by Gary Lincoff, is apparently a reference to the
tendency of the stalk or “leg” to stain bluish, though it often won’t stain. Other species: P.
caerulipes of eastern North America also blues only slightly if at all; it has a more or less
evanescent veil and grows on hardwood logs and debris. Like P. stuntzii, it is weakly
hallucinogenic.

Psilocybe cubensis (Magic Mushroom)

CAP 1.5-8(10) cm broad, broadly conical or oval or bell-shaped (often with an umbo) when
young, gradually expanding to convex, broadly umbonate, or plane; surface smooth or
with small whitish veil remnants when young, viscid when moist, soon dry, color variable:
whitish with a brown to yellowish center, or entirely yellow to yellowish-buff to yellow-
brown, or sometimes cinnamon-brown when young and sometimes dingy olive in old age;
bruising and aging bluish; margin sometimes hung with veil remnants. Flesh firm, white,
staining blue or blue-green when bruised. GILLS close, adnate to adnexed or seceding to
free; pallid, soon becoming gray, then deep purple-gray to nearly black; edges whitish.
STALK 4-15 cm long, 0.4-1.5 cm thick, equal or more often thicker below, dry, white or
sometimes yellowish to yellow-brown, aging or bruising blue or blue-green; smooth. VEIL
membranous, white or bluish-stained, usually forming a thin, fragile, superior ring on stalk
which is blackened by falling spores. SPORE PRINT dark purple-brown to blackish;
spores 11-17 * 7-12 microns, elliptical, smooth, thick-walled, with a large apical germ pore.
Cystidia present on faces of gills, but chrysocystidia absent.
HABITAT: Solitary or in groups on dung and manure, especially in cattle pastures; widely
distributed in the tropics and subtropics—Colombia, Central America, Mexico, etc.—and
in the Gulf Coast region of the United States. Since it is being cultivated on a widespread
basis, it may eventually turn up in the warmer parts of California—as it did in Santa Cruz
during muggy summer weather (on compost from a “magic mushroom” farm). It can be
cultivated on a variety of simple grain or compost mediums, but strict temperature control
and sterile conditions are necessary to induce growth and prevent contamination.
EDIBILITY: Hallucinogenic (see p. 895). It is not as powerful on a dry weight basis as
P. cyanescens, but is larger. At one time the demand for “magic mushrooms” far out¬
stripped the supply, with the result that many of the “psilocybin” mushrooms sold on the
street were actually grocery store mushrooms (Agaricus bisporus)laced with LSD or other
substances (one sample I examined proved to be a soggy chanterelle soaked inammonia!).
Now that cultivation procedures have been refined, you’re much more likely to get the real
thing. It is often sold dried, though drying decreases the potency. The mycelium is also
“active.” Ironically, natives of Oaxaca, Mexico, consider this species inferior to others,
such as P. mexicana (perhaps because it was unknown to them until the Spaniards intro¬
duced cattle). Instead of using it themselves, they sell it to eager gringos!
Psilocybe cubensis is best told by its shape, whitish to golden-brown cap color, and tendency to stain
blue when handled, especially on the stalk. It is common on dung in tropical and subtropical regions.

COMMENTS: This is the largest, handsomest, and best known of all the “psilocybin”
mushrooms. The brownish to yellow or pallid cap, membranous ring on the stem, and
tendency to bruise blue are the main fieldmarks—plus its growth in fields, usually on cow
patties. It resembles the yellow-capped Stropharias (e.g., S. semiglobata) so closely that
it is called Stropharia cubensis by many mycologists. The Stropharias, however, do not
stain blue. In shape it resembles Agrocybe praecox, but the spore print is darker, and it
has a decidedly different “aura”—or so I’m told by at least one certified fruitcake! Other
species. The “Landslide Mushroom,”/5, caerulescens, is another common, blue-staining,
hallucinogenic species that occurs along the Gulf Coast as well as in Mexico. It has an
evanescent veil, bitter taste, and an olive-black cap when young that may become redder
(e.g., reddish-brown) in age. As its name implies, it grows in recent landslides, as wellas on
sugar cane mulch and other debris. P. tampanensis, discovered in Florida, is a slender¬
stemmed hallucinogenic species that forms 1-2” underground “tubers.”

STROPHARIA
Small to medium-sized, saprophytic mushrooms. CAP usually viscid when moist and often brightly
colored; typically convex to plane or umbonate. GILLS typically attached, dark brown to gray,
purple-gray, or black at maturity. STALK often fleshy, but sometimes slender; central. VEIL
present, usually forming an annulus (ring) on stalk. VOLVA absent. SPORE PRINT deep brown
to purple-brown, purple-black, or black. Spores typically elliptical and smooth, with a germ pore.
Chrysocystidia usually present on gills. Cap cuticle filamentous. Acanthocytes usually present in
mycelium.

THIS is a medium-sized genus of brightly colored mushrooms with a well-developed,

persistent veil, attached gills, and dark (purple-brown to black) spores. The cap is
usually viscid and some shade of yellow, yellow-brown, orange, red, green, blue, or white.
In most cases the veil is membranous and forms a distinct annulus (ring) on the stalk, but
in some species, such as S. semiglobata, it forms only a fibrillose zone (as in Naematoloma
and Psilocybe), and in others, e.g., S. ambigua, it leaves copious remnants on the cap
margin. An interesting and apparently unique microscopic feature of Stropharia is the
presence of acanthocytes (needlelike calcium oxalate crystals) in the mycelium of many
—if not all—species.
Stropharia is most apt to be mistaken for Agaricus, which has chocolate-brown spores
and free gills which are frequently pink when young, and for Agrocybe, which has a

374
STROPHARIA 375

browner (never purple-brown) spore print and dry to only slightly viscid cap. Stropharia
intergrades to some extent with Psilocybe, but in the latter (as defined here) either the stalk
bruises bluish-green or the veil does not form an annulus or the cap is brownish to buff.
Most Stropharia species are found in humus, grass, or dung, but a few grow on decayed
wood or wood chips. Woodland species can be mistaken for Naematoloma, but as a rule
they rarely grow in the clusters typical of that genus, and their veil is more persistent.
Many Stropharias are attractive, but only the large and distinctive S. rugoso-annulata
is commonly eaten. Some species may actually be poisonous, though there are conflicting
opinions on this point. Seven representatives of the genus are described here.

Key to Stropharia
1. Cap brick-red to reddish to orange and small (typically less than 6 cm broad) .
. (see Naematoloma aurantiaca Sc others, p. 382)
1. Differently colored and/or larger . 2
2. Cap and/or stalk blue to green when fresh (or partially so), but often fading or developing
yellowish tones in age .S. aeruginosa Sc others, p. 380
2. Not as above . 3
3. Growing in dung, manure, grass, compost, or mulched or landscaped areas . 4
3. Not as above; usually growing in woods . 9
4. Veil membranous, usually forming an annulus (ring) that is often striate or grooved (on upper
surface); stalk generally 1-2 cm thick or if thinner then usually rather short . 5
4. Not as above; stalk usually at least 5 cm long and slender . 7
5. Cap 4-15 cm or more broad, wine-red to reddish-brown to tan, yellow-brown, or even grayish-
brown; stalk at least 1 cm thick .S. rugoso-annulata, p. 378
5. Not as above; usually smaller and more slender; cap white to yellowish or yellow-brown . . 6
6. Cap white or tinged yellowish to ochre at center . . S. melanosperma (see S. coronilla, p. 377)
6. Cap golden-brown to yellowish or creamy ..S. coronilla, p. 377
7. Stalk viscid or slimy below the veil, at least when moist .S. semiglobata Sc others, p. 376
7. Stalk not viscid . 8
8. Cap smooth (without scales) and white to yellowish when fresh, and usually less than 6 cm broad;
stalk neither cottony nor scaly .S. umbonatescens Sc others (see S. semiglobata, p. 376)
8. Not as above . 9
9. Cap scaly or fibrillose and not viscid; stalk thick (at least 1 cm) and scaly below the ring
(annulus) . S. kauffmanii, p. 380
9. Not as above . 10
10. Cap small (less than 5 cm broad), without scales, chestnut-brown to olive-brown or olive-gray
when moist, sometimes fading to yellow-brown or buff as it dries; stalk only 2-4 mm thick;
annulus (ring) often greenish- or bluish-tinged .(see Psilocybe, p. 368)
10. Not as above . 11
11. Stalk slender, brownish, with small scales below the annulus (ring); cap tawny-orange to yellow¬
ish to brown or olive-brown, small (less than 8 cm broad), often umbonate and with small,
often concentrically arranged scales (but these sometimes washed off); growing on rotten
wood, sawdust, debris, etc.; especially common in the Pacific Northwest .
.S', squamosa (see S. kauffmanii, p. 380)
11. Not as above . 12
12. Cap typically some shade of yellow or cream; veil typically shredding, most of it remaining on
cap margin or forming only a slight annulus (ring) on stalk . 13
12. Not as above; veil usually forming a large annulus on stalk . 14
13. Stalk usually at least 0.6 cm thick, often cottony or shaggy; found in woods S. ambigua, p. 377
13. Stalk more slender (less than 1 cm thick), not shaggy or only slightly so; growing in a wide
variety of habitats .S. riparia (see S. ambigua, p. 377)
14. Cap yellowish, often with darker (brownish) spots; found under hardwoods and in lawns and
other open places in eastern U.S. (especially the South) . . S. hardii(see S', coronilla, p. 377)
14. Cap dull brown or shaded with gray, purple, etc.; found in northern North America, usually
under conifers .S. hornemannii, p. 379
Stropharia semiglobata. Left: Small specimens. Right: A larger one. It resembles Psilocybe cubensis,
but does not stain blue. The veil does not always form a distinct ring on stalk. Note long slender stem.

Stropharia semiglobata (Hemispherical Stropharia)

CAP 1-5 (6) cm broad, hemispherical (rounded) or broadly bell-shaped, becoming
convex or rarely plane; surface smooth, viscid or slimy when moist, pale yellow to straw
colored, yellowish-buff, or yellow-brown; margin often paler, sometimes hung with
whitish veil remnants. Flesh pale or watery yellowish, thin. GILLS typically adnate but
sometimes seceding, at first grayish, then dark purple-brown to black. STALK 5-8 (13) cm
long, 2-6 mm thick, typically long and slender, equal or slightly enlarged at base; somewhat
fibrillose above the veil, viscid or slimy below (when moist); white to yellowish. VEIL slimy,
delicate, forming a fragile, superior, fibrillose ring or zone on stalk which is soon blackened
by falling spores, or sometimes disappearing entirely. SPORE PRINT dark purple-brown
to black; spores 15-19 x 7.5-10 microns, elliptical, smooth. Chrysocystidia present ongills.
HABITAT: Solitary or in small groups on dung, manure, rich soil, straw, and grazed or
fertilized grass; widely distributed, and one of our most common dung fungi. In our area it
fruits whenever it is damp, often in the company of Psilocybe coprophila and the Pan-
aeolus campanulatus group.
EDIBILITY: Edible, but slimy and mediocre according to most sources. As usual, how¬
ever, Captain Charles Mcllvaine disagrees, stating that, “the caps are equal to any
mushroom—tender, good, and harmless.”
COMMENTS: Our most common and uninteresting Stropharia, this species is sometimes
mistaken for the considerably more interesting Psilocybe cubensis, which has a more
prominent ring and stains blue or green when bruised. The viscid to slimy yellowish cap,
slender viscid or slimy stem, and dark gills set S. semiglobata apart. The veil may form a
slight ring, but is not as membranous as that of S. coronilla. There are several closely
related species or variants which are difficult to distinguish, including: S. stercoraria, with
cap usually plane at maturity, stalk not viscid or only slightly so, and spores larger; S.
siccipes, with a dry, sometimes rooting stalk and smaller spores; S. (-Psilocybe) umbona-
tescens, with a conical to distinctly umbonate cap, dry stalk, and similarly-sized spores;
and Psilocybe {-Stropharia) merdaria (see comments under P. coprophila). All of these are
partial to dung, manured ground, and grass, while S. albonitens, another similar species,
grows on the ground in a variety of habitats. It has a whitish or yellow-tinged, often umbo¬
nate cap and a membranous but thin or evanescent veil. Agrocybe species are also similar
in color and shape, but have browner gills and a brighter or browner spore print. See also
S. riparia (under S. ambigua).
Stropharia coronilla, various stages of development. This grass-inhabiting species looks like a small
Agaricus, but has attached gills and a striate or grooved annulus (ring). Note relatively short stalk.

Stropharia coronilla (Garland Stropharia)

CAP 2-6 cm broad, convex to plane or slightly uplifted in age; surface usually smooth (but
in one form with small orangeish scales), slightly viscid when moist, golden-brown to
yellowish, yellowish-buff, or creamy. Flesh soft, white. GILLS adnate or at times adnexed
in age, close, pallid becoming grayish, then purplish or purple-gray to purple-black.
STALK 2-5 cm long, 3-6(10) mm thick, usually rather short, more or less equal, not viscid;
whitish, minutely scaly or cottony above the ring, fibrillose to smooth below; base often
with white mycelial threads. VEIL membranous, white, forming a persistent, median to
superior ring on stalk; ring usually striate or grooved on upper surface and soon darkened
by spores (but in one form not striate). SPORE PRINT dark purple-brown to blackish;
spores 7-11 * 4-5.5 microns, elliptical, smooth. Chrysocystidia present on gills.
HABITAT: Scattered to gregarious on lawns and baseball fields, also in pastures and other
grassy areas; widely distributed but not particularly common, at least in our area. It fruits
most any time but is most frequent in the fall.
EDIBILITY: Dubious—poisonous according to some; hardly worth experimenting with.
COMMENTS: This attractive little Stropharia is often mistaken for an Agaricus, but has
attached rather than free gills and a frequently grooved or lined (striate) ring. It is shorter
than S. semiglobata, the ring is more prominent, and the stalk is not viscid. Forms with a
deeply grooved ring have been called S. bilamellata. A similar, widespread species with a
paler (whitish or tinged creamy to ochre at center) cap, S. melanosperma, grows on dung
or in pastures. S. hardii is a slightly larger (cap up to 10 cm broad) eastern species that grows
in grass as well as in woods. Its veil is more apt to disappear in age and it has smaller spores.
Like S. coronilla, it is easily confused with Agaricus, but has attached rather than free gills.

Stropharia ambigua (Questionable Stropharia) Color Plate 89

CAP 3-15 cm broad, obtuse to convex, becoming plane or even uplifted in age; surface
smooth, viscid or slimy when moist, yellow to yellowish-brown to yellowish-buff, tawny, or
sometimes nearly white; margin hung with cottony white veil remnants. Flesh white, thick,
soft. GILLS pale gray, gradually darkening to purplish-gray or purplish-black; close,
typically adnate but sometimes seceding. STALK 6-18 cm long, 0.5-2 cm thick, more or less
equal, often long; stuffed or hollow; silky and white above the veil, clothed with soft, dry,

377
378 STROPHARIACEAE

delicate, cottony white scales below (but these sometimes wearing off); often yellowish
toward base in age; base often with white mycelial threads attached. VEIL soft, white,
cottony, leaving shreds or strands on the cap margin and sometimes a superior ring or
ragged zone on the stalk. SPORE PRINT dark purplish to nearly black; spores 11-14*6-
7.5 microns, elliptical, smooth. Chrysocystidia present on gills.
HABITAT: Solitary to scattered or in groups in rich humus, usually under conifers, but
also with alder and other hardwoods; known only from the Pacific Coast. It is fairly
common in our area from late fall through early spring, especially in dank, cold places (in
rain forests, along woodland streams and gullies, etc.).
EDIBILITY: Edible ? According to one authority, it tastes “like old leaves.” As I do not
make a habit of chewing on old leaves, I cannot attest to the validity of this comparison.
COMMENTS: There is nothing ambiguous or questionable about this elegant, stately
fungus. It is our most common woodland Stropharia and at its best is one of the most
exquisitely beautiful of all mushrooms—well worth seeking out. The soft, delicate white
scales that sheathe the stem and the strands of veil tissue on the cap margin are very striking.
Amanita gemmata is somewhat similar in color but has white gills; S. hornemannii has a
duller cap and more prominent ring. Other species: S. riparia (=S. magnivelaris?) is a
somewhat similar but slightly smaller and slimmer (stalk less than 1 cm thick) species that is
fairly common in the West in a variety of habitats (under aspen and alder, along streams,
under mountain conifers, etc.). Its veil is thinner than that of S. ambigua and more kleenex-
like, and often leaves remnants on the cap near the margin rather than dangling from the
margin itself. Also, the stalk is not nearly so shaggy as that of S. ambigua, and the cap,
although similar in color, is more apt to be slightly umbonate.

Stropharia rugoso-annulata (Wine-Red Stropharia)

CAP 4-15 (20) cm broad, obtusely bell-shaped or convex becoming broadly umbonate to
plane; surface smooth, slightly viscid or dry, color variable: wine-red to purple-brown
to reddish-brown when fresh, but fading to tan, straw-color, or even grayish as it ages. Flesh
thick, fairly firm, white. GILLS adnate or notched (but sometimes becoming free in age),
crowded, at first whitish but soon gray and finally purple-gray to purple-black with whitish
edges. STALK 7-12 (25) cm long, 1-3 (7) cm thick, often enlarged at base, white or dis¬
coloring yellowish to brownish in age; base often with white mycelial threads attached.

Left: Close-up of the shaggy veil of Stropharia ambigua. Right: Stropharia rugoso-annulata. Cap
color ranges from wine-red to tan. The cultivated version is often much more robust.
STROPHARIA 379

VEIL membranous, white; forming a thick, persistent, superior ring on stalk that is soon
blackened by falling spores; ring grooved or lined (striate) on upper surface and often split
radially into segments. SPORE PRINT deep purple-brown to black; spores 10-15 x 6-9
microns, elliptical, smooth. Chrysocystidia present on gills.
HABITAT: Scattered to gregarious in mulch, wood chips, straw, lawns, gardens, and
other cultivated areas; widely distributed. It is quite common in New England and has
also turned up in Washington. I have yet to find it in our area, but it is bound to turn up
sooner or later (probably sooner).
EDIBILITY: Edible, and the best of the genus for the table. It is easily grown at home and
widely cultivated in Europe (particularly eastern Europe). The flavor is fairly good—
reminiscent of “undercooked potatoes soaked in burgundy,” according to Rick Kerrigan.
COMMENTS: This large, handsome Stropharia is easily recognized by its wine-red to tan
cap, purple-black spores, and grooved, often segmented or clawlike annulus (ring). Its
growth in planted areas suggests that it is an “alien,” but its origin is unknown. The gills are
never pink as in Agaricus, and they are attached to the stalk—at least when young—and the
spores are purple-black rather than chocolate-brown.

Stropharia hornemannii
CAP 4-12(15) cm broad, obtuse to convex, becoming broadly umbonate or plane; surface
viscid or slimy when moist, smooth or with a few whitish scales (veil remnants) near margin;
dull brown to dingy purple-brown, grayish-brown, grayish-purple, or smoky reddish-
brown, often fading in age to yellow-brown or grayish-tan. Flesh thick, soft, white; taste
rather disagreeable. GILLS typically adnate but sometimes seceding, broad, close, pale
gray becoming purple-gray to dull purple-brown to nearly black. STALK 5-15 cm long,
0.5-2.5 cm thick, more or less equal, silky-smooth above the ring, sheathed with soft, dry,
delicate, cottony white scales below, at least when young; base often with white mycelial
threads. VEIL membranous, white, forming a persistent, flaring or skirtlike, superior ring
on stalk which is darkened by falling spores. SPORE PRINT purple-brown to purple-
black; spores 10-14 x 5.5-7 microns, elliptical, smooth. Chrysocystidia present on gills.

Stropharia hornemannii has a shaggy stalk like that of S. ambigua, but its veil is much more mem¬
branous and forms a prominent annulus (ring) on the stalk.
380 STROPHARIACEAE

HABITAT: Solitary, scattered, or in small groups on ground or rotting wood under

conifers; widely distributed in northern North America. Like S. ambigua, it is quite
common in the late summer and fall in the Pacific Northwest and northern California, but
does not seem to range as far south as that species (I have yet to find it in our area). In Mt.
Rainier National Park in Washington the two species can often be found growing together
on the Longmire Trail.
EDIBILITY: Unknown, but not worth experimenting with because of its poor taste.
COMMENTS: This beautiful mushroom resembles S. ambigua, but has a duller (browner
or grayer) cap, and its veil forms a prominent, well-developed annulus (ring) rather than
leaving copious strands on the cap margin. S. depilata is apparently a synonym. Other
species: Pholiota albivelata is somewhat similar, but smaller (cap 4-8 cm broad, pinkish-
brown to vinaceous-brown), with a white stalk that is scurfy or scaly below the mem¬
branous, persistent, striate annulus. It is placed in Pholiota because it has cinnamon-
brown to dark yellow-brown spores; however, it has acanthocytes on the mycelium,
suggesting that, like P. subcaerulea, it belongs in Stropharia. It occurs under conifers
in the Pacific Northwest and northern California, but rarely fruits in large numbers.

Stropharia aeruginosa (Blue-Green Stropharia) Color Plate 91

CAP 2-6 (8) cm broad, broadly bell-shap£d to convex becoming broadly umbonate or
nearly plane; surface viscid when moist, smooth or with a few whitish scales (veil remnants)
near margin; bright green to blue-green when fresh, developing yellow tones in age. Flesh
soft, white or tinged blue. GILLS typically more or less adnate (but may secede), fairly
close, at first pallid but soon grayish, finally purple-brown or chocolate-brown. STALK
3-8 cm long, 3-8 (12) mm thick, more or less equal, smooth and pallid above the ring,
colored like the cap or paler below, and usually with small cottony scales; often slightly
viscid when moist. VEIL membranous, white, soft, forming a fragile, superior ring on stalk
which often disappears in age. SPORE PRINT dark purple-brown to purple-black; spores
6-10 x 4-5 microns, elliptical, smooth. Chrysocystidia present on gills.
HABITAT: Solitary or in small groups in rich soil, humus, woody debris, or sometimes
even in grass; widely distributed. I have not found it in our area, but it is quite common in
southern California, especially in the winter under oak. In the Southwest I’ve collected
it under aspen in the summer, and in the Pacific Northwest I’ve seen it under conifers.
EDIBILITY: Not recommended. It is deemed poisonous by many authors, but is
supposedly eaten in Europe.
COMMENTS: This is one of the most distinctive of all the agarics, easily identified by
its viscid, greenish to blue-green or yellow-green cap and gray to chocolate-brown or
blackish-brown gills. Two similar species occur under conifers in the Pacific Northwest:
S. albocyanea, with a paler (whitish to yellowish) cap and stalk tinged variously with
blue or blue-green; and Pholiota subcaerulea, with a bluish cap (when fresh) and paler
(cinnamon-brown) spores and mature gills.

Stropharia kauffmanii Color Plate 90

CAP 5-15 cm broad, convex to plane; surface not viscid; covered with brown to yellow-
brown or grayish-brown scales on a dull yellowish or tan background; margin some¬
times hung with veil remnants. Flesh thick, white. GILLS adnate or notched, narrow, thin,
close or crowded, pallid becoming gray, then purple-gray to purple-black; edges often
eroded. STALK 6-10 cm long, 1.5-3 cm thick, equal or slightly swollen at base; whitish or
creamy, with erect or recurved fibrillose scales, especially below the ring; base often with
white mycelial threads. VEIL membranous, forming a fragile, white, superior ring on stalk
STROPHARIA 381

which is soon darkened by falling spores, or often disappears in age. SPORE PRINT dark
purple-brown to purple-black; spores 6-8 x 4-4.5 microns, elliptical, smooth, apical germ
pore absent or minute. Chrysocystidia present on gills.
HABITAT: Solitary, scattered, or in groups in rich humus, around brush piles and decayed
woody debris, etc.—usually under hardwoods such as alder, cottonwood, and aspen,
spring through fall. It occurs in northern California and the Pacific Northwest, but in my
experience is more common in the aspen forests of the Southwest and southern Rockies
than anywhere else. There it fruits, like most other mushrooms, in the summer.
EDIBILITY: A tempting mushroom, but I can find no information on it.
COMMENTS: The dry (non-viscid) scaly cap is unusual for a Stropharia, and combines
with the scaly stem and violet-gray gills to distinguish this beautiful species. Its closest
relative is probably Pholiota fulvosquamosa, which differs in having brown gills and
brown spores. Both species are in some respects more typical of Agaricus than of the
Strophariaceae, but have gills attached to the stem (at least when young). Another species
with a scaly stalk, 5. (-Psilocybe) squamosa, has a smaller, viscid cap (see key to Stro¬
pharia). It is widely distributed, but particularly common in the Pacific Northwest.

NAEMATOLOMA
Small to medium-sized mushrooms found mostly on wood. CAP smooth, often brightly colored,
usually not viscid. GILLS attached, dark at maturity. STALK usually slender, more or less central.
VEIL typically present but evanescent or forming only a slight annulus (ring) on stalk. VOLVA
absent. SPORE PRINT usually deep brown to purple-brown, rarely cinnamon-brown. Spores
typically elliptical, with a germ pore. Chrysocystidia present on gills. Cap cuticle filamentous.

THIS is a small group of dark-spored mushrooms with a brick-red to cinnamon-brown,

yellow, greenish-yellow, or orange-red cap. A veil is usually present in young specimens,
but does not normally form a ring, and the cap is not usually viscid, thereby helping to
distinguish it from Stropharia. Psathyrella is often confused with Naematoloma, for
good reason—the two genera were once grouped together in the genus Hypholoma (a name
now used interchangeably with Naematoloma). However, Psathyrellas can be told in the
field by their fragile flesh, white or pallid stalk, and brown to buff or whitish cap, and in the
laboratory by their cellular cap cuticle. Pholiota can also be confused with Naematoloma,
but as defined here has a brown to cinnamon-brown spore print.
Naematolomas are partial to cold weather, but occur throughout the mushroom season.
The common species in our area fruit in tufts or large clusters on decaying wood, but may
appear terrestrial if the wood is buried. Some species, however, fruit in a scattered to
gregarious pattern on the ground, usually in cold northern bogs or under conifers in lignin-
rich humus. Naematolomas may also occasionally turn up on lawns, but none grow on
dung. They are not choice edibles, and at least one, N. fasciculare, is poisonous. Four
species are described here.

Key to Naematoloma
1. Cap more or less brick-red, the margin usually pallid; typically growing in clumps on dead
hardwoods in eastern North America .N. sublateritium (see N. aurantiaca, p. 382)
1. Not as above . 2
2. Cap orange to red or sometimes reddish-brown to brick-red; growing scattered to gregarious
or tufted on ground, in wood chips, mulch, etc.N. aurantiaca & others, p. 382
2. Not as above (but cap may be orange or tawny at center and yellow at margin) . 3
3. Stalk white; fruiting body fragile; cap cuticle cellular .(see Psathyrella, p. 361)
3. Not as above . 4
382 STROPHARIACEAE

4. Typically growing in tufts or clusters on wood (sometimes buried) or roots, or occasionally

densely gregarious on sawdust or wood chips . 5
4. Typically growing widely scattered to gregarious (but not normally clustered) on ground or in
bogs or moss, mainly under conifers .TV. dispersum & others, p. 384
5. Gills yellow to greenish when young . N. fasciculare, below
5. Gills pallid to grayish or grayish-brown when young .TV. capnoides & others, p. 383

Naematoloma aurantiaca (Orange Naematoloma) Color Plate 93

CAP l.5-5.5 cm broad, convex becoming broadly umbonate or plane; surface slightly
viscid or dry, smooth, bright scarlet to red-orange or orange, or at times brick-red to rusty-
reddish to reddish-brown; margin often hung with whitish veil remnants. Flesh pallid, not
bruising blue. GILLS close, pallid or yellowish when young, then grayish-brown or gray¬
ish-olive and finally purple-brown to purple-black in old age; adnate or notched, some¬
times seceding. STALK (2) 3-7 (10) cm long, 2-6 (10) mm thick, equal or with the base
slightly swollen or narrowed; white or tinged yellow above, developing bright orange to
reddish-orange stains over the lower half; base sometimes with white to yellow mycelial
threads. VEIL membranous but very thin, whitish, soon disappearing or forming a slight,
easily-obliterated ring on stalk. SPORE PRINT dark purple-brown; spores 10-14 * 6-9
microns, elliptical, smooth. Chrysocystidia present on faces of gills.
HABITAT: Scattered to gregarious on wood chips, sawdust, and humus rich in lignin,
but often appearing on lawns, in gardens, etc.; distribution uncertain and erratic, but
fairly common in California. It is common in the parks of the San Francisco Bay area. It
also occurs in Los Angeles, and I have found it in Santa Cruz growing from fallen
eucalyptus seed pods. It likes the same habitats as Psilocybe cyanescens, but has a longer
season, fruiting from fall through spring, or even in the summer if it is wet enough.
EDIBILITY: Unknown.
COMMENTS: Also known as Stropharia aurantiaca, this bright, attractive little mush¬
room is easily recognized by its reddish-orange cap, dark spore print, and tendency to grow
in parks or gardens. It is probably an introduced species, but its origin is unknown. Other
species: Stropharia thrausta (-S. squamosa var. thrausta) is a similarly colored species with
a frequently umbonate cap and reddish to orange scaly stem. It is widely distributed (I
have seen it in New Mexico), but does not occur in our area. TV. sublateritium of eastern
North America has a brick-red cap with a paler margin and much smaller spores. It fruits in
clusters on dead hardwoods in the fall and early winter, and is edible and quite common.

Naematoloma fasciculare (Sulfur Tuft) Color Plate 92

CAP (1) 2-5 (9) cm broad, at first broadly conical or bell-shaped, soon becoming convex,
then broadly umbonate to plane; surface smooth, not viscid, bright sulfur-yellow to
greenish-yellow, or at times yellow-orange (especially when young), the center sometimes
darker (orange-tan to orange-brown); margin often hung with small veil remriants. Flesh
thin, yellow; taste very bitter (rarely mild). GILLS close, typically adnate but sometimes
seceding, at first sulfur-yellow, becoming greenish-yellow or olive, then finally dusted
purple-brown to nearly black with spores. STALK 5-12 cm long, 3-10(15) mm thick, equal
or tapering downward, yellow to tawny, but often developing rusty or brownish stains from
base upward; often curved or sinuous, dry, firm. VEIL thin, pale yellow, evanescent, or
leaving slight vestiges on cap margin or an obscure fibrillose zone on stalk which is
subsequently blackened by falling spores. SPORE PRINT purple-brown to deep purple-
gray; spores 6-8 * 3.5-5 microns, elliptical, smooth. Chrysocystidia present on faces of gills.
HABITAT: Gregarious, usually in tufts or dense clusters on decaying wood of both
hardwoods and conifers, but sometimes growing from buried wood or roots and thus
NAEMATOLOMA 383

appearing terrestrial; widely distributed. It is abundant in our area in the fall and winter,
less so in the spring, and is one of the first woodland mushrooms you’re liable to encounter.
EDIBILITY: Poisonous. In Europe and Asia it has caused several deaths; in America only
gastrointestinal upsets have been reported. Fortunately, the bitter taste is a deterrent.
COMMENTS: The yellow to greenish-yellow gills, dark spores, bitter taste, and clustered
growth habit are the principal fieldmarks of this attractive, cosmopolitan mushroom
Hypholoma fasciculare is a synonym. In age the clustered caps often assume a grayish or
purple-brown tinge from a coating of spore dust. N. capnoides is quite similar in cap
color, but its gills are gray to purple-brown (never yellow), and it has a mild taste and grows
only on conifers. Several similar Naematolomas grow on the ground (see N. dispersum).
Other species: N. subviride is a small southern version of N. fasciculare;N. dispersum var.
idahoense has a tawny to cinnamon-brown cap and bitter taste, and grows in clusters on
conifers. For similar brown-spored species, see comments under Pholiotamalicolagroup.

Naematoloma capnoides (Conifer Tuft)

CAP 2-7 cm broad, convex or slightly umbonate to plane; surface smooth, not viscid,
yellow to tawny, orange-brown, rusty-brown, or cinnamon, the margin often yellower and
hung with veil remnants. Flesh thin, pallid; taste mild. GILLS close, usually adnate but
often seceding, at first pallid, then grayish, finally dark gray to purple-brown. STALK5-10
cm long, 3-8 (10) mm thick, equal or tapered downward, dry, slender, pallid or yellowish
above, often rusty-brown to tan or brownish below. VEIL evanescent or leaving small
patches of tissue on cap margin and sometimes an obscure fibrillose zone on stalk. SPORE
PRINT purple-brown to deep purple-gray; spores 6-7.5 * 3.5-4.5 microns, elliptical,
smooth. Chrysocystidia present on faces of gills.
HABITAT: Gregarious, usually in clusters, on rotting conifers; widely distributed, but
especially common in the Pacific Northwest. I have seen it in our area in the fall and winter
on Douglas-fir, but it is not common.
EDIBILITY: Edible, but thin-fleshed and not particularly tasty.
COMMENTS: Also known as Hypholoma capnoides, this nondescript Naematoloma
resembles its ubiquitous relative, N. fasciculare, in cap color, but has a mild taste and
grayish rather than yellow or greenish-yellow immature gills. The stalk is often quite long
in relation to the cap. Several Pholiotas look similar (see Pholiota malicola group), but
have brown or cinnamon spores. Other species: A pale-capped version occurs under
mountain conifers in the spring; it may or may not be a distinct species; N. radicosum of
eastern North America is similar in color but has a deeply rooting base and bitter taste.

Naematoloma capnoides. This slender-stemmed conifer-lover grows in tufts or clusters. It resembles

N. fasciculare (see color plate) but does not have yellow or greenish gills.
384 STROPHARIACEAE

Naematoloma dispersum (Dispersed Naematoloma)

CAP 1-4 cm broad, bell-shaped, but sometimes expanding to convex or even plane with an
umbo; surface smooth, not viscid, tawny to tawny-orange, fading to yellowish; margin
often hung with veil remnants. Flesh thin; taste typically somewhat bitter. GILLS usually
adnate, but sometimes seceding, close, pallid, becoming dingy olive or olive-gray, then
finally purple-brown with paler edges. STALK 6-12 cm long, 2-5 mm thick, equal, usually
long and slender, rather tough and pliant but sometimes also brittle; yellowish above and
brown to dark reddish-brown below. VEIL fibrillose or cobwebby, evanescent or leaving a
fibrillose zone on upper stalk. SPORE PRINT purple-brown; spores 7-10 * 4-5 microns,
elliptical, smooth. Chrysocystidia present on faces of gills.
HABITAT: Widely scattered to gregarious in humus and debris under conifers; wide¬
spread, but particularly common in logged-over areas of the Pacific Northwest from late
summer through early winter. It occurs in northern California, but not in our area.
EDIBILITY: Unknown.
COMMENTS: Also known as Hypholoma dispersum, this is one of several Naemato¬
loma species that characteristically grow in a scattered to gregarious fashion rather than in
clusters. The tendency of the gills to develop olive tints in age is one of its distinctive fea¬
tures, as is its tall slender stalk and purple-brown spore print. It may grow in small tufts, but
not in the large clumps typical of N. fasciculare, and the stem is not white as in Psathyrella.
There are a number of similar Naematolomas that are difficult to distinguish, but of little
interest to the average collector since none are good edibles. Their ranks include: N.
olivaceotinctum, with a greenish-tinted cap (and often gills) in age; N. squalidellum and
A. poly trichi, with yellow gills that become greenish-yellow in age; A. ericaeum, a southern
species with a slightly viscid brown to ochre cap; A. udum, with a browner cap and larger
spores (14-18 microns long), “growing on muck in bogs, especially along rabbit runways”
(A.H. Smith); A. elongatum, with dull cinnamon spores; and A. myosotis, growing in
bogs, with a viscid cap and dull cinnamon spores (the latter two species are also placed in
Pholiota). The above species are not as fragile as the Psathyrellas and do not have white
stems. All tend to grow in groups on troops on the ground rather than in clumps on wood.

PHOLIOTA
Mostly medium-sized mushrooms found on wood or woody debris or sometimes on ground. CAP
usually viscid and/or scaly. GILLS typically adnexed to adnate to slightly decurrent. STALK
central to somewhat off-center but not lateral, usually fleshy but frequently slender; often scaly
below the veil. VEIL present; membranous, slimy, or fibrillose, disappearing or often forming an
annulus (ring) on stalk. VOLV A absent. SPORE PRINT dull brown to cinnamon-brown or some¬
times rusty-brown. Spores mostly elliptical, smooth, usually with a germ pore. Chrysocystidia
often present on faces (sides) of gills. Cap cuticle typically filamentous.

PHOLIOTA is the largest genus of brown-spored, wood-inhabiting agarics. The fruiting

body is usually larger and fleshier than in Galerina and Tubaria, a veil is always present
in young specimens, and the stalk is well-developed and central to somewhat off-center.
The veil can be fibrillose, cottony, slimy, or membranous, and in most cases leaves visible
remains on the stem in the form of an annulus (ring) or a coating of scales over the lower
portion of the stalk.
Among the brown-spored wood-rotters, Crepidotus lacks both a veil and stem, Paxillus
has distinctly decurrent gills and lacks a veil, while Gymnopilus may have a veil but has a
dry cap and brighter (rustier or oranger) spores. Naematoloma and Stropharia intergrade
somewhat with Pholiota, but are traditionally separated by their darker (purple-brown
to blackish) spore color, as is Psathyrella. A few Pholiotas are terrestrial, and these are
PHOLIOTA 385

more difficult to distinguish. In the field they can usually be told from Cortinarius and
Hebeloma by their viscid or slimy cap, presence of a veil, frequently slender build (only
rarely is the stalk bulbous), and sometimes clustered growth habit; microscopically they
are distinct by virtue of their smooth spores.
Many Pholiotas qualify as“LBM’s” and are not likely to interest the average mushroom
hunter. However, some are quite striking (e.g., P. squarrosa and the P. aurivella group).
As few are worth eating and several are mildly poisonous, the genus is best avoided by
beginners. (Some species listed as edible in older books have since been shuffled to genera
such as Rozites and Agrocybe.) In their monograph on North American Pholiotas,
Alexander Smith and L.R. Hesler recognize over 200 species. Since microscopic exami¬
nation is required to determine the shapes or types of sterile cells (cystidia) on the gills,
many of these species cannot be identified in the field. The coniferous forests of the West
are uniquely rich in Pholiota species, but for some reason our area is poorly represented.

Key to Pholiota
1. Growing on recently charred soil or wood (ashes) . 2
1. Not typically growing in ashes . 3
2. Stalk typically 5-10 mm thick .P. brunnescens, p. 393
2. Stalk thinner (2-6 mm) P. highlandensis, P. carbonaria, & others (see P. brunnescens, p. 393)
3. Cap blue to blue-green when fresh (but may discolor yellowish or tan in age); found in the West,
usually under conifers .P. subcaerulea (see Stropharia aeruginosa, p. 380)
3. Not as above (if blue or greenish-blue when young, then found in eastern North America) . 4
4. Growing on wood (occasionally buried) or in wood chip mulch . 5
4. Growing on the ground . 15
5. Cap more or less bright orange-pink when fresh and lacking scales; taste bitter .
... P. astragalina, p. 387
5. Not as above . 6
6. Growing on dead wood of mountain conifers in the spring( when or shortly after the snow melts);
cap and veil viscid to slimy; cap and stalk lacking prominent scales; stalk typically at least
6 mm thick; cap yellowish to rusty-tawny but not brightly colored; known only from the West
.P. sp. (unidentified) (“Snowbank Pholiota”—see photo below)
6. Not growing when the snow is melting, or if so then not as above . 7

Left: These young specimens of Pholiota squarrosoides (see comments under P. squarrosa) are
flagrantly scaly, but older caps are viscid and not as scaly (see color plate).Right “Snowbank
Pholiota.” This unidentified (and probably unnamed) species is common in the northern Sierra
Nevada and Cascades on dead conifers soon after the snow melts. The yellowish-buff to tawny or rusty
cap is viscid or slimy and has few if any scales; the veil is also viscid.
 STROPHARIACEAE

Growing on poplar, cottonwood, aspen, or willow; stalk 1 -4 cm thick at apex, usually thicker
at base, lacking large erect scales (but may have patches of tissue); cap white to creamy or
buff when fresh (but may darken to ochre or brownish in age) .P. destruens, p. 395
Not as above; color or habitat different or stalk thinner and/or scalier . 8
Both cap and stalk yellowish-brown to dark rusty-brown and covered with powdery or
granulose scales when fresh (those on cap may wash off or disintegrate); cap 1-4 cm broad
and dry (not viscid!); stalk slender (1.5^4 mm thick); taste usually bitter or metallic; odor not
distinctive .Phaeomarasmius erinaceellus (-Pholiota erinaceella)
Not with above features . 9
Cap bright yellow to orange, bright tawny, golden, or bright rusty-brown when fresh (but
sometimes darker rusty-brown when very young); cap usually decorated with large scales,
spots, patches, or “straps” of tissue (which may be darker and may wash off) . 10
Not as above; if brightly colored then cap bald or with a few scattered fibrillose veil remnants
(one species tawny to whitish with erect or recurved scales) . 11
Scales on cap and stalk bright yellow; cap dry to slightly viscid .P. flammans, p. 391
Scales usually darker than background color; cap distinctly viscid to slimy when moist, often
shiny in dry weather .P. aurivella group & others, p. 390
Cap and lower stalk becoming reddish-brown to dark vinaceous-brown at maturity (but often
orange-brown when young) and decorated with brown fibrillose scales; cap viscid; usually
solitary or in twos or threes, mainly in eastern North America and the Southwest .
.P. albocrenulata, p. 392
Not with above features . 12
Cap and stalk with prominent erect or recurved scales (which may be obliterated or flattened
somewhat in age); stalk typically less than 1.5 cm thick . 13
Not as above (but cap or stalk may have recurved scales and both may have flattened scales 15
Cap never viscid; gills often (but not always) greenish-tinged in age; odor mild or garlicky;
growing on hardwoods (especially aspen) and conifers.P. squarrosa, p. 389
Cap often dry at first but a viscid layer beneath the scales usually evident in age; gills not greenish-
tinged; odor mild or slightly fruity; on wood or ground . 14
Typically found on hardwood stumps or logs; cap and stalk very scaly when young, the scales
tawny or paler .P. squarrosoides (see P. squarrosa, p. 389)
Usually growing on the ground, less commonly on wood; cap and stalk very scaly to only slightly
so, the scales some shade of brown (darker than in above species) .P. terrestris, p. 389
Growing in deep moss, bogs, or mucky places; stalk equal and usually hollow, long and thin
(length usually at least 40 times the width); cap small, yellow or olive; mostly northern . . 34
Not as above . 16
Lower stalk (i.e., below the veil) with distinct scales (at least in most specimens), the scales some¬
times recurved but often small . 17
Not as above . 19
Cap smooth (without scales) and hygrophanous: rusty-brown to reddish-brown or orange-
brown and striate when moist, fading to yellowish, ochre, or buff as it loses moisture (often
two-toned in intermediate stages); often found in large clusters .P. mutabilis, p. 395
Not as above . 18
Growing on ground or wood chips (often along roads or trails), usually in tufts or clusters; cap
with scales at first (which may wash off or wear away), dark brown to light brown, grayish-
brown, yellow-brown, or tawny; common .P. terrestris, p. 389
Not as above . 19
Cap yellow to bright ochre or orange when fresh . 20
Cap some other color . 23
Cap distinctly viscid when moist . 21
Cap not viscid . 22
Lower portion of stalk (i.e., below the veil) with distinct scales . 23
Not as above .P. malicola group & others, p. 388
Fruiting body staining orange-brown when bruised P. multifolia (see P. malicola group, p. 388)
Not as above .P. malicola group & others, p. 388
Cap viscid or slimy when moist, often with adhering debris when dry, not hygrophanous . 24
Cap dry or hygrophanous or sometimes slightly viscid (but if so, then soon drying out) . . 30
PHOLIOTA 387

24. Veil membranous, typically forming a well-developed annulus (ring) on stalk .

.P. albivelata (see Stropharia hornemannii, p. 379)
24. Not as above; veil typically disappearing or merely forming a slight ring or fibrillose zone 25
25. Cap whitish to pinkish-buff, pinkish-gray, or even grayish P. Ienta(see P. lubrica group, p. 392)
25. Not as above . 26
26. Cap shaded variously with green, olive, purple, brown, and/ or gray whenyoung, oftenyellowto
orange in age; found mainly in eastern North America (especially the South) . P. polychroa
26. Not as above . 27
27. Stalk lacking scales, even when young; cap usually olive-brown to brown or ochre with a yellow
to dingy greenish-yellow margin .P. spumosa, p. 394
27. Not as above; stalk cottony or with small scales below the veil (at least when young and fresh)
and/or differently colored . 28
28. Cap usually with rows of concentrically-arranged veil remnants when young and often ap¬
pearing fibrillose or streaked in age.P. decorata (see P. lubrica group, p. 392)
28. Not as above . 29
29. Veil slimy or viscid in wet weather; cap bright to dark reddish-brown, vinaceous-brown, or
brown and completely smooth (bald) .P. velaglutinosa (see P. lubrica group, p. 392)
29. Veil not slimy or viscid; cap brown to rusty-brown, orangish, etc., the margin often paler or
even whitish . P. lubrica group & others, p. 392
30. Veil usually forming a ring (annulus) on stalk or leaving skinlike fragments on cap margin 31
30. Veil disappearing or leaving only a slight hairy zone on stalk (check several specimens!) . . 32
31. Stalk thick or slender, lacking obvious scales or cottony material below the veil; cap smooth
(bald), usually at least 4 cm broad when mature; white rhizomorphs (mycelial threads) often
present at base of stalk or in surrounding humus; cap cuticle cellular (seeAgrocybe, p. 467)
31. Not as above; fruiting body usually small and stalk usually slender; cap cuticle not cellular
.(see Galerina, Tubaria, & Allies, p. 399)
32. Cap and stalk pale yellow to pale cinnamon or pinkish-cinnamon; cap not translucent-striate
when moist; cap convex to plane or slightly umbonate . . P. scamba{see P. spumosa, p. 394)
32. Not as above . 33
33. Stalk with scattered patches of veil remnants over lower portion; cap convex to plane at maturity,
with an incurved margin when young; gills not decurrent; very common in the spring and early
summer under mountain conifers (on rotten wood or in lignin-rich humus), but also occurring
in other habitats .P. vernalis (see P. mutabilis, p. 395)
33. Not as above .(see Galerina, Tubaria, & Allies, p. 399)
34. Cap viscid when moist, olive or olive-tinged .
.P. myosotis (-Ndematoloma myosotis) (see N. dispersum, p. 384)
34. Cap not viscid, yellow to olive-yellow . .
. P. elongatipes (-Ndematoloma elongatum) (see N. dispersum, p. 384)

Pholiota astragalina (Pinkish-Orange Pholiota)

CAP 2 -5 cm broad, bell-shaped or obtuse becoming convex, umbonate, plane, or with an
uplifted, often wavy margin in old age; surface smooth, viscid or slimy when wet but soon
dry; bright reddish-orange to pinkish-orange, the margin sometimes paler, fading some¬
what in age and often developing blackish discolorations; margin often hung with veil
remnants when young. Flesh thin, orange to yellow; taste bitter. GILLS typically adnexed
or notched or even free, close, bright yellow or yellow-orange, discoloring where bruised or
in age. ST ALK 5-12 cm long, 4-7 mm thick, equal or tapered toward base, fibrillose, with¬
out scales, hollow, sometimes sinuous, pale yellow, the base often oranger or discoloring
brownish. VEIL yellowish, leaving remnants on cap margin or disappearing. SPORE
PRINT brown; spores 5-7 * 3.5-4.5 microns, elliptical, smooth. Chrysocystidia present.
HABITAT: Scattered to gregarious or in small clusters on rotting conifers; widely dis¬
tributed. It is especially common in the Pacific Northwest and northern California in
the late summer and fall; I have not seen it in our area.
EDIBILITY: Inedible because of the bitter taste.
388 STROPHARIACEAE

COMMENTS: The brilliant pinkish-orange cap plus the yellow gills, brown spores, and
growth on conifers make this one of our most distinctive as well as beautiful Pholiotas. It
is reminiscent of Naematoloma, but has paler (browner) spores and a viscid cap when wet.

Pholiota malicola group (Forgettable Pholiota)

CAP 3-8 (15) cm broad, convex becoming plane or slightly umbonate; surface viscid or
dry, smooth or with a few veil remnants at margin, yellow to ochraceous-tawny, orange, or
orange-buff, fading somewhat in age; margin often wavy. Flesh pallid or yellowish; odor
mild or faintly fragrant; taste mild. GILLS close, adnexed or notched, yellowish becoming
rusty-brown or cinnamon-brown. STALK 4-15 cm long, 0.4-1 (2.5) cm thick, equal or
tapered downward or enlarged at base; solid, dry, fibrillose but not scaly; pallid or yellow¬
ish above, darker (tawny or colored like cap) below, becoming rusty-brown in age from
falling spores. VEIL fibrillose, disappearing or forming a slight ring or zone of fibrils near
top of stalk. SPORE PRINT rusty-brown; spores 7.5-11 * 4.5-5.5 microns, elliptical,
smooth, with an apical germ pore. Chrysocystidia absent.
HABITAT: In groups or clusters on rotting logs and stumps, wood chips, etc.; widely
distributed. It is fairly common in our area in the fall and winter.
EDIBILITY: Unknown.
COMMENTS: This forgettable Pholiota is a member of the so-called “P. alnicola com¬
plex”—agroupofdifficult-to-distinguish, scale-less(but not veil-less), yellowish to rusty or
tawny, wood-inhabiting species. They are reminiscent of Naematolomas in their color
and clustered growth habit, but differ in having rusty-brown or cinnamon-brown spores.
They also approach Gymnopilus, but the species in that genus have oranger spores.
Clumps growing in wood chip mulch might be confused with P. terrestris, but that species
has small scales on the stalk below the veil. Closely related and/ or superficially similar
Pholiotas include: P. flavida, with slightly smaller spores and a frequently fragrant or at
least distinctive odor; the “true” P. alnicola, with a mild to bitter taste and yellowish or
olive-tinged cap; P. spinulifera and P. fibrillosipes, with smaller spores, a viscid cap and
conspicuous cystidia on the gill faces, found in groups or clusters in soil, mulch, and saw¬
dust (veil remnants whitish in former, yellow to orange in latter);/*, subochracea, with a
viscid, pale yellow to ochre cap, long (5 cm or more) stem, chrysocystidia, and small
spores (5-6 microns long), found in the Pacific Northwest; P.prolixa, like P. subochracea
but larger-spored, common in eastern North America, often in large clusters in low
hardwood forests or bottomlands; and P. multifolia, also eastern, with a dry yellow cap
and tendency to bruise rusty-brown or orange-brown. None of these are worth eating.
Pholiota malicola group commonly grows in tufts or clusters on dead wood and in wood chip mulch.
These rather small specimens have a yellow cap and practically smooth (scale-less) stalk.
Pholiota terrestris is common on lawns and along roads and trails. It usually grows in tufts or clusters
on the ground, rarely on wood. N ote scaly stalk (the scales can be large or small); spore print is brown.
(Joel Leivick)

Pholiota terrestris (Terrestrial Pholiota)

CAP (1) 2-8 (10) cm broad, obtusely conical or convex becoming plane or somewhat
umbonate; surface usually with dry fibrillose scales, but viscid or slimy in wet weather
beneath the scales, which sometimes wear off; color variable: dark brown to light brown,
grayish- or yellow-brown, or tawny, the scales darker; margin sometimes streaked, often
hung with veil remnants. Flesh white to watery yellow or brown, thin. GILLS attached
(usually adnate), close, at first pallid to grayish, then dull brown to dull cinnamon-brown.
STALK 3-10 (13) cm long, (2) 4-10 mm thick, equal or narrowed below, slender, solid or
becoming hollow, dry, pallid to buff, or brownish toward base; covered with brown scales
or patches below the veil. VEIL fibrillose, whitish, forming a slight superior, fibrillose ring
or zone on stalk, or disappearing. SPORE PRINT brown; spores 4.5-7 * 3.5^4.5 microns,
elliptical, smooth. Chrysocystidia present on gills.
HABIT AT: In groups or clusters on the ground, especially along roads, paths, and in other
disturbed areas; also on lawns, lignin-rich debris, rarely on or around old stumps. It is
widely distributed and especially common along the west coast (including our area)
in the fall and winter. Single individuals are occasionally found, but tufts or clusters are
the rule.
EDIBILITY: Edible but thin-fleshed, insipid, and usually wormy to boot.
COMMENTS: Our most common Pholiota, this is the only veiled, brown-spored mush¬
room that habitually grows in clusters on the ground (several dark-spored Psathyrellas
also do). The scaly stem and evanescent veil are good secondary fieldmarks, but the color,
viscidity, and degree of scaliness exhibited by the cap vary considerably according to age
and weather conditions. The veil does not form a prominent ring as in the white-spored
honey mushroom (Armillariella mellea), but after collapsing it traps falling spores and
turns brown as in Cortinarius. P. squarrosoides (see comments under P. squarrosa) is
very closely related to P. terrestris, but has paler scales and is not terrestrial.

Pholiota squarrosa (Scaly Pholiota) Color Plates 96,97

CAP 3-10 (15) cm broad, obtuse or convex becoming broadly bell-shaped to slightly
umbonate or plane; surface dry, pale tan to straw color, buff, or pale yellow-brown, or in
age darker yellow-brown or sometimes greenish-yellow toward the margin; covered with a
dense layer of upright or recurved, often darker (brown) scales; margin incurved at first

389
390 STROPHARIACEAE

and often fringed copiously with veil remnants. Flesh pale yellowish, rather pliant; odor
mild in some forms, distinctly garlic- or onionlike in others; taste mild or rancid. GILLS
crowded, adnexed to adnate to slightly decurrent, pale yellowish to buff or tinged gray,
then often developing a greenish tinge before finally becoming brown or dull rusty-brown.
STALK 4-12 cm long, 0.5-1.5 cm thick, equal ortapered downward, solid, smooth above
the veil, covered with erect or recurved scales below (like on cap); colored like cap or
becoming darker brown or reddish-brown below. VEIL membranous-fibrillose, forming
a fragile, often torn, superior ring on stalk or only leaving shreds on cap margin. SPORE
PRINT dull rusty-brown; spores 5.5-9 * 3.5-5 microns, elliptical, smooth, with a germ
pore. Chrysocystidia present on gills.
HABITAT: In tufts or dense clusters on wood, usually at the bases of trees (both hard¬
woods and conifers); widely distributed. It is very common on aspen and spruce in the
Rocky Mountains and Southwest during the summer. It is also common on aspen in the
Sierra Nevada in the summer and fall, but I have never seen it on the coast.
EDIBILITY: Not recommended. Some people eat it regularly but others have suffered
severe stomach upsets and old specimens are often rancid-tasting.

COMMENTS: A beautiful and memorable mushroom in its prime, the erect or recurved
scales on the cap and stalk plus the tan to pale tan color and brown spore print set this
species apart. The cap is not viscid as in most Pholiotas and the garlic odor, when present, is
also distinctive. It is usually found on wood, whereas P. terrestris, which is closely related,
typically grows on the ground. Other species: P. squarrosoides (COLOR PLATE 98)
is a very similar and edible scaly species with a somewhat paler (whitish to pale tawny or
yellowish or light brown) cap and mild odor. Its gills are never greenish and in age or wet
weather the cap becomes viscid from a gelatinous layer beneath the scales. It occurs on
hardwoods such as maple and alder, and is much more common than P. squarrosa in
northern California and the Pacific Northwest. In addition to the color plate, a young
cluster is shown on p. 385. Another similar but unidentified species with a very distinctive
citrus fragrance occurs in New Mexico.

Pholiota aurivella group (Golden Pholiota) Color Plate 95

CAP (3) 5-16 cm broad, broadly bell-shaped or convex becoming broadly umbonate or
plane; surface very sticky-gelatinous or slimy when moist (but may dry out), pale to dark
yellow, tawny, golden-orange, or rusty-orange (or sometimes rusty-brown when young),
decoratd with darker scales (the scales large and flattened to slightly recurved and
triangular to strap-shaped or spotlike) that sometimes wear away or wash off in age;
margin often hung with veil remnants when young. Flesh pallid to yellowish, soft in age.
GILLS close, adnate or notched, pallid to yellow becoming brown to rusty-brown or
even brownish-orange in age. STALK 4-15 cm long, 0.4-2.5 cm thick, equal ortapered in
either direction, central or off-center, dry and more or less smooth above the veil, scaly
below (the scales usually not viscid); yellow to pale yellow-brown or colored like the cap
(but often paler). VEIL fibrillose, whitish or yellowish, forming a slight ring or fibrillose
zone on upper stalk or disappearing. SPORE PRINT brown; spores 8.5-10 x 5-6.5 microns
(but see comments!), elliptical, smooth, with a germ pore. Chrysocystidia often present.
HABITAT: Gregarious (often tufted or clustered) on living or dead hardwoods and
conifers; widely distributed. In our area this species“complex” occurs rarely on hardwoods
in the fall and winter, but it is a very prominent fungal feature of the coniferous forests
of the Rocky Mountains, Southwest, Sierra Nevada, and Pacific Northwest, especially in
the summer and fall. In the Southwest I have also seen large fruitings on aspen.
Pholiota aurivella group. Note extremely slimy cap with large scattered scales. Cap color ranges from
yellow to bright tawny- or rusty-orange. These specimens, which have bright rusty caps, were growing
on a dead oak; they are probably P. limonella or P. squarroso-adiposa(see comments below).

EDIBILITY: To be avoided. Some books list it as edible, but many people have suffered
gastric upsets after eating members of this species “complex.” The texture is rather soft
and gelatinous anyway, and it is said to taste “like marshmallows without the sugar.”
COMMENTS: This striking mushroom and its look-alikes (see below) are easily recog¬
nized by their scaly stalk, brown spores, and yellow to orange, viscid cap with large
spotlike darker scales. The honey mushroom (Armillariella mellea) is somewhat similar,
but is not as brightly colored and has white spores. Distinguishing species within the
P. aurivella “complex,” however, is not so easy. Many variants have been described (e.g.,
P. connata, with a thinly viscid stalk and P. abietis, with pale brown immature gills), but
recent cultural studies have revealed the presence of three widespread, non-interbreeding
species which differ principally in spore size: P aurivella, with spores 8.5-10 x 5-6.5
microns, P. limonella (-P. squarroso-adiposa), with spores 6.5-9.5 * 3.5-5.5 microns
and apparently the most common of the three in North America; and P. adiposa, a hard¬
wood-lover with even smaller spores (5-6 * 3-4 microns) and often viscid scales on the
stalk. To complicate matters, there are some other closely related species, including:
P. aurivelloides, with even larger spores than P. aurivella;P. hiemalis, found on northern
conifers usually late in the fall, with viscid scales on the stalk and pallid, yellow-edged gills
when young; and P. filamentosa, with a lemon-yellow to greenish-yellow cap and a thick,
persistent annulus (ring) on the stalk, also found with conifers. Whew!

Pholiota flammans (Flaming Pholiota; Yellow Pholiota)

CAP 3-8 (10) cm broad, convex or obtuse becoming broadly umbonate to nearly plane;
surface brilliant yellow or at times dark yellow or tawny at the center, covered with bright
yellow scales which may wear off in age; dry or in wet weather sometimes viscid beneath
the scales; margin usually fringed with veil remnants. Flesh fairly firm, yellow. GILLS
usually adnexed or notched, bright yellow becoming rustier in age, close. STALK (3)5-10
cm long, 4-10 mm thick, equal or slightly thicker at base, bright yellow or the base slightly
darker; smooth above the veil, sheathed with a dense layer of recurved yellow scales below;
not viscid. VEIL bright yellow, disappearing or forming a slight superior ring or fibrillose-
cottony zone on stalk. SPORE PRINT brown; spores 3-5 x 2-3 microns, oblong to ellip¬
tical, smooth. Chrysocystidia present on gills.

391
392 STROPHARIACEAE

HABITAT: Solitary or tufted on conifer logs and stumps; widely distributed in northern
North America, not common. In California it fruits in the fall and winter, but is rare.
EDIBILITY: Edible, but not choice (see comments on edibility of P. aurivella).
COMMENTS: The brilliant yellow color sets apart this beautiful mushroom. It is most
likely to be confused with the P. aurivella group, but the cap is dry to only slightly viscid and
the scales on the stalk and cap are yellow rather than rusty, cinnamon, or brown. P. adiposa
(see comments under the P. aurivella group) is somewhat similar, but favors hardwoods.

Pholiota albocrenulata
CAP 3-10 (15) cm broad, broadly conical or convex becoming broadly umbonate to nearly
plane; surface viscid or slimy when moist, orange-brown to dark rusty-brown or reddish-
brown, becoming dark vinaceous-brown in age, and adorned with scattered brown fibril-
lose scales (veil remnants); margin often fringed with veil remnants. Flesh thick, whitish;
taste mild or bitter. GILLS close, notched or adnate to slightly decurrent, whitish be¬
coming grayish and finally brown, the edges finely scalloped and white or beaded with
tiny white droplets. STALK 3-10 (15) cm long, 0.5-1.5 cm thick, more or less equal, rather
fibrous, stuffed or hollow; pallid or grayish above, brown to reddish-brown (like cap)
below, with scattered brown scales below the veil; often curved. VEIL fibrillose-cottony,
forming a slight superior fibrillose ring or zone on stalk, or disappearing. SPORE PRINT
brown; spores 10-15 (18) * 5-8 microns, elliptical, smooth.
HABITAT: Solitary or in small groups (twos and threes) on stumps, logs, and living trees,
usually of hardwoods (especially maple and elm); widely distributed, but seldom found
in quantity and apparently absent on the west coast. I have collected it twice in New Mexico
in August—once on a ponderosa pine and once on an aspen.
EDIBILITY: Said to be harmless, but seldom eaten.
COMMENTS: Though not often encountered, this is a striking mushroom by virtue of
its reddish-brown to dark brown, scaly, viscid cap and white-edged gills. Microscopically
it is close to Stropharia, and is placed in that genus by some mycologists (along with
several other Pholiotas, e.g., P. subcaerulea and P. albivelata).

Pholiota lubrica group (Lubricous Pholiota)

CAP 3-10 cm broad, convex becoming plane or sometimes with an uplifted, wavy margin;
surface viscid or very slimy when moist, smooth (but see comments), color variable: dark
reddish-brown to rusty-brown, rusty-orange, or ochraceous-tawny at the center, often
paler (yellowish or even whitish) toward the margin, which is often hung with veil remnants
when young. Flesh fairly thick, whitish to watery yellow or greenish-yellow. GILLS adnate
to adnexed, close; whitish, yellow, or sometimes greenish-yellow, becoming brown or dull
cinnamon as the spores mature. STALK 5-10 cm long, (0.4) 0.8-1.5 cm thick, more or less
equal, smooth above the veil, usually with small scales below which may wear off; white or
yellow, the scales often darker; sometimes brownish-stained in age. VEIL fibrillose to
somewhat membranous, forming a slight superior ring or fibrillose zone on stalk. SPORE
PRINT brown; spores 5.5-7 * 3-4.5 microns, elliptical, smooth. Chrysocystidia absent.
HABITAT: Solitary, scattered, or in groups in humus and on woody debris in woods;
widely distributed. Fairly common in our area in the late fall, winter, and early spring,
especially under pine.
EDIBILITY: Unknown.
Left: Pholiota velaglutinosa has a vinaceous-brown cap and slimy veil. Right: Pholiota ferrugineo-
lutescens has a rusty-orange cap, dry veil, and scaly stalk. Both species are discussed under the
Pholiota lubrica group.

COMMENTS: The above description will more or less fit a large number of Pholiotas with
a viscid-slimy cap and scaly stalk that grow solitary to gregarious (but not often clustered)
in humus or on rotting wood. They are particularly prevalent under conifers but also occur
with hardwoods, and their identification is best left to pholiotologists. The “true” P.
lubrica is said to have a dark brown to reddish-brown cap with a paler or whitish margin.
Some of the other commoner species or variants are: P. ferruginea, with an oranger cap;
P. ferrugineo-lutescens, occasional in our area, with a slightly oranger cap and a thicker
white stem that stains yellow; P. sublubrica, with brownish veil remnants on the cap and
a thicker (1-1.5 cm) stem; P. velaglutinosa, with a bright to dark reddish-brown or brown
cap that lacks veil remnants and a viscid or glutinous veil, sometimes abundant in our
coastal pine forests (see photograph);/*, lenta, with a pale cap (whitish to buff or sometimes
tinged ochre or gray or pinkish) that features whitish veil remnants; andP. decorata, with
a fibrillose-streaked cap that is dark vinaceous-brown to reddish-brown with a paler
margin and has rows of concentrically-arranged veil remnants when young, common in
the Pacific Northwest and Rocky Mountains. Also see P. spumosa, which lacks scales on
stalk, and P. albivelata (under Stropharia hornemannii), which has a membranous ring.

Pholiota brunnescens (Charcoal Pholiota)

CAP 2 -7 cm broad, convex to plane or with an uplifted, often wavy margin; surface viscid
or very slimy when moist, smooth or with small scattered whitish veil remnants;
chestnut-brown to dark reddish-brown to orange-brown, tawny, or dark yellow-brown,
sometimes fading in age to dull orange; margin often paler. Flesh rather soft, dingy
brownish; odor usually mild. GILLS adnate to adnexed, crowded, narrow, whitish or
grayish or pale yellowish, becoming dull cinnamon-brown to browninage. STALK4-6(9)
cm long, (4)7-10 mm thick, equal, whitish to pale yellow, often darkening somewhat below
in age or staining tawny when handled; covered with numerous small yellowish, fibrillose
scales (usually arranged in concentric belts) below the veil. VEIL fibrillose, lemon-yellow,
usually disappearing or forming only a slight superior ring on stalk. SPORE PRINT
brown; spores 6-7 * 4-4.5 microns, elliptical, smooth. Chrysocystidia absent.
HABIT AT: Scattered to densely gregarious or clustered on wood or soil in recently burned
areas; locally common in its favored habitat throughout the mushroom season. It is
known only from the West, but the similar P. highlandensis(see comments) is widespread.

393
Pholiota brunnescens is one of several charcoal-loving Pholiotas. Note slimy cap and the scales
on the stalk. The scales are yellow when young but turn brown as they trap discharged spores.

EDIBILITY: Unknown.
COMMENTS: This is one of several closely-related Pholiotas that fruit only on charred
soil or wood. It is distinguished by its sticky-slimy, tawny to orangish to dark reddish-
brown cap, scaly stem, brown spores, and yellow veil. The other common ash-lovers
have slimmer stems (2-6 mm thick). They include: P. highlandensis (-P. carbonaria of
Europe), widely distributed, with a whitish veil and slightly smaller cap,P. carbonaria, with
a rusty-red to reddish veil; P. fulvozonata, with an orange-brown or russet-colored veil;
and P. subangularis, with a small cap that is only slightly viscid (if at all) and a pallid veil.

Pholiota spumosa (Slender Pholiota)

CAP 2-6 (8) cm broad, obtusely conical or convex becoming umbonate or plane; surface
smooth or appearing fibrillose or streaked, viscid or slimy when moist; color variable but
usually olive-brown when young becoming brown to tawny or tawny-ochre at the center
and yellow to dingy greenish-yellow toward the margin. Flesh yellow or greenish-yellow,
soft, thin; odor mild. GILLS close, adnate or notched, yellow to pale greenish-yellow,
becoming grayish or tawny and finally brown. STALK 3-7 (10) cm long, 4-6 (8) mm thick,
more or less equal, fibrillose but not scaly, yellow to pale greenish-yellow above, becoming
sordid brownish below or from the base upward. VEIL pale yellowish or whitish, delicate,
fibrillose, disappearing or leaving slight remnants on cap margin and stalk. SPORE
PRINT brown or dull rusty-brown; spores6-9 * 4-4.5 microns, elliptical, smooth. Chryso-
cystidia absent.
HABIT AT: Solitary to widely scattered to gregarious or tufted onground and debris under
conifers, widely distributed. Fairly common in our area in the fall and winter under pine,
but rarely fruiting in large numbers.
EDIBILITY: Unknown
COMMENTS: This species belongs to a complex group of Pholiotas with yellow to
greenish-yellow flesh and gills, a viscid to slimy cap, and an evanescent veil. The veil is not
slimy as in P. velaglutinosa, nor is the stem scaly below the veil as in the P. lubrica group.
Other species: P. graveolens is similar but has a strong odor. In our area I have also found
P. scamba, a small species (cap 1.5-3 cm broad) with a whitish to pale yellow or cinnamon-
tinged cap and thin (1 -3 mm) fibrillose or woolly stalk. It grows in groups or small clusters
on dead conifer logs, sticks, etc. Also see comments under the P. lubrica group.

394
PHOLIOTA 395

Pholiota destruens (Destructive Pholiota)

CAP 5-20 cm broad, convex becoming broadly convex or rarely plane; surface slightly
viscid when moist, white to creamy, buff, or at times ochre to brownish, covered with soft
or cottony, whitish to buff scales or patches which may be come matted or washed off in
age; margin often shaggy from veil remnants. Flesh thick, white, firm. GILLS adnate or
notched, close, white becoming dull brown to deep rusty-cinnamon in age. ST ALK (3) 5-15
cm long, 1 -3 cm thick, equal or enlarged below, central or off-center, solid, hard, white, but
often developing brownish stains below in age; smooth above the veil, at first clothed with
whitish to buff scales and patches below. VEIL cottony, white, forming a slight superior
ring on stalk, or disappearing. SPORE PRINT cinnamon-brown; spores 7-9.5 * 4-5.5
microns, elliptical, smooth, with a germ pore. Chrysocystidia absent.
HABIT AT: S olitary or in groups or clusters on dead cottonwood and poplar or sometimes
aspen or willow, especially on the cut ends of logs; widespread. It is particularly com¬
mon in the valleys and bottomlands of the West, where Lombardy poplar and cotton¬
wood are so prevalent. I have seen it in Oregon, New Mexico, and the Sacramento Valley
in California, but not on the coast. It usually fruits late in the season.
EDIBILITY: Edible, but rather tough and poorly-flavored.
COMMENTS: This large Pholiota is distinct by virtue of its pale overall color, soft whitish
veil remnants (scales or patches) on the cap, thick hard stalk, brown spore print, and
occurrence on poplar (or sometimes willow). Its name refers to the fact that it rapidly
destroys the wood on which it feeds.

Pholiota mutabilis (Changeable Pholiota)

CAP 1.5-6 cm broad, obtuse becoming convex, broadly umbonate, or even plane; surface
smooth, lubricous or slightly viscid when wet, hygrophanous: rusty-brown to orange-
brown, reddish-brown, or tawny when moist, fading from the center outward to yellowish-
brown, ochre, or yellowish-buff as it dries (often two-toned: yellowish at center and
browner toward margin); margin translucent-striate only when moist. Flesh thin, white
or tinged brown. GILLS adnate to slightly decurrent, close, pallid soon becoming brown or
dull cinnamon. STALK 3-10 cm long, 2-10(12) mm thick, equal or tapered toward base,
stuffed or hollow, smooth and whitish above the ring, becoming brownish below and
covered with numerous small, often recurved scales (at least when fresh); base sometimes
blackish-brown in age. VEIL whitish, forming a small membranous or fibrillose superior
ring on stalk, or sometimes disappearing. SPORE PRINT cinnamon-brown; spores
5.5-7.5 * 3.5-5 microns, elliptical, smooth, with a germ pore. Chrysocystidia absent.
HABITAT: Typically in clusters—often large—on logs, stumps, or occasionally buried
wood; widely distributed and very common, late summer through early winter, but I have
not seen it in our area. Although it is said to prefer hardwoods, I usually find it on conifers
in northern California and the Pacific Northwest. The fruitings are sometimes so massive
that the substrate (log or stump) is hidden from view.

EDIBILITY: Edible, but not recommended. Experienced collectors sometimes harvest

the large clusters, but it is easily confused with the poisonous Galerina autumnalis and
numerous other “LBM’s” of unknown edibility.
COMMENTS: Also known as Kuehneromyces mutabilis and Galerina mutabilis, this
brown-spored “LBM” is best recognized by it smooth, hygrophanous, often two-toned
cap, scaly stalk with a ring, and penchant for growing in dense clusters. The poisonous
Galerina autumnalis is quite similar, but lacks scales on the stem, has roughened spores,
and does not usually grow in large clusters. Several Naematoloma and Psathyrella species
Pholiota mutabilis is a nondescript “LBM” that usually fruits in clusters. Note how the cap is striate
when moist and becomes two-toned as it loses moisture. Also note how stalk is scaly below the ring.

are also similar but have darker spores. Other species: P. vernalis is closely related and
similar, but lacks scales on the stem (or has only a few patches) and does not grow in such
large clusters. It is common in the mountains of western North America shortly after
the snow melts in the spring, but occurs elsewhere also.

CORTIN ARIACEAE

ALMOST every brown-spored, terrestrial, woodland mushroom you find will belong to
the Cortinariaceae, and many wood-inhabiting ones will also. Like their white-spored
counterparts, the Tricholomataceae, they are a vast, diverse, and baffling group—with
even more species, but fewer genera. The gills are not deeply decurrent and/ or poroid as in
the Paxillaceae, and the cap cuticle is typically filamentous and the spores pore-less, in
contrast to the Bolbitiaceae. The latter characters are microscopic, but with a little practice
the two can be distinguished in the field. The Cortinariaceae are primarily woodland fungi,
a dominant fungal feature of cool temperate forests, whereas the Bolbitiaceae are warm-
weather fungi found mostly in grass, gardens, and dung. Also, with the exception of
Galerina and Tubaria, the Cortinariaceae tend to be larger, fleshier, and/or less fragile
than the Bolbitiaceae.
As applied to the Cortinariaceae, the term“brown-spored” is somewhat misleading. The
spore color actually ranges from orange or bright rusty-orange (Gymnopilus) to rusty-
brown (Cortinarius) to dull brown, ochre-brown, or yellow-brown. Mushrooms with a
filamentous cap cuticle and purple-brown to purple-black spores are traditionally placed
in the Strophariaceae. However, the difference between “brown” and “purple-brown” is
not always clearcut, and the two families seem to intergrade via Pholiota (a brown-spored
member of the Strophariaceae that is keyed out here in the Cortinariaceae) and Galerina.
Another family, the Crepidotaceae, is recognized by some authorities; it is largely tropical
and composed almost entirely of “LBM’s” included here under the Cortinariaceae.

396
CORTINARIACEAE 397

There are very few esteemed edibles in the Cortinariaceae. The gypsy mushroom
(Rozites caperata) is perhaps the best known and most widely collected, but there are many
more species that are definitely known to be poisonous (particularly in Galerina,
Cortinarius, Hebeloma, and Inocybe), and hundreds of others have yet to be tested.
Identification is very difficult in this family, and having access to a microscope does little
to expedite the tedious and labyrinthine identification process, because so little has been
published on the family and so many species are poorly known or still unnamed. Several
representatives from each common genus are included in this book.

Key to the Cortinariaceae

1. Typically growing on wood (may be buried!), wood chips, bark mulch in nurseries, sawdust,
fabrics, or ashes (on burnt ground or wood); neither the flesh nor the gills lilac or violet . . 2
1. Typically growing on the ground (but occasionally on very rotten wood); gills and/ or flesh
violet or lilac in some cases. 8
2. Stalk poorly developed (lateral to off-center) or absent at maturity; on wood or various fabrics
.Crepidotus, p. 405
2. Stalk well-developed, central to somewhat off-center but not lateral; not found on fabrics . 3
3. Spore print rusty-orange to bright rusty-brown; cap not normally viscid or slimy; gills usually
yellow to orange or rusty-orange; stalk fleshy though sometimes slender Gymnopilus, p. 407
3. Not as above; spore print different colored (including duller rusty-brown or ochre-brown) 4
4. Fresh fruiting body small and dark red to wine-colored . . Galerina, Tubaria, & Allies, p. 399
4. Not as above . 5
5. Veil absent . 6
5. Veil present (check young specimens because it may disappear) . 7
6. Gills free at maturity; spore print pinkish-cinnamon .(see Pluteaceae, p. 253)
6. Not as above . 25
7. Stalk fleshy, or if thin then not particularly fragile and sometimes scaly below the veil; cap
usually larger than 5 cm broad (but sometimes smaller), sometimes dry but more often viscid
and usually not translucent-striate when moist; cap sometimes brightly colored, sometimes
growing in large clusters .(see Strophariaceae, p. 367)
7. Stalk thin (less than 5 mm), usually hollow and fragile and without scales; cap small (usually
less than 4 cm) and dry, or if viscid then typically also hygrophanous and translucent-striate
when moist; cap usually dull-colored, not typically growing in large clusters . 25
8. Spore print reddish or with a greenish or olive tinge when moist; gills reddish when young; cap
powdery and small (up to 5 cm broad) and dull grayish to brownish; not common .
. (see Agaricaceae, p. 310)
8. Not as above; very common . 9
9. Membranous veil present when young, usually forming a distinct annulus (ring) on stalk . 10
9. Veil absent, or if present then cobwebby, silky, hairy, or slimy but not membranous . 13
10. Stalk thick and fleshy (at least 8 mm thick, usually more) and/ or gills purplish when young 11
10. Not as above . 12
11. Veil covering the gills well into maturity or tending to shred radially rather than break away from
the cap; fruiting body tending to develop underground (occasionally surfacing in wet weather);
associated with mountain conifers . Cortinarius, p. 417
11. Not as above; developing above the ground .Rozites & Phaeolepiota, p. 411
12. Cap distinctly viscid or slimy when fresh, sometimes with scales; stalk often scaly or cottony or
shaggy below the veil .(see Strophariaceae, p. 367)
12. Cap not viscid or only slightly so and not scaly; stalk not normally cottony, shaggy, or scaly
(but may be fibrillose) .Galerina, Tubaria, & Allies, p. 399
13. Stalk tapered below to form a “tap root” that extends deep into humus; veil absent', spore print
typically rusty-brown to cinnamon-brown .Phaeocollybia, p. 413
13. Not as above; “tap root” absent, or if present then a veil also present( when young) and/ or spores
differently colored . 14
398 CORTINARIACEAE

14. Gills brightly colored (yellow, orange, red, blue, green, purple), at least when young; cap not
typically translucent-striate when moist . 15
14. Gills not brightly colored (but may be dingy yellow, tawny, etc.) and/ or cap translucent-striate
when moist . 17
15. Cobwebby, silky, or hairy veil present when young; cap viscid or dry; spore print rusty-brown
to cinnamon (but not rusty-orange); gills variously colored; common . . Cortinarius, p. 417
15. Veil absent; cap not typically viscid; spore print orange to rusty-orange or dull brown to yellow-
brown or olive-brown; gills yellow to orange or rusty-orange; not very common . 16
16. Spore print orange to rusty-orange or bright rusty-brown .Gymnopilus, p. 407
16. Spore print brown to olive-brown .(see Paxillaceae, p. 476)
17. Volva present at base of stalk; spore print reddish or pinkish-cinnamon (seePluteaceae, p. 253)
17. Not as above; volva absent or spore color browner or rustier . 18
18. Cap typically viscid or slimy when moist, usually smooth (bald) or with a few scattered scales
or veil remnants . 19
18. Cap typically not viscid, but sometimes slightly tackyin wet weather; cap surface smooth or silky
to fibrillose, woolly, scaly, or powdery . 24
19. Stalk distinctly viscid or slimy from the remains of a slimy veil; spore print typically rusty-brown
to cinnamon . Cortinarius, p. 417
19. Not as above . 20
20. Cap usually small (up to 4 cm broad) and striate when moist; stalk thin, i.e., 1-3 (5) mm thick;
fruiting body fragile; often found in moss .Galerina, Tubaria, & Allies, p. 399
20. Not as above . 21
21. Veil present, at least when young (check several specimens if unsure), usually but not always
leaving traces on the stalk in the form of hairs, scales, or an annulus (ring) . 22
21. Veil absent in all stages .Hebeloma, p. 463
22. Spore print typically rusty-brown to cinnamon-brown; veil cobwebby or silky; stalk usually
lacking scales below the veil, usually (but not always) at least 1 cm thick; spores roughened
. Cortinarius, p. 417
22. Not as above ... 23
23. Odor often radishlike or spermatic and/ or taste bitter; gill edges often whitish; stalk apex often
powdery, scurfy, or with small flakes; cap viscid but not often slimy; spore print dull brown
(not cinnamon- or rusty-brown); not typically growing in large clusters from a common base
.Hebeloma, p. 463
23. Not as above; sometimes growing in clusters .(see Strophariaceae, p. 367)
24. Spore print yellow-brown to tan to dull brown, or if rusty-brown or cinnamon then fruiting body
often tawny or yellowish, translucent-striate when moist, and often growing in moss; spores
smooth or roughened . 25
24. Spore print typically rusty-brown to cinnamon-brown; cobwebby or silky veil present when
young, often leaving hairs on stalk; stalk thick or thin but not usually fragile; spores roughened
. Cortinarius, p. 417
25. Odor spermatic or like green corn (crush the cap if unsure!); spore print dull brown or dull yellow-
brown .Inocybe, p. 455
25. Not as above . 26
26. Spore print dull brown or dull yellow-brown; cap usually opaque and small or medium-sized,
often umbonate and/ or with easily-splitting margin (especially in age); surface of cap usually
silky, hairy, or scaly; gill edges often whitish; gills typically not decurrent; very apex of stalk
often minutely powdered, flaky, or scurfy; often growing in groups but not usually in clusters
.Inocybe, p. 455
26. Not as above . 27
27. Veil present at least when very young; stalk with small scales (often somewhat concentrically
arranged) below the veil .(see Strophariaceae, p. 367)
27. Not as above; veil absent or present; stalk not normally scaly, but may have fibrils .28
28. Stalk typically 4 mm or more thick; cap typically 4-10 cm broad .29
28. Not as above (usually smaller) .Galerina, Tubaria, & Allies, p. 399
29. Spore print reddish- or pinkish-cinnamon .(see Entolomataceae, p. 238)
29. Not as above . (see Bolbitiaceae, p. 466)
 399

GALERINA, TUBARIA, & Allies

Small to minute, mostly brown mushrooms found on wood, moss, or ground. CAP typically smooth
or nearly so, often translucent-striate toward margin when moist. GILLS brown to rusty or tawny at
maturity, typically adnexed to adnate (Galerina) or slightly decurrent (Tubaria). STALK thin
(generally less than 5 mm thick), fragile or cartilaginous, usually hollow and central. VEIL absent
or present, sometimes forming an annulus(ring) on stalk. VOLVA absent. SPORE PRINT ochre-
brown to cinnamon-brown or brown. Spores smooth or roughened, usually lacking a germ pore.
Cystidia usually present on gill edges. Cap cuticle usually filamentous.

THIS is an artificial grouping of little brown mushrooms(“LBM’s”) with brownish spores

and a thin, usually fragile stem. Galerina is a large genus( over200 species) formerly divided
among Pholiota and the obsolete genus Galera. It can be recognized in the field by its small
size, yellowish to brown color, and conical to convex cap, plus the thin stem and brown
spores. Pholiota intergrades with Galerina, while Conocybe species also have brown
spores but are usually more conical and have a different type of cell structure in the cap
cuticle (cellular rather than filamentous). Also, Conocybes usually grow in grass, dung, or
cultivated soil, whereas Galerinas are partial to wood, humus, and moss in forests and bogs
(however, some Galerinas grow in grass and some Conocybes in moss).
Tubaria is a small but ubiquitous genus with slightly decurrent gills and a convex to plane
or depressed cap. It is most likely to be confused with Agrocybe, which does not usually
have decurrent gills, and with Pholiota, with which it intergrades. Several other small
genera are also treated or mentioned here(e.g., Naucoria, Alnicola, and Simocybe), but
they are differentiated largely on microscopic characters and are of little interest to the
average mushroom hunter.
Galerina and Tubaria are especially common during dry spells when other mushrooms
are scarce. Several Galerinas are known to contain deadly amanita-toxins. Many others
have not been tested and all are difficult to identify—a compelling reason to avoid eating
all Galerinas, Tubarias, and other “LBM’s.”
Two Galerinas and two Tubarias are described here, plus one rare but distinctive
mushroom of uncertain disposition, Naucoria vinicolor. If your“LBM” does not key out
convincingly, throw it away! For more comments on “LBM’s,” see p. 32.

Key to Galerina, Tubaria, & Allies

1. Fruiting body dark red to vinaceous (wine-colored) when fresh . . Naucoria vinicolor, p. 404
1. Not as above . 2
2. Veil present when young, usually forming a small or large annulus (ring) on the stalk (check
several specimens if possible) . 3
2. Veil absent, or if present then not forming an annulus . 11
3. Annulus prominent, distinctly membranous and often flaring and/or movable; cap dry and not
translucent-striate; cap conical or bell-shaped when young (but may expand), 0.5-2.5 cm
broad; spores with a germ pore, cap cuticle cellular .(see Conocybe, p. 470)
3. Not as above; cap usually striate when moist (at least at margin) and/or differently shaped 4
4. Growing on wood or wood chips. 5
4. Growing on ground (in humus, moss, etc.) . 9
5. Cap thinly to thickly viscid when moist . 6
5. Cap not viscid . 7
6. Cap thinly viscid and translucent-striate when moist; stalk lacking obvious scales below the
annulus (ring); spores roughened .Galerina autumnalis, p. 401
6. Not as above; spores smooth . (seePholiota, p. 384)
7. Cap often with hoary white fibrils or particles when young; gills usually adnate to decurrent
. Tubaria confragosa, p. 403
7. Not as above . 8
400 CORTINARIACEAE

8. Stalk with small or prominent scales or patches of veil remnants below the veil; spores smooth
. (seePholiota, p. 384)
8. Not as above; spores usually roughened .Galerina marginata (see G. autumnalis, p. 401)
9. Cap reddish-cinnamon when moist; often growing on lawns; known only from the Pacific
Northwest .Galerina venenata (see G. autumnalis, p. 401)
9. Not as above; usually growing in moss, grass, or swampy ground and/ or differently colored 10
10. Cap translucent-striate when moist.Galerinapaludosa (see G. autumnalis, p. 401)
10. Not as above . (seeAgrocybe, p. 467)
11. Gills usually slightly decurrent; cap convex to plane or slightly depressed (not conical and
not typically tawny or yellow) . 12
11. Not as above . 13
12. Cap and stalk white to grayish; cap often somewhat hairy, especially toward margin; found
in woods; widespread but not common .(see Clitocybe & Allies, p. 148)
12. Cap brown when fresh but often fading to buff or whitish as it dries; found in many habitats
. Tubaria furfuracea & others, p. 402
13. Cap more or less olive-brown; growing on wood; spores smooth .... Simocybe centunculus
13. Not with above features . 14
14. Stalk or base of stalk blue to blue-green in age; growing in moss or grass (see Conocybe, p. 470)
14. Not as above . 15
15. Cap bright yellow when fresh (but often fading) and conspicuously striate and viscid; entire
fruiting body withering or dissolving quickly...(see Bolbitius, p. 473)
15. Not as above . 16
16. Spore print ochre-brown to pinkish; odor usually strong and fishy or cucumberlike; cap bell¬
shaped to conical when young, reddish-brown to dark brown or blackish (margin often paler);
stalk also dark, minutely velvety; gills with giant cystidia . . . (see Tricholomataceae, p. 129)
16. Not as above . 17
17. Growing on decaying wood; cap hairy or scaly or silky and/ or odor spermatic; cap not viscid
.(seelnocybe, p. 455)
17. Not as above . 18
18. Cap convex to plane or broadly umbonate . 19
18. Cap conical to bell-shaped or acutely umbonate . 24
19. Lower stalk usually with small scales or veil remnants; veil present at least in young specimens;
cap often viscid when moist; spores smooth; usually found on wood, sometimes on ground, but
not usually in moss . (see Pholiota, p. 384)
19. Not as above; spores smooth or roughened; growing in many habitats, often in moss .... 20
20. Cap viscid when moist and conspicuously striate; flesh very thin; stalk white or yellow and very
fragile (2-3 mm thick); entire fruiting body withering or decaying quickly; cap cuticle cellular
.(see Bolbitius, p. 473)
20. Not as above . 21
21. Gills adnate; cap brown but often fading to tan, buff, or even whitish as it loses moisture; spore
print ochre to dull brown or tan; spore walls nearly colorless and thin, easily breaking; common
. Tubaria furfur acea & others, p. 402
21. Not as above . 22
22. Spore print brown; found mostly under alder and willow .23
22. Spore print usually rusty-brown to cinnamon-brown (brighter than above); found in many
habitats. 24
23. Cap dark reddish-brown .Alnicola scolecina
23. Cap brown to tan or yellow-brown .Alnicola melinides, A. escharoides, & others
24. Cap usually yellowish to yellow-brown or tawny or sometimes rusty-brown; veil present or
absent; usually growing in moss or grass or on wood; cap usually translucent-striate when
moist; stalk fragile; spores roughened or smooth . . . Galerina heterocystis & others, p. 402
24. Not as above; sometimes growing in moss, but usually on ground and not normally in grass or
on wood; veil always present (at least when very young); stalk not often fragile; spores
roughened .(see Cortinarius, p. 417)
Galerina autumnalis is a deadly poisonous “LBM.” Growth on wood, brown spores, and scale-less
stalk with a small annulus (ring) are the main features.

Galerina autumnalis (Deadly Galerina)

CAP 1-4 (6.5) cm broad, convex to nearly plane or slightly umbonate; surface smooth,
viscid when moist, dark brown to yellow-brown or tawny, fading to tan or yellowish as it
dries; margin translucent-striate when moist. Flesh thin, watery brown; odor mild or
slightly farinaceous. GILLS attached (slightly decurrent to adnexed) but often seceding,
close, yellowish to pale brown becoming rusty-brown or brown. STALK 2-10 cm long, 3-
6 (10) mm thick, equal or thicker below, dry, hollow, pallid to brownish, often darker
below in age, fibrillose below the veil; base often with white mycelial strands. VEIL fibril-
lose or somewhat membranous, usually forming a thin, superior, white ring on stalk which
is subsequently darkened by falling spores or often disappears in age. SPORE PRINT
rusty-brown; spores 8-11 * 5-6.5 microns, elliptical, roughened and/ or wrinkled.
HABITAT: Scattered to gregarious or tufted on rotting wood and debris of both hard¬
woods and conifers; widely distributed. Fairly common in our area from fall through early
spring, especially during relatively dry years.
EDIBILITY: DEADLY POISONOUS—it contains amanita-toxins! Fortunately, it is
rarely eaten because of its diminutive dimensions and mundane appearance.
COMMENTS: Since this drab “LBM” is deadly poisonous, it is important to learn its dis¬
tinguishing characteristics: (1) the rusty-brown spores (2) the small size and thin stem (but
in areas of high rainfall overgrown specimens often occur) (3) the veil which usually (but
not always!) forms a thin whitish superior annulus (ring) (4) the growth on wood (some¬
times buried or very decayed). The annulus may turn brown in age or even disappear, so
it is best to avoid any mushroom that is remotely similar, including the edible Pholiota
mutabilis, which typically grows in large clusters and has small scales on the stem. There
are several very similar deadly poisonous Galerinas with a thin annulus, including: G.
marginata, with a moist but not viscid cap, found mainly on decaying conifers; and G.
venenata, with a reddish-cinnamon cap that fades to pinkish-buff or whitish in age,
growing on lawns or buried wood and known only from the west coast. Other species: G.
paludosa also has a white superior annulus, but it has a long thin stem and grows in bogs;
its edibility is unknown.

401
402 CORTINARIACEAE

Galerina heterocystis
CAP 5-20 mm broad, bluntly conical becoming bell-shaped or convex or sometimes
umbonate; surface smooth, hygrophanous: pale yellow to pale cinnamon to tawny and
translucent-striate when moist, paler(more or less buff) when dry. Flesh very thin, fragile.
GILLS close, usually attached but not decurrent, pale yellowish becoming pale cinnamon-
brown. STALK 1-8 cm long, 1-3 mm thick, more or less equal, tubular, fragile, pallid to
pale yellowish darkening to brown or cinnamon; lower portion faintly fibrillose. VEIL
absent or rudimentary. SPORE PRINT pale cinnamon-brown; spores 11-17 * 6.5-8.5
microns, more or less elliptical, roughened to nearly smooth.
HABITAT: Scattered to gregarious in damp mossy or grassy places (usually in or near
woods); widely distributed. In our area this species and its numerous look-alikes are
especially common during relatively dry weather when other mushrooms are scarce.
EDIBILITY: Unknown. Do not experiment!
COMMENTS: There are dozens of Galerinas that will more or less fit the above descrip¬
tion, and they can only be differentiated microscopically. Unlike G. autumnalis, the veil is
evanescent or even absent, but like that species the cap is usually translucent-striate when
fresh. Some are found in the woods or at their edges, others frequent seepage areas, still
others grow on logs or in bogs; most are moss-inhabiting. Conocybe species may also
key out here, but are generally more sharply conical, not as translucent, and have a cellular
cap cuticle; they favor lawns, dung, or cultivated ground, but a few grow in moss. Psilo-
cybe and Psathyrella species are also similar, but have darker spores. Other Galerinas
with an evanescent or absent veil include: G. semilanceata, with a fibrillose veil, usually
found on the ground; G. cedretorum, larger (cap 1-3 cm), with a more or less convex cap
and no veil, found in humus or debris under conifers; G. hypnorum, minute, with a rudi¬
mentary veil, found on mossy logs; G. triscopa, also minute, but with a sharply conical
cap when young, growing on logs; and G. tibicystis, growing only in Sphagnum bogs.
None of these should be eaten.

Tubaria furfuracea (T otally T edious T ubaria)

CAP 1-3 (4) cm broad, convex becoming plane or slightly depressed; surface smooth to
finely fibrillose or often with minute whitish flecks and patches (veil remnants); hygro¬
phanous but not viscid, brown to reddish-brown, cinnamon-brown, or tan when moist,
fading to buff, pinkish-buff, or whitish as it dries (often fading in center first); margin striate
when moist. Flesh thin, brownish. GILLS close, adnate to slightly decurrent, pale tawny to
cinnamon or brown. STALK 2-6 cm long, \A mm thick, equal or slightly thicker below,
colored more or less like cap or paler, sometimes with whitish flecks, fibrillose, fragile;
base usually with whitish mycelium. VEIL whitish, fibrillose, evanescent. SPORE PRINT
ochre-brown to pale ochre; spores 6-9 * 4-6 microns, elliptical, smooth.
HABITAT: Scattered to gregarious on ground, sticks, and woody debris in wet places—

Tubaria furfuracea, the “Totally Tedious Tubaria.” Note adnate to slightly decurrent gills and ten¬
dency of cap to fade as it loses moisture. A similar viscid-capped species is also common locally.
Tubaria furfuracea is a quintessential “LBM.” These specimens are taller than most and their gills
are rather widely spaced. Could they be a distinct species? Does anyone know? Does anyone care?

woods, vacant lots, landscaped areas, along trails, etc.; common and widely distributed. It
seems to be most abundant when and where other mushrooms are scarce—perhaps
because it is only likely to be noticed when and where other mushrooms are scarce. I have
seen enormous fruitings in wood chip mulch at Golden Gate Park in San Francisco.
EDIBILITY: Unknown.
COMMENTS: This is it, folks—your quintessential “LBM” (see p. 32)—as boring as it is
ubiquitous and as innocuous as it is inconspicuous. The brownish spores, adnate to
decurrent gills, thin stalk, and whitish-flecked, hygrophanous cap are the most distinctive
(or least undistinctive) fieldmarks. To say more about it would do the more interesting
mushrooms in this book an injustice. Other species: The “Totally Tedious Tubaria” is
easily mistaken for the “Truly Trivial Tubaria,” T. pellucida, which, however, is slightly
smaller (cap 0.5-1.5 cm broad) and has slightly smaller spores. T. tenuis, otherwise known
as the “Truly Trivial AND Totally Tedious Tubaria,” is also smaller, but has a completely
glabrous (bald) cap and widely spaced gills. A small unidentified viscid-capped species
also occurs in our area. There are various other “LBM’s” belonging to obscure genera
such as Simocybe and Alnicola. You can find more information on these in the key to
Galerina, Tubaria, & Allies, but don’t you have something more exciting to do?

Tubaria confragosa (Not S o Tedious Tubaria)

CAP 1-5 cm broad, broadly convex to more or less plane or slightly uplifted; surface moist
or dry but not viscid; hygrophanous, brown to vinaceous-brown or reddish-brown to
reddish-cinnamon when moist, markedly paler (buff to cinnamon-buff) as it dries out;
smooth or often appearing hoary at first from a thin layer of whitish fibrils or minute scales
(especially toward the margin, which is striate when moist). Flesh thin, fragile, colored
like cap. GILLS cinnamon to rusty-brown to reddish-cinnamon to brown, adnate to
slightly decurrent, close. STALK 2-8 cm long, 1.5-6 mm thick, equal or thicker below, soon
hollow, colored like cap or paler, usually with fibrils or a few small scales below the ring;
base typically with white mycelial mat. VEIL usually forminga membranous, oftenflaring
superior ring on stalk, but sometimes disappearing or leaving only a fibrillose zone.
SPORE PRINT brown to dark reddish-cinnamon; spores 6.5-9 * 4-6 microns, broadly
elliptical, smooth.
HABIT AT: Gregarious (often clustered) on rotting logs, sawdust, etc.; widely distributed.
I have seen large fruitings in the fall and winter in wood chip mulch, sometimes accom¬
panied by T.furfur acea.

403
Tubaria confragosa, the “Not So Tedious Tubaria,” is reminiscent of the more common “Totally
Tedious Tubaria” (T. furfuracea), but usually has an annulus (ring) on the stalk.

EDIBILITY: Unknown.
COMMENTS: The presence of a well-defined annulus (ring)—at least in many specimens
—rescues this “LBM” from the obscurity it so richly deserves. It is slightly larger than
T. furfuracea, and is more likely to be cespitose (clustered). It is also known asPhaeo-
marasmius confragosus and Pholiota confragosa, but the hygrophanous, non-viscid cap
and smooth to only slightly scaly stalk separate it from the Pholiotas of this book.

Naucoria vinicolor Color Plate 94

CAP 1-4 cm broad, convex to plane or obtusely umbonate; surface smooth or finely
fibrillose or occasionally fibrillose-scaly, not viscid, dark red to wine-red (vinaceous), the
center often darker. Flesh thin, tinged cap color; odor mild. GILLS slightly decurrent to
adnate, adnexed, or notched, close; dark red or vinaceous when young, soon becoming
cinnamon or rusty-brown as spores mature. STALK 1-7 cm long, 2-6 mm thick, equal or
slightly thicker below, colored more or less like cap but often overlaid with a fine whitish
silky-fibrillose coating. VEIL fibrillose, vinaceous, disappearing or forming a slight hairy
zone on stalk which turns cinnamon from falling spores. SPORE PRINT cinnamon-
brown; spores 6.5-8 * 4-5 microns, elliptical or bean-shaped, smooth.

HABITAT: In small tufts or clusters (or sometimes solitary) on dead wood; known only
from California, but perhaps more widely distributed. I have found it several times on oak
in the fall and winter, but it is rare.
EDIBILITY: Too rare to be of value. Greg Wright, who has found it in Los Angeles
County, reports that it is harmless, with a “mealy, moderately mushy” texture and a flavor
“that suggests bland beef.” (I just can’t wait to try it! Can you?)
COMMENTS: This interesting mushroom was originally described in 1909. It does not
belong to the genus Naucoria in its modern sense, but has not officially been transferred to
another genus—perhaps because of its rarity, but perhaps also because its “correct” genus
is in doubt! The fibrillose veil and cinnamon-brown spores give it the aspect of a small
Cortinarius (e.g., C. sanguineus), but it always seems to grow on wood, usually in small
clusters. Its wine-red color and small size plus the spore color and growth on wood are the
distinguishing field marks. Lactarius fragilis sometimes grows on wood but has a latex
and/ or a fragrant odor, lacks a veil, and has white or yellowish spores. Naematoloma
aurantiaca is similarly colored but has darker spores and is usually terrestrial.

404
 405

CREPIDOTUS
Small to medium-sized mushrooms typically growing shelflike on wood. CAP usually round to
kidney-shaped in outline, surface smooth or hairy. Flesh soft, thin. GILLS usually brownish at
maturity. STALK usually absent or rudimentary, or if present then lateral or off-center. VEIL and
VOLVA absent. SPORE PRINT dull brown to yellow-brown, cinnamon-brown, or pinkish-
brown. Spores smooth or roughened, lacking an apical germ pore. Cystidia often present on gills.
Cap cuticle filamentous.

THIS is a fairly common but lackluster group of wood-inhabiting mushrooms with little
or no stem. They superficially resemble the oyster mushrooms (Pleurotus) as well as
Phyllotopsis and Claudopus, but have brown spores. Other brown-spored, wood-
inhabiting mushrooms (Pholiota, Galerina, etc.) have well-developed stems. Crepidotus
is given its own family, the Crepidotaceae, by some mycologists.
Crepidotus species are worthless as food—they are flaccid and decay rapidly. They
favor decaying hardwood logs, branches, and twigs (especially oak), but a few species
occur on conifers or even in soil—in which case they may have a lateral to off-center (but
not central) stem. Two widespread representatives are depicted here.
Key to Crepidotus
l. Fruiting body tough and leathery or woody.(see Polyporaceae& Allies, p. 549)
1. Not as above; fruiting body fleshy . 2
2. Cap white or whitish . 3
2. Cap tawny to pale ochre, brownish, or red (but may sometimes fade to whitish in age) .... 4
3. Cap 1-4 cm broad and nearly smooth (bald) C. applanatus & others (see C. herbarum, below)
3. Cap smaller (up to 2 cm broad) and distinctly hairy or downy . C. herbarum & others, below
4. Cap bright red (scarlet to cinnabar-red), small (up to 15 mm broad); gill edges also scarlet to red;
found on hardwoods in eastern North America; rare .C. cinnabarinus
4. Not as above; cap not red . 5
5. Found on fabrics or old carpets, seat covers of abandoned cars, mattresses, “rotting blue jeans,”
or on wood; fruiting body yellow-brown to brown or cinnamon; stalk present, often darker,
usually off-center, curved, and slender .Melanotus textilis
5. Not as above; found on wood or lignin-rich humus .6
6. Gills often forked or veined, especially near base of cap .(see Paxillaceae, p. 476)
6. Not as above . 7
7. Gills yellow to orange or ochre-orange . C. crocophyilus(see C. mollis, p. 406)
7. Gills whitish to grayish, brownish, or dull cinnamon . C. mollis, p. 406

Crepidotus herbarum (Little White Crep)

CAP 0.5 -2 cm broad, kidney-shaped to nearly round in outline; surface hairy or downy,
white, not viscid. Flesh very thin, white. GILLS fairly well-spaced, white becoming pale
ochre or brownish; radiating from point of attachment to substrate. STALK absent or
rudimentary. SPORE PRINT pale yellow-brown; spores 6-8 * 3 A microns, pip-shaped to
lance-shaped or somewhat elliptical, smooth.

Crepidotus variabilis (see comments under C. herbarum) is one of several small whitish shelving
species with brown or pinkish-brown spores.
406 CORTINARIACEAE

HABITAT: Scattered or in groups or troops on fallen branches, twigs, herbaceous stems,

anddebris(usually of hardwoods); widely distributed. Common in our area throughout the
mushroom season, but often overlooked.
EDIBILITY: Unknown, and like most of us, destined to remain so.
COMMENTS: This is one of several small whitish Crepidotus species that typically grow
on twigs and branches, sometimes completely covering their host. It is likely to be mistaken
for a Claudopus or small Pleurotus, but has brownish spores. Similar species include:
C. versutus, with dull brown, elliptical spores 9-10 microns long; C. variabilis, with pale
brown to pinkish-brown, minutely warty, elliptical spores 5-7 microns long; C.fusisporus,
with pinkish-buff, fusiform (elongated) spores; C. applanatus, very common and widely
distributed, with round, minutely spiny spores and a larger (1 -4 cm), nearly smooth, white
cap that often becomes brownish in age; and C. maculans, also larger and smooth, but
usually blackish-spotted in age.

Crepidotus mollis (Jelly Crep; Flabby Crepidotus)

CAP 1 -5 (8) cm broad, fan- or kidney-shaped to nearly round in outline, convex to plane;
surface gelatinous in wet weather beneath a dense to rather sparse coating of fulvous to
rusty-ochre to brown fibrils (hairs) or small fibrillose scales; in age often smooth or with
very few fibrils and varying in color from tawny to pale ochre to brown, or fading to whitish.
Flesh soft, thin, pallid, soon flaccid. GILLS close, whitish becoming brown or dull cinna¬
mon; radiating from base of cap. STALK absent or rudimentary. SPORE PRINT dull
brown to yellowish-brown; spores 7-11 * 4.5-6.5 microns, elliptical, smooth.
HABITAT: Usually in groups or overlapping tiers on the bark of dead hardwoods (or
rarely conifers); very widely distributed and common. I n our area it is frequent throughout
the mushroom season, especially on live oak.
EDIBILITY: Unknown.
COMMENTS: Also known as C. fulvotomentosus and C. calolepis, this is our most
common and conspicuous Crepidotus, often fruiting in attractive masses on dead oaks.
The relatively large size (for a Crepidotus), gelatinous texture when wet, brown spores and
mature gills, and fibrillose scales on the cap when young are the main fieldmarks. In age it
becomes quite flabby and is likely to attract your attention for this reason if for no other.
C. crocophyllus is a similar, widely distributed species with yellow to ochre-orange imma¬
ture gills. I have found it several times in our area.

Crepidotus mollis is a common shelving mushrooms with a flabby cap and brown spores. Youngcaps
shown here are covered with brown fibrils or scales, but older or rain-battered caps can be bald.
 407

GYMNOPILUS
Medium-sized to large mushrooms found mostly on wood. CAP smooth or scaly, dry. GILLS
notched to slightly decurrent, usually yellow to rusty-orange. STALK more or less central, fleshy.
VEIL usually (but not always) present, sometimes forming an annulus (ring). VOLVA absent.
SPORE PRINT orange to rusty-orange to bright rusty-brown. Spores typically elliptical and
roughened, without an apical pore. Cystidia present, at least on gill edges. Cap cuticle filamentous.

THIS is a clearcut group of rusty-orange-spored mushrooms formerly divided among

Pholiota and the defunct genus Flammula. The fruiting body is typically reddish-brown to
rusty-orange to yellow, and a veil is often present. The vast majority of species grow on
wood but at times may appear terrestrial. Pholiota and Cortinarius are the genera most
often confused with Gymnopilus. Pholiota, however, usually has a viscid cap and duller
(brown to cinnamon-brown) spores, while Cortinarius grows on the ground.
Gymnopilus species are found primarily in the woods, but sometimes turn up on cut
stumps, wood chip mulch, and in nursery flats and flower pots. They are quite common,
but rarely fruit in the large numbers typical of, say, Hebeloma, Cortinarius, and Inocybe.
About 75 species of Gymnopilus are known from North America and over 25 from
California. Several are quite striking but none are good edibles. A few—specifically G.
spectabilis, G. validipes, and G. aeruginosus—are “pupil-dilating” (hallucinogenic).
Four species are depicted here.

Key to Gymnopilus
1. Cap usually tinged or variegated with blue or blue-green when young (sometimes also with
other colors) or staining bluish when bruised .2
1. Not as above; blue or blue-green shades absent . 3
2. Cap with fibrils or scales; veil present, at least when young . G. aeruginosus, p. 409
2. Cap nearly smooth; veil absent .G. punctifolius(see G. aeruginosus, p. 409)
3. Cap with fibrils or scales of a radically different color than the background(i.e., reddish, pinkish,
brownish, etc.); not common .4
3. Cap smooth or with fibrils or scales that are roughly the same color as background (i.e., some
shade of yellow, orange, rusty-orange, or reddish-brown); common . 6
4. Cap scales or fibrils red to purple-red, reddish-brown, or pink . 5
4. Cap scales or fibrils tawny to dark brown to blackish-brown .
.G. fulvosquamulosus& G. parvisquamulosus(see G. luteofolius, p. 409)
5. Fruiting body (especially cap) sometimes with bluish or blue-green stains (examine several
specimens!); veil evanescent, not typically forming a distinct annulus (ring) .
.G. aeruginosus & others, p. 409
5. Not as above; veil often forming a slight annulus .G. luteofolius & others, p. 409
6. Cap medium-sized togigantic(up to 40 cm broad or more!); stalk 1-7 cm thick; veil membranous
or fibrillose, often forming an annulus (rng) on stalk; usually growing in clusters (but occa¬
sionally solitary) .G. spectabilis group, p. 410
6. Not as above; smaller and veil absent or evanescent; not often clustered .7
7. Veil present when young (but often disappearing in age) . 8
7. Veil absent(check young specimens!) . 10
8. Veil whitish .G. penetrans & G. flavidellus (see G. sapineus, p. 408)
8. Veil pale yellow or yellowish .9
9. Cap golden-yellow to tawny-orange, often with minute scales or scattered fibrillose patches
. G. sapineus, p. 408
9. Cap darker (tawny to cinnamon, russet, or reddish-brown), smooth .
.G. luteocarneus(see G. sapineus, p. 408)
10. Usually growing on ground; taste mild . G. terrestris(see G. sapineus, p. 408)
10. Typically found on wood; taste usually bitter G. liquiritae& G. beltulus( see G. sapineus, p.408)
Gymnopilus luteocarneus (see comments under G. sapineus, below) is one of several small orangish
species with an evanescent veil or no veil at all. Mature caps can be somewhat larger than the ones
shown here, but never approach the size of the G. spectabilis group. Left: A young specimen on wood
and an older one on a cone. Right: Mature specimen; note how stalk base is darkened by falling spores.

Gymnopilus sapineus (Common and Boring Gymnopilus)

CAP (1) 2-5 (9) cm broad, convex to nearly plane or sometimes obscurely umbonate;
surface dry, usually with minute scales or scattered patches of fibrils, often cracking in age;
golden-yellow to tawny-orange, the margin sometimes paler. Flesh yellowish, firm; taste
usually bitter. GILLS attached (usually adnate), close, yellow becoming rusty-yellow to
rusty-cinnamon in age. STALK2.5-7 cm long,3-7(10) mm thick, equal or tapered slightly
below; yellowish-buff to yellow, becoming brownish-yellow in age or when handled; fibril-
lose. VEIL yellowish, fibrillose, disappearing or leaving a few hairs near top of stalk which
turn rusty from falling spores. SPORE PRINT amber- to rusty-orange to bright rusty-
brown; spores 7-10 x 4-5.5 microns, elliptical, minutely roughened.
HABITAT: Solitary to scattered or in small tufts or groups on rotting logs, cones, and
humus rich in lignin; widely distributed, but rarely fruiting in large numbers. In our area
it is common under conifers (especially pine) throughout the mushroom season. Similar
species turn up occasionally in nursery flats and flower pots.
EDIBILITY: Unknown; the small size and bitter taste are deterrents.
COMMENTS: There are several small, eminently undistinguished and evidently indis¬
tinguishable Gymnopilus species that will more or less fit the above description. As a group
they can be recognized by their yellow to orange-brown color, modest size, growth on
wood, dry cap, and rusty-orange spores. Some have an evanescent fibrillose veil, others
have no veil at all. G. sapineus is distinct by virtue of the small scales or fibrillose patches
on the cap. The others are hardly worth differentiating, but some of them are: G. luteo¬
carneus, common in California on conifers, with a smooth, darker (tawny-cinnamon to
russet) cap and pale yellowish, evanescent veil; G. penetrans and G. flavidellus, with a
smooth yellow to yellow-orange cap and whitish evanescent veil, the latter with a stalk
that stains orange-brown when handled; and the following species with no veil and a
smooth, tawny to orange, rusty, or cinnamon-brown cap: G. liquiritiae, growing on wood,
taste bitter; G. bellulus, on wood, taste bitter, but cap only 1-2.5 cm broad and smooth to
minutely scurfy; and G. terrestris, usually terrestrial under conifers, with a mild taste.
Left: A close relative of G. aerwgj>?o.s«s(perhapsthesame?),thisunidentified Gymnopilus has dark red
fibrils on the cap and stalk when young and often exhibits bluish stains. N ote absence of annulus(ring)
on stalk (see comments below). Right: Close-up of G. luteofolius showing cap fibrils and annulus.
\

Gymnopilus aeruginosas
CAP (2) 5-15 (23) cm broad, convex to nearly plane; surface dry, fibrillose-scaly (or
cracked in age); color variable: at first dull bluish-green or variegated with green, yellow,
salmon, red, or vinaceous, sometimes fading to buff or pinkish-buff in age; scales tawny to
reddish to dark brown. Flesh whitish or tinged blue or green; taste bitter. GILLS adnexed
to adnate or slightly decurrent, often seceding; buff to yellow-orange or ochre, close.
STALK (3) 5-12 cm long, (0.4) 1 -1.5 (4) cm thick, more or less equal, colored more or less
like the cap, smooth or fibrillose, dry. VEIL yellowish, fibrillose, often scanty, leaving an
evanescent zone of hairs near top of stalk. SPORE PRINT rusty to rusty-orange or rusty-
cinnamon; spores 6-9 * 3.5^4.5 microns, elliptical, roughened.
HABITAT: In groups or clusters on stumps, logs, and sawdust of both hardwoods and
conifers; widely distributed, but most common in the Pacific Northwest. I have seen large
fruitings in the fall, winter, and spring in wood chip mulch.
EDIBILITY: Hallucinogenic. The blue-green tones are indicative of psilocybin and/
or psilocin, as in Psilocybe.
COMMENTS: The tendency of the cap to exhibit a blue-green tinge when young (often
variegated with pink, vinaceous-red, and/ or other colors) plus the scaly cap, yellowish
gills, rusty-orange spores, and growth on wood make this species quite distinctive. G.
harmoge may be a synonym. A similar unidentified species (perhaps the same?) with
vinaceous-red fibrils on the cap and stalk when young (see photo) is common on wood
chips. Its cap often shows bluish or blue-green stains and it is hallucinogenic. G. puncti-
folius is colored like G. aeruginosus, but has a nearly smooth (bald) cap and no veil.

Gymnopilus luteofolius
CAP 2-12 cm broad, convex or obtuse becoming nearly plane; surface dry, at first covered
with dense, dark red to purple-red or reddish-brown, fibrillose scales, these fading slowly
to pinkish-red or yellowish-red; surface finally yellowish in old age as scales disperse;
margin inrolled at first. Flesh thick, reddish to lavender, then fading to yellowish; taste
bitter. GILLS notched to adnate or slightly decurrent, fairly close, yellow becoming bright
rusty-orange or rust-colored as spores mature. STALK 3-10 cm long, 0.3-2 cm thick,
fleshy, equal or enlarged below (or tapered downward if clustered); solid, dry, fibrillose,
more or less colored like cap, becoming yellowish or rusty-stained. VEIL fibrillose to
somewhat membranous, yellowish; forming a hairy, superior ring on stalk which may
disappear or trap falling spores. SPORE PRINT bright rusty-orange; spores 5.5-8.5 *
3.5-4.5 microns, elliptical, roughened.

409
410 CORTINARIACEAE

HABIT AT: In groups or clusters on decaying coniferous wood, sawdust, and humus rich in
lignin (rarely on hardwoods); widespread, but not common. I have seen one fantastic
local fruiting, under pine at New Brighton Beach State Park, with Pluteus cervinus and
large clusters of Pholiota terrestris and Naematoloma fasciculare (all certified wood-
lovers) also present.
EDIBILITY: U nknown, but the closely related G. aeruginosus is said to be hallucinogenic.
COMMENTS: This distinctive mushroom can be told by the dark red fibrils or scales on
the cap (and often the stem), plus the yellow gills, rusty-orange spores, and relatively
persistent veil. The much more common Tricholomopsis rutilans features the same at¬
tractive color combination, but has white spores and lacks a veil, while G. aeruginosus and
relatives have a transient veil and often have bluish stains. G. pulchrifolius is a smaller spe¬
cies with a pink to pale pink cap. Others with distinctively colored cap scales include: G.
fulvosquamulosus, with tawny to brown scales on a yellow background; and G.
parvisquamulosus, with dark brown to blackish-brown scales on an ochraceous-tawny
background. Both have a mild taste and fibrillose to slightly membranous veil. I have
found them in our area, but they are rare.

Gymnopilus spectabilis group Color Plate 99

(Big Laughing Mushroom; Giant Gymnopilus)
CAP 5-40 cm or more broad, convex becoming broadly convex or nearly plane; surface
dry, smooth to silky or fibrillose, often breaking up to form small scales; bright yellow-
orange to yellowish-buff when young (or at times nearly whitish from overlaid fibrils),
often somewhat darker in age (rusty-orange to golden-tawny to orange-brown or reddish-
brown); margin at first incurved, sometimes wavy, sometimes with veil remnants. Flesh
thick, firm, yellowish; taste bitter. GILLS notched to adnate or slightly decurrent, close,
ochre-buff or pale yellow becoming more or less rusty-orange to rusty-brown in age.
STALK (3) 5-25 cm long, 1-6 (10) cm thick, usually swollen in the middle or below, the
base often narrowed; solid, firm, dry, rusty-orange to rusty-yellow or paler, fibrillose below
the ring. VEIL pallid to pale yellowish or rusty-stained, membranous or fibrillose,
sometimes disappearing, but usually forming a superior ring on stalk; ring soon stained
with rusty-orange spores, often collapsing or disappearing in age. SPORE PRINT bright
rusty-orange; spores 7-10.5 x 4.5-6 microns, elliptical, roughened or wrinkled.
HABITAT: Usually in clusters (occasionally solitary) on or around stumps and trees;
widely distributed. This species “complex” favors conifers on the west coast, hardwoods
in eastern North America. In our area it is quite common on old pine stumps in the fall,
winter, and early spring, and occasionally turns up on eucalyptus also.
EDIBILITY: Inedible due to the bitter taste. Forms in Asia and eastern North America
apparently contain psilocybin and/ or psilocin and are hallucinogenic (hence the Japanese
name, “Big Laughing Mushroom”). On the west coast, however, it is apparently “in¬
active.” An Ohio woman had an unforgettable experience after inadvertently nibbling on
one. She found herself in an alien world of fantastic shapes and glorious colors, and while
concerned friends were rushing her to the hospital, she was heard to mutter, “If this is the
way you die from mushroom poisoning, then I’m all for it. ..”
COMMENTS: This species “complex” is easily recognized by its overall yellow-orange
to rusty-orange color, rusty-orange spores, robust size, bitter taste, and frequent presence
of a ring formed by the veil. Several Pholiota species are similar, but have duller spores and
viscid caps; the honey mushroom (Armillariella mellea) is also somewhat similar, but has
white spores and whitish or flesh-colored gills, while the jack-o-lantern mushrooms
Gymnopilus spectabilis group is easily told by its yellow to rusty-orange color (see color plate),
membranous veil, and rusty-orange spores. Note how small the penny is! This large western form is
considered to be a separate species, G. ventricosus, by some authorities.

(Omphalotus) have white or yellow-tinted spores, lack a veil, and grow only on hardwoods.
“Typical” G. spectabilis grows on both hardwoods and conifers and has a somewhat
fibrillose veil. It was originally called Pholiota spectabilis and is also known as G.junonius
(probably its “correct” name). The large, non-hallucinogenic, conifer-loving westernform
with a swollen stalk and membranous veil is called G. ventricosus {if truly distinct). It is
one of our largest and most spectacular mushrooms, commonly attaining pizza size(more
than one foot in diameter) with clusters weighing 10 pounds or more. Young specimens
present an entirely different appearance: squat and compact with hard, very thick, yellow
flesh and very narrow (shallow) gills. Other species in the G. spectabilis^complex” include:
G. subspectabilis, with larger spores; and G. validipes of eastern North America, a hallu¬
cinogenic species with a mild to only slightly unpleasant taste and a fibrillose veil.

ROZITES & PHAEOLEPIOTA

Medium-sized to large, terrestrial, woodland mushrooms. CAP dry; often wrinkled (Rozites)
or powdery-granulose (Phaeolepiota). GILLS attached, tawny to orange-buff to brownish at
maturity. STALK central, fleshy, smooth to fibrillose (Rozites) or powdery-granulose (Phaeo¬
lepiota). VEIL present, membranous, usually forming a distinct annulus (ring) on stalk. SPORE
PRINT rusty-brown to tawny or orange-buff Spores elliptical, roughened to nearly smooth.

THOUGH not closely related, these two small genera are treated together here because
they have a number of features in common: both grow on the ground, both have yellow-
brown to rusty-brown spores and a well-developed membranous veil that forms a distinct
annulus on the stalk, and both were originally placed in the genus Pholiota.
Rozites has somewhat wrinkled or roughened spores and has been aptly characterized
as “a Cortinarius with a membranous veil.” Phaeolepiota, on the other hand, has a granu-
lose or powdery cap and stem and has consequently been called “a brown-spored Cysto-
derma.” (Some mycologists go so far as to place it alongside Cystoderma in the Agari-
caceae; others retain it in Pholiota of the Strophariaceae.)
Each genus includes a single well-known species. Both are edible and quite good, but
some people are apparently “allergic” to Phaeolepiota.

411
412 CORTINARIACEAE

Key to Rozites & Phaeolepiota

1. Cap and stalk covered with a granulose or powdery layer (but the granules sometimes wearing
or washing off, especially those on the cap); stalk 1.5 cm thick or more; found mainly in the
Pacific Northwest .Phaeolepiota aurea, below
1. Cap and stalk not powdery or granulose . 2
2. Gills purplish when young or some part of fruiting body purple-tinged (stt Cortinarius, p. 417)
2. Not as above; purple or violet shades absent . 3
3. Cap distinctly viscid or slimy when moist . 4
3. Cap not viscid (or only very slightly so and soon drying out) . 5
4. Gills covered by a cobwebby partial veil when young; annulus (ring) on stalk formed by the outer
(universal) veil; spore print rusty- to cinnamon-brown.(see Cortinarius, p. 417)
4. Not as above . (seePholiota, p. 384)
5. Cap warm tan to yellow-brown or orange-brown, often wrinkled radially (especially in age) and
sometimes with a thin silky or hoary bloom when young; spore print rusty-brown; found in
forests .Rozites caperata, below
5. Not as above; cap dark brown to olive-brown to tan, creamy, etc.; spore print dull to dark brown
but not rusty-brown; found in forests as well as suburban habitats . . (seeAgrocybe, p. 467)

Rozites caperata (Gypsy Mushroom) Color Plate 101

CAP 5-15 cm broad, oval becoming somewhat bell-shaped to broadly convex, plane, or
obscurely umbonate; surface dry, usually distinctly wrinkled or corrugated radially, at
first covered with a thin white to grayish coating of silky fibrils (especially at center); warm
tan to yellow-brown or orange-brown, margin often paler. Flesh thick, white, firm. GILLS
adnate to adnexed or notched, close, at first pallid, soon dull tawny or brown, sometimes
transversely banded with darker and lighter zones. STALK 5-13 cm long, 1-2.5 cm thick,
equal or slightly enlarged at base, solid, firm, white to pale tan or pale ochre; apex often
striate or scurfy, base sometimes with an obscure volvalike zone. VEIL white, membran¬
ous, forming a more or less median ring on stalk. SPORE PRINT rusty-brown; spores
11-15 * 7-10 microns, elliptical, roughened or warty. Some cystidia present on gill edges.
HABITAT: Scattered or in groups on ground in woods; widely distributed in northern
regions. It favors mossy, old-growth coniferous forests but also grows under hardwoods,
especially when huckleberry is in the vicinity. I have seen large fruitings in Washington
and Idaho in the late summer and fall. It also occurs in northern California and the Sierra
Nevada, but I have yet to find it in our area.
EDIBILITY: Edible, and in my humble fungal opinion, the best of the Cortinariaceae.
It is especially good with rice after a long, hard day of backpacking. The tough stems should
be discarded.
COMMENTS: Originally called Pholiota caperata, the gypsy mushroom is easily told
by its warm brown or yellowish, wrinkled cap that has a hoary sheen when young, plus
the membranous veil which forms an annulus(ring) and the rusty-brown spores. Agrocybe
praecox is somewhat similar, but does not have a wrinkled cap and usually grows in culti¬
vated ground; Phaeolepiota aurea differs in its powdery-granulose cap and stem, while
similarly-colored Cortinarius species do not have a membranous veil.

Phaeolepiota aurea
CAP 6-20 (30) cm broad, obtuse to convex becoming broadly convex, plane, or broadly
umbonate; surface dry, granular to somewhat powdery (the granules sometimes wearing
away in age), orange to orange-tan, tawny-yellow, or golden-brown, often somewhat paler
PHAEOLEPIOTA 413

in age; margin usually hung with veil remnants. Flesh thick, pallid or yellowish. GILLS
adnate to notched or free, close, pallid or pale yellowish becoming tawny to orange-
brown. STALK 5-15 (25) cm long, (1)2^4 (6) cm thick, thicker toward base, orange to buff
or colored like cap and granulose or powdery below the ring. VEIL membranous, colored
like cap, sheathing the stalk and breaking to form a superior flaring or funnel-like ring
which eventually collapses or becomes skirtlike; ring smooth on upper surface and granu¬
lose on underside. SPORE PRINT pale yellow-brown to orange-buff; spores 10-14 * 5-6
microns, elliptical, smooth to minutely roughened.
HABITAT: In groups or clusters in rich humus and soil under both hardwoods and
conifers; known from the Pacific Northwest and Alaska (also Europe), fruiting in the
late summer and fall. It is rather rare, but when it fruits it often does so in large quantities.
An ideal place to look for it is under alder along roads and trails.
EDIBILITY: Edible for most people, but mildly poisonous to some. It is a tempting
specimen, but try it cautiously if you must.
COMMENTS: Also known as Pholiota aurea and Togaria aurea, this large, beautiful
mushroom is as distinctive as it is rare. No other large, brownish-spored mushroom is
golden-brown to pale orange with a granulose coating on both the cap and stem.
Cystoderma species are granulose but much smaller and have white spores; Agaricus
augustus is similarly colored but has chocolate-brown spores and an almondy odor, while
Pholiota species, Rozites caperata, and Agrocybe praecox lack the granulose coating.

PHAEOCOLLYBIA
Medium-sized, terrestrial, woodland mushrooms. CAP usually viscid or slimy when moist. GILLS
adnexed to free, usually rusty-brown or cinnamon at maturity. STALK with along, tapered, deeply
rooting base or “tap root. ” VEIL and VOLVA absent. SPORE PRINT cinnamon- to rusty-brown.
Spores elliptical, roughened, without an apical germ pore. Cystidia present, at least on gill edges.

THE presence of a “tap root” and absence of a veil distinguish this small, well-marked
genus from its closest relative, Cortinarius. The “tap root” (pseudorhiza) is merely an
extension of the stem that roots deeply in the humus. It gradually tapers to a point, be¬
coming so thin and fragile that it can’t be traced to its origin (probably tree roots). Most
other agarics with “tap roots” (e.g., Caulorhiza and Oudemansiella) have white spores.
Phaeocollybias are quite rare in most regions(including ours), but where they do occur
they often fruit in large numbers. The Pacific Northwest is an exception, for it is there
that the genus is most common and diverse. Many types seem to occur only with Sitka
spruce, while several favor western hemlock, redwood, or Douglas-fir; few, if any, occur
with hardwoods. About two dozen species are known from N orth America, many of them
restricted to the Pacific Northwest. Little is known of their edibility. Three representatives
are described here and several others are keyed out.

Key to Phaeocollybia
1. Cap dark green to olive, at least when fresh and moist (but may fade as it loses moisture) .. 2
1. Cap never greenish or olive .4
2. Gills violet or lilac when young .P. fallax (see P. o/ivacea, p. 414)
2. Gills whitish to pale brown when young, or at least not violet . 3
3. Stalk typically slender (less than 1 cm thick at apex); cap 1 -6 cm broad .
.P. festiva& P. pseudofestiva (see P. olivacea, p. 414)
3. Stalk typically about 1 cm thick (or more) at apex; cap 3-11 cm broad . . .. P. olivacea, p. 414
414 CORTINARIACEAE

4. Gills violet or lilac when young .P. lilacifolia(see P. californica, p. 415)

4. Gills pallid to yellowish, tan, brownish, or cinnamon when young .5
5. Cap typically with grayish tones (grayish-brown to grayish-buff) at maturity or when faded 6
5. Not as above; cap dark brown to amber-brown, cinnamon, reddish, or orangish .7
6. Stalk typically 8 mm thick or more at apex .P. gregaria(see P. olivacea, p. 414)
6. Stalk typically 3-7 mm thick at apex .P. scatesiae(see P. olivacea, p. 414)
7. Stalk slender (up to 7 mm thick at apex) and cap rather small (up to 5 cm broad) .8
7. Stalk thicker than above (usually at least 5 mm at apex) and cap 3-7 cm broad or larger .. 10
8. Stalk 1.5-3 mm thick at apex; cap not viscid or only slightly so .
.P. similis(sQQ P. calif ornica, p. 415)
8. Not as above; stalk usually at least 3 mm thick at apex, or if thinner than cap viscid or slimy 9
9. Cap amber-brown to yellowish to salmon or salmon-ochre; spores at least 8 microns long . . .
.P. attenuata& P. laterarius(see P. californica, p. 415)
9. Cap cinnamon-brown to reddish or orangish P. radicataSi others (see P. californica, p. 415)
10. Cap (4) 6-25 cm broad, cinnamon to dark reddish-brown to liver-colored, dark brown, or
umber; surface viscid or slimy; stalk 1-4 cm thick at apex . 11
10. Not as above; either smaller, slimmer, or oranger than above, or cap not viscid . 12
11. Cap dark brown to umber or sometimes dark pinkish-brown; cap conical when young.
.P. spadicea(see P. kauffmanii, p. 416)
11. Cap usually redder or more liver-colored than above and/ or not conical when young.
.P. kauffmanii & others, p. 416
12. Cap slimy or viscid when moist .P. californica Sc others, p. 415
12. Cap not viscid . P. deceptiva (see P. californica, p. 415)

Phaeocollybia olivacea (Olive Phaeocollybia)

CAP 3-11 cm broad, obtusely conical or convex becoming plane or uplifted, but usually
retaining an umbo; surface smooth, viscid or slimy when moist, hygrophanous: deep green
to olive-green when moist, fading as it dries (to olive-buff, etc.). Flesh thin, olive; odor
cucumberlike or radishlike when fresh. GILLS pallid becoming pale brown, then rusty-
brown; adnexed or free, close, edges often eroded or wavy. STALK 10-22 cm long or more,
(0.5) 1-2.5 cm thick at apex, equal above or swollen at ground level, then tapering below
the ground to form a “tap root”; cartilaginous, smooth, stuffed with a pith; watery olive
to yellowish above, rusty-orange to reddish-brown below or occasionally overall. SPORE
PRINT rusty-brown; spores 8-11 ><5-6 microns, elliptical with snoutlike apex, roughened.
HABITAT: Scattered to densely gregarious, often in large rings, on ground in mixed
woods and under conifers; fruiting in the fall and winter, known only from the west coast.
It is said to be common in southern Oregon and northern California, but is rare in our area.
Budding boletivore Craig Mitchell has located one very prolific fairy ring near Big Basin
State Park.
EDIBILITY: Unknown
COMMENTS: This striking mushroom is easily recognized by its slimy olive-green cap,
rather thick rooting stem, rusty-brown spores, and absence of a veil. It is somewhat
unusual among the Phaeocollybias in that it often fruits in mixed woods (i.e., it may or
may not be associated with conifers). There are three other Phaeocollybias with an olive
cap when fresh; P. festiva, P. pseudofestiva, and P.fallax. All are considerably slimmer,
smaller, and more conical (cap 1-6 cm broad, stalk less than 1 cm thick at apex) than P.
olivacea, and all have slightly smaller spores. P. fallax is easily told by its violet or lilac
gills when young, but the other two can only be differentiated microscopically. Also
worth mentioning are two species which have grayish-toned caps: P. scatesiae, with a
slim (3-7 mm) stalk, dingy yellowish gills when young, and a dull cinnamon cap that fades
Phaeocollybia olivacea. Greenish to olive-gray cap, rusty-brown spores, and “tap root” are
distinctive.

to grayish-brown, often fruiting in massive clusters of up to several hundred fruiting

bodies; and P. gregaria, with a thicker (8-15 mm) stalk and a yellowish-gray to brownish-
gray or grayish-buff cap. All of the above species occur with northern conifers and are
very rare or absent in our area.

Phaeocollybia californica Color Plate 100

CAP (2) 3-7(10) cm broad, conical with an inrolled margin when young, often expanding
in age to plane but retaining a distinct umbo; surface smooth, viscid when moist, hygro-
phanous: amber-brown to cinnamon-brown or orange-brown when moist, fading as it
dries, but in age often rusty-spotted or darkening to reddish-brown. Flesh thin; odor
pungent (somewhat radishlike). GILLS adnate to adnexed or even free, pallid or tinged
cap color, darkening to brown or rusty-brown in age; close. STALK 10-20 cm long or
more, 4-10 mm thick, equal above, gradually tapering below the ground to form a “tap
root”; cartilaginous, hollow, smooth, colored more or less like cap or paler, or more often
dark reddish or reddish-brown below and paler (apricot-buff) above. SPORE PRINT
rusty-brown; spores 8-11 * 5-6 microns, elliptical with a small “beak” at apex, roughened.
HABIT AT: Densely gregarious or in large loose clusters on ground under conifers, often
forming arcs or fairy rings; known only from the west coast, not common. In our area it
is quite rare, but one large fairy ring under redwood and Douglas-fir fruits in colossal
quantities every winter.
EDIBILITY: Unknown.
COMMENTS: The presence of a “tap root,” absence of a veil, rusty-brown spores, and
amber-brown to orangish or reddish-brown color are the distinctive features of this
species and its look-alikes (see below). Caulorhiza umbonata also features a “tap root,”
but is much more common and has pallid gills, a non-viscid cap, and white spores. Other
dark cinnamon to orangish Phaeocollybias include: P. piceae, with a bitter taste and
more or less mild odor, favoring Sitka spruce; P. dissiliens, also fond of Sitka spruce, but
with distinctly smaller spores and a thicker stalk that easily splits lengthwise; P. lilacifolia,
with violet gills when young; P. attenuata, with a thinner stem (3-6 mm thick at apex) and a
viscid, amber-brown to yellowish-buff cap; P. laterarius, also slender but with a salmon
to pale salmon-ochre cap when moist (known from Michigan); P. similis, with a very
thin stalk (1.5-3 mm) and a non-viscid or only slightly viscid cap; andP. deceptiva, with
a cinnamon to yellowish-brown, non-viscid cap and a thicker (8-15 mm) stalk that does

415
Phaeocollybia californica. Note conical cap, “tap root,” and absence of veil. Also see color plate.

not markedly redden with age. Finally, there is a group of species (e.g., P. radicata, P.
jennyae, P. christianae, P. sipei) with a slender (3-6 mm) stem, small cap, and smaller
spores (less than 8 microns long) that are best differentiated from each other microscopi¬
cally. Nearly all of the above occur in the Pacific Northwest; a few are more widely dis¬
tributed or have counterparts (e.g., P. rufipes) in eastern North America. None are as
large as P. kauffmanii and none show the olive or grayish tones of P. olivacea and the
species listed under it.

Phaeocollybia kauffmanii (Giant Phaeocollybia)

CAP 7-25 cm broad, obtuse or convex when young becoming broadly umbonate to nearly
plane in age; surface smooth, viscid to very slimy; cinnamon to pale reddish-cinnamon
or reddish-brown becoming more or less liver-colored in age, but fading to reddish-orange,
rusty-reddish, or apricot as it loses moisture; margin remaining inrolled for a long time,
not striate. Flesh thick, firm, colored like cap or paler; odor usually farinaceous, at least
when crushed. GILLS close or crowded, free or adnexed, buff becoming brown to rusty-
brown in age. STALK (15) 20-40 cm long, 1.5^1 cm thick at apex, tapered downward to
form a long “tap root” below the ground; pinkish-brown to pinkish-buff above, reddish-
brown to dark purplish-brown below and darkening with age (sometimes nearly black);
smooth; stuffed with a pith, the outer rind tough and cartilaginous. SPORE PRINT
cinnamon-brown; spores 8-11 * 4.5-7 microns, elliptical with an apical beak, roughened.
HABITAT: Solitary, scattered, or in groups under conifers (particularly Sitka spruce)
in the late summer and fall; known only from the west coast. It is fairly common during
some seasons, particularly in Oregon and northern California, rare during others. I have
not seen it in our area.

EDIBILITY: Unknown. Its large size is tempting but the glutinous cap is not.
COMMENTS: This giant of the genus can usually be recognized by its size alone. The
colors are quite variable depending on age and moisture, but are usually darker and
redder than those of P. calif ornica and the species discussed under it. No other Phaeo¬
collybia has a stalk so consistently thick nor a cap so broad. The obtuse rather than conical
shape of young caps is also distinctive. Other species: P. oregonensis is a very similar but
slightly smaller species with smaller spores; P. spadicea is a fairly large species (cap 4-12
cm broad, stalk 1 -2 cm thick) with a dark brown to umber cap when moist. Both occur in
the Pacific Northwest.
416
 417

CORTINARIUS
Woodland fungi nearly always growing on the ground. CAP viscid or dry, smooth to fibrillose or
scaly. GILLS typically attached but not often decurrent; variously and often brightly colored when
young but usually rusty-brown to cinnamon-brown in age. ST ALK thick or thin but usually fleshy,
central, the apex typically not powdery or dandruffy. VEIL present as a cobwebby or silky cortina
which often leaves hairs on stalk; fibrillose or slimy universal veil often present also. VOLVA
typically absent (but rimmed bulb sometimes present). SPORE PRINT rusty-brown to cinnamon-
brown or at times ochraceous-tawny. Spores round to elliptical, roughened, lacking an apical pore.
Cystidia usually absent on faces (sides) of gills but sometimes present on the edges. Cap cuticle
typically filamentous.

CORTINARIUS is the largest genus of gilled mushrooms, with an estimated 1000

species—many of them still unclassified. Cortinarii can be dry or slick, thin or thick, large
or small, chunky or tall, but as a group they are easy to recognize, for there are three funga-
mental features that unite all 1000+ species: rusty-brown to cinnamon-brown spores, the
presence of a cortina, and a terrestrial woodland growth habit.
The cortina, which is the hallmark of Cortinarius, is an exquisitely constructed veil of
silky or cobwebby fibers. It shrouds the gills of the young mushroom (COLOR PLATE
105), but collapses as the cap opens, often leaving hairs on the stem which are subsequently
stained rusty-brown by spores (COLOR PLATE 103). In some species the cortina
may completely disappear, but the combination of rusty-brown spores (and often, rusty-
brown mature gills) and growth on the ground is a fairly infallible indicator of Cortinarius.
In many species the cortina is augmented by a universal veil that leaves scaly belts, fibrils,
and/ or a sheath of slime on the stem (and sometimes the cap).
Among the brown-spored mushrooms, Cortinarius is most likely to be confused with
Inocybe and Hebeloma, which are terrestrial and may have a cortina, but which have
duller brown spores and often have whitish-edged gills and a powdery or dandruffy stalk
apex; with Phaeocollybia, which has rusty-brown spores but lacks a cortina; and with
Gymnopilus and Pholiota, both of which are primarily wood-inhabiting. (Cortinarii
can occasionally be found on wood in an advanced stage of decay, but you will never find
them on the mossy crotch of a live oak or the cut end of a felled fir.)
Within these parameters—presence of a cortina and often a universal veil, rusty-brown
to cinnamon-brown or “ochraceous-tawny” (whatever that may mean) spores, and
growth on the ground near or under trees—Cortinarii vary tremendously in shape, size,
color, and habit. Many are Boring Ubiquitous Mushrooms (“BUM’s”), but a sizable
number are big, bold, and blatantly beautiful. In fact, the Cortinarii, along with the
waxy caps (Hygrophoraceae), are the most colorful of all the agarics: blues, violets,
yellows, oranges, browns, and olives predominate, but reds, bright greens, and whites
are not rare. To make any headway in identification it is essential to know the color
of fresh individuals—particularly the color of the immature gills, since it changes as
soon as the spores begin to form. As Cortinarii grow older, the bright colors and
individuality of their youth are drowned in brownness, until the enormity of their
uniformity makes identification of species hopeless (on the other hand, the sameness of
their mundane-ness helps to signify that they are Cortinarii).*
Y ou should also note whether the cap and stalk are viscid when they arefresh and moist.
Scent is another important variable—many species have indifferent odors, but some are
enticingly aromatic (e.g., C. percomis), others are downright disgusting (e.g., C. cam-
phoratus), while still others are aromatic and disgusting (e.g., C. subfoetidus).
Even if you take meticulous notes on fresh specimens in all stages of development, you
will still find Cortinarii difficult to identify. Perhaps this is why professional mycologists

*Note: These lines were composed at 3:00 A.M. under the influence of severe boredom.
 CORTINARIACEAE
418

seldom bother to name the multitudes of Cortinarii they encounter, and why Elias Fries,
the “father” of mushroom taxonomy, said in 1838: No genus is more natural nor more
sharply distinguished from others . . . but the species are so intimately related among
themselves that to distinguish the separate ones is almost to be despaired of. Cortinarius
species deserve to be better known, however, so don’t let your inability to name them
prevent you from learning to recognize them. Naming them, after all, is not an end in itself,
but a means of expediting the recognition process. If you really must have labels, you can
give them descriptive nicknames of your own.
The labyrinthine process of identifying a Cortinarius can be shortened to some extent
by learning to place each species in its proper subgenus. Five (or sometimes seven) large
subgenera have traditionally been recognized, and even raised to the rank of genus by
some mycologists:
Subgenus Myxacium: both cap and stalk viscid or slimy.
Subgenus Bulbopodium: only the cap viscid; stalk with an abrupt, often rimmed basal
bulb (see photograph on p. 439 and Color Plate 105).
Subgenus Phlegmacium: only the cap viscid; stalk equal to club-shaped or swollen, but
lacking a rimmed bulb.
Subgenus Telamonia: neither the cap nor the stalk viscid; cap hygrophanous and
smooth or occasionally fibrillose.
Subgenus Cortinarius (now divided into four subgenera—see below): neither the cap
nor the stalk viscid; cap often fibrillose or scaly but not typically hygrophanous.

The three subgenera with viscid caps are the easiest to recognize. Myxacium is especially
distinctive by virtue of the slimy universal veil that coats the cap and stalk. Bulbopodium
and Phlegmacium feature many large and/ or colorful species but intergrade somewhat
(the bulb can be somewhat abrupt or slightly rimmed), causing some investigators to
recognize only the latter.
The hygrophanous-capped subgenus Telamonia is the largest and most difficult group
of Cortinarii, with at least 400 species—most of them brownish or cinnamon-brown and
many of them small and nondescript (in other words, typical“LBM’s” in the grand, bland
tradition of Inocybe, Galerina, and Tubaria).
Members of the dry-capped subgenus Cortinarius are now divided up by Cortinarius-
categorizers into four subgenera, based largely on the types of pigments they contain:
Subgenus Cortinarius (in its most restricted sense): contains a single striking deep
violet, fibrillose-scaly species (C. violaceus).
Subgenus Sericeocybe: has violet to pale violet or lilac-white colors.
Subgenus(or genus) Dermocybe: composed mainly of small, slender-stemmed, bright¬
ly colored, conifer-loving species with yellow, olive, orange, or red pigments called
anthraquinones (which are water-soluble and hence make wonderful dyeing agents).
Subgenus Leprocybe: a large and diverse group whose color ranges from olive to yellow,
yellow-brown, rusty-orange, or brown, but whose pigments differ from those found
in Dermocybe.

Cortinarius is one of the most prominent features of our woodland fungal flora, but its
fruiting behavior is notoriously erratic. Sometimes there is one sudden spectacular,
Russula-like eruption of Cortinarii; at other times they fruit in a rather uninspired,
desultory fashion throughout the rainy season. A few species, such as C. glaucopus, are
common every year, but many types will be exceedingly abundant one season, then rare
or absent for the next two or ten (or even twenty!) years.
Most if not all Cortinarii are mycorrhizal. They attain their maximum numbers and
diversity in the cool coniferous forests of the north temperate zone, but are also common
under hardwoods (particularly “Bulbopodiums” and “Phlegmaciums”). In our area each
Cortinarius crocolitus (see comments under C. collinitus, p. 431) is one of hundreds of Cortinarii. Its
stalk is marked by yellow-brown or ochre belts formed by the universal veil.

forest type offers its own characteristic clique of Cortinarii: e.g., C. infractus, C. cotoneus,
and C. collinitus with tanoak; C. glaucopus, C. regalis, and C. sodagnitus with live oak;
C. cinnamomeus, C. phoeniceus var. occidentalism and C. obtusus with pine; and C.
balteatus, C. cylindripes, and C. vibratilis with ericaceous trees and shrubs (huckleberry,
manzanita, and madrone). It is in northern latitudes or higher altitudes(i.e., with northern
conifers, especially spruce), however, that the Cortinarii come into their own, and their
astonishing diversity becomes overwhelming.

Some of the older popular mushroom manuals list Cortinarius as a “safe” (albeit
mediocre*) genus—but nothing could be farther from the truth. Cortinarius, in fact,
contains several deadly poisonous species (particularly in the subgenus Leprocybe—see
C. gentilisl) whose effects on the human body are greatly delayed (for up to three weeks!).
Therefore it is best not to eat ANY Cortinarius, especially in view of the large number of
poorly known or difficult-to-identify species. (Even some of those proclaimed to be edible
may not be safe, since there is no assurance that the eater identified them correctly.)
Cortinarius is a tempting potential food source, however, so if you are adamant about
experimenting, then stick to the ones with viscid caps (none of which, as yet, are known
to be deadly poisonous), and wait at least two weeks—rather than the usual two days—
after your first “test” before indulging in more.
Because Cortinarii are so prominent and colorful, the selection of species in this book
is fairly sizable, at least in comparison to other giant—but more mundane—genera such as
Inocybe and Psathyrella. Over 130 species of Cortinarius are mentioned (not all of them
native to California) and 34 are fully described. Remember, however, that these 130+
species represent only a fraction of the total number that occur. The fieldmarks listed are
often insufficient for positive identification, and many of the descriptions are meant to
serve as models or “archetypes” for a large number of species (e.g., brown “Telamonias”
with a club-shaped stalk and medium-sized cap are typified by C. laniger). Thus most of the
names in this chapter should be seen as suggestions—rather than statements—as to what
your mysterious Cortinarius might be. Those wishing to verify their identifications should
consult a more definitive source, such as Alexander Smith’s keys to North American
Cortinarii (which are not available to the public), or Daniel Stuntz’s key to species of the
Pacific Northwest and Meinhard Moser’s keys to those of Europe, which are (see Sug¬
gested Readings and Primary References).

*Even Captain Charles Mcllvaine, that intrepid turn-of-the-century toadstool-tester who sung the praises of such
flaccid, featureless, fleshless fungi as Coprinus ephemerus (“choice as a flavoring”) and Psathyrella candolleana
(“tender one of the best”), was non-plussed by the Cortinarii, for on more than one occasion he characterized their
flavor as “more or less that of rotten wood.” However, one colleague of mine has a dissenting interpretation of
Mcllvaine’s comparison: he reasons that Mcllvaine’s tastes (which included a fondness for stinkhorns) were so
bizarre and eccentric that “rotten wood” was undoubtedly high up on his list of“best edibles,” and consequently,
so were the Cortinarii!
420 CORTINARIACEAE

Key to Cortinarius
l. Fruiting body typically developing underground (but cap may surface in wet weather), the veil
membranous or somewhat membranous and persistently covering the gills (i.e., either re¬
maining intact through maturity or shredding radially but not rupturing); found mostly under
mountain conifers ..2
1. Not as above; if developing underground then veil not persistent and membranous .4
2. Cap purplish or lavender-tinged .C. velatus{see C. magnivelatus, p. 442)
2. Cap white to yellow, ochre, or yellow-brown (not purplish) .3
3. Cap whitish when fresh; veil tough, often not rupturing . C. magnivelatus, p. 442
3. Cap yellow to yellow-brown or tan (or rusty-stained); veil not as tough or thick as above, often
shredding radially .C. verrucisporus & others (see C. magnivelatus, p. 442)
4. Cap viscid or slimy when moist (it may dry out, but look for adhering debris) . 5
4. Cap not normally viscid (in wet weather it may be slightly viscid or tacky but never slimy) 61
5. Cap radially corrugated or coarsely wrinkled, yellow-brown to ochre, tawny, or rusty-brown;
gills purplish at first but soon brown to rusty-cinnamon; stalk with a basal bulb when young,
but bulb often obscure in age; found under hardwoods in eastern North America, often in
large numbers .C. corrugatus
5. Not as above .6
6. Fruiting body massive; stalk 3-7 cm thick and lacking a basal bulb; flesh also very thick; cap
brownish to russet at the center (often with small flattened scales) and yellowish at margin
(lacking pronounced violet tones) . C. ponderosus & others, p. 432
6. Not as above; either smaller or with violet on cap or stalk with a basal bulb .7
7. Stalk (at least the lower part) viscid or slimy when moist, sometimes enlarged below but typically
without an abrupt, rimmed basal bulb (in very wet or very dry weather the viscidity may not be
evident) . 8
7. Stalk typically not viscid (but if cap is slimy, some of the slime may drip off onto stalk base);
stalk with or without an abrupt, rimmed basal bulb . 21
8. Fruiting body small (cap 2-5 cm broad) and gray, the cap and young gills with a slight lilac tinge
that often disappears in age; stalk whitish with a bluish-tinged apex when young; found under
conifers; rather rare. C. sterilis
8. Not as above .9
9. Fruiting body purple, violet, or bluish in some part, at least when young and fresh . 10
9. Violet or bluish shades completely absent . 17
10. Stalk typically club-shaped (thicker at base) . 11
10. Stalk equal, tapered downward, or spindle-shaped (swollen slightly in the middle and tapered
below) . 13
11. Cap violet or lavender well into maturity C. iodes& others (see C. cylindripes group, p. 430)
11. Not as above . 12
12. Center of cap rusty or tawny becoming chestnut-brown in age, the margin paler; gills violet-
tinged when very young . C. castaneicolor (see C. cylindripes group, p. 430)
12. Not as above .C. griseoluridus & others (see C. cylindripes group, p. 430)
13. Lower portion of stalk with conspicuous transverse or concentric bands, plaques, or scaly belts
formed by the universal veil; cap yellow-brown to rusty-orange or rusty-brown at maturity
(margin may be bluish or violet when young) . C. collinitus group, p. 431
13. Not as above (stalk occasionally with some scaly belts, but if so then cap darker or differently
colored) . 14
14. Cap purple or lilac, at least when young .C. cylindripes group & others, p. 430
14. Not as above . 15
15. Cap blackish to deep chestnut-brown to dark olive-brown, often fading to cinnamon-brown or
yellowish-brown in age, the margin often becoming radially wrinkled or grooved; stalk long,
usually rooting deeply in soil; common under northern conifers . 16
15. Not as above; cap differently colored, at least when young (i.e., paler or brighter) .
. C. pseudosalor & C. delibutus (see C. cylindripes group, p. 430)
CORTINARIUS 421

16. Cap blackish to deep chestnut-brown when young, fading to cinnamon in age; gills not violet
or bluish when young .C. vanduzerensis, p. 432
16. Cap colored as above or ranging to olive- or yellow-brown; gills violet or bluish-tinged when
young . C. elatior(see C. collinitus group, p. 431)
17. Stalk usually club-shaped or thickened below, at least when young; fruiting body sometimes
rather small . 18
17. Stalk more or less equal or tapered downward; fruiting body medium-sized or larger .... 20
18. Fruiting body medium-sized (cap usually 5 cm broad or more); stalk usually at least 1 cm thick
at apex, typically with ochre-brown patches of veil tissue below; odor and taste not distinctive
.C. pallidifolius(see C. collinitus group, p. 431)
18. Not as above; fruiting body rather small (cap usually less than 5 cm broad); odor typically
fragrant or taste of cap surface bitter . 19
19. Odor fragrant; flesh, young gills, and stalk yellow . . C. ciirinifolius(see C. percomis, p. 436)
19. Odor typically mild but taste very bitter (at least of the cap surface); flesh and stalk whitish
.C. vibratilis, p. 429
20. Lower portion of stalk typically with transverse or concentric bands, plaques, or scaly belts
of universal veil tissue; found under both hardwoods and conifers C. collinitus group, p. 431
20. Not as above; associated principally with conifers . C. mucosus, p. 429
21. Immature gills sooty olive to olive, greenish, or yellow-green, or soon becoming these colors 22
21. Immature gills differently colored (including yellow) . 28
22. Stalk lacking an abrupt, rimmed basal bulb; cap sooty-olive when young (may be browner or
tawnier in age), the skin often bitter-tasting; gills usually sooty-olive also .
. C. infractus& others, p. 435
22. Not as above; stalk usually with an abrupt, often rimmed basal bulb, at least when young 23
23. Immature gills with violet faces or entire gills with a fleeting violet phase at first .24
23. Gills never violet .25
24. Gills at first with violet faces and olive edges C. montanus& others(see C. cedretorum, p.439)
24. Gills violet-tinged when very young but soon entirely olive to greenish or greenish-yellow . . .
.C. scaurus group, p. 440
25. Cap reddish to reddish-brown or orange-brown when fresh, at least at the center .
.C. orichalceus& C. rufo-olivaceus (see C. scaurus group, p. 440)
25. Not as above ..'. 26
26. Cap olive; gills olive to olive-yellow; stalk yellowish or olive-tinged; found mainly under conifers
in the Pacific Northwest . C. prasinus (see C. scaurus group, p. 440)
26. Not as above; usually more brightly colored, at least in part; widespread .27
27. Gills green; stalk pale bluish when fresh; found in eastern North America under hardwoods
.C. virentophyllus (see C. scaurus group, p. 440)
27. Not as above; especially common in the West .C. scaurus group, p. 440
28. Some part of fruiting body lilac, violet, purple, or bluish when fresh and young(check cap sur¬
face, stalk, immature gills, and flesh) .. 29
28. Fruiting body completely lacking violet, lilac, or bluish tones .48
29. Flesh (at least in base of stalk) staining wine-red when bruised . 72
29. Not as above (but flesh may stain purple or dull lilac) . 30
30. Gills and/or flesh staining purple or dull lilac when bruised (if already purple, then staining
darker purple) . C. mutabilis group & others, p. 437
30. Not as above . 31
31. Immature gills yellow or purplish with yellow or olive edges . 32
31. Not as above . 33
32. Flesh partly violet or lilac and partly yellowish or whitish when fruiting body is sliced open
lengthwise (see Color Plate 105); stalk apex not bluish.C. cedretorum & others, p. 439
32. Not as above; stalk apex often bluish; gills entirely olive or with olive edges .
.C. montanus (see C. cedretorum, p. 439)
33. Whitish fibrillose or felty veil remnants usually present on cap (often in the form of a patch)
.C. calyptratus & C. calyptrodermus (see C. rega/is, p. 443)
33. Not as above . 34
422 CORTINARIACEAE

34. Stalk with an abrupt, rimmed basal bulb, at least when young .
34. Stalk merely equal or club-shaped, lacking a sharply defined, rimmed basal bulb
35. Basal bulb with a whitish volva- like sheath; cap grayish to bluish-gray; growing mainly under
spruce . .C. volvatus(sQQ C. regalis, p. 443)
35. Not as above . 36

36. Cap bright yellow to yellow-brown or ochre (the center sometimes oranger), even when young
and fresh.C. calochrous & others (see C. fulmineus, p.441)
36 Cap differently colored (but may develop ochre or yellowish shades in age) .37
37. Cap dark reddish-brown to chestnut, becoming paler reddish-brown in age; gills pale lilac at
first, darkening to reddish-brown; found mainly under conifers, especially in southern Oregon
and northern California .C. subpurpureophyllus
37. Not as above . 38
38. Cap dark olive or greenish to steel-gray or brown, often streaked or mixed with ochre, blue, or
gray, and often developing rusty to fulvous tones when older (especially toward center);
immature gills gray to bluish-gray to bluish-violet (or occasionally violet) .
.C. glaucopus group & others, p. 437
38. Not as above; immature gills lilac, violet, or lavender, or if bluer then cap typically bluish to
bluish-gray or at least differently colored from above. 39
39. Associated with conifers in the Pacific Northwest; fruiting body modest in size (stalk typically
less than 1.5 cm thick at apex); gills lilac to pinkish-lilac when young . C. olympianus, p. 439
39. Not as above; especially common under hardwoods .40
40. Fruiting body (or at least the immature gills and stalk apex) with a bluish tinge when fresh
... C. caesiocyaneus (see C. olympianus, p. 439)
40. N ot as above (gills and stalk apex more violet or purple than blue) .41
41. Mature fruiting body with a small but sharply defined basal bulb; taste usually bitter (at least
of the cap cuticle); cap with violet or lilac tones at least at the margin; common under oak in
California .C. sodagnitus group, p.438
41. Not as above; found mainly under hardwoods in eastern North America .42
42. Cap and basal bulb discoloring ochre or buffy-tan fairly quickly .
.C. velicopia(see C. olympianus, p.439)
42. Not as above . C. michiganensis, C. aggregatus, & others (see C. olympianus, p. 439)
43. Cap usually lilac or lavender at least at the margin; stalk usually very thick (2-5 cm) or if not then
odor usually sweetish (like overripe pears) . 44
43. Not as above .45
44. Stalk very thick (2-5 cm); flesh thick and firm .C. balteatus&i others, p. 433
44. Stalk typically 2 cm thick or less; odor usually sweetish .C. subfoetidus, p. 434
45. Stalk with pale tawny patches or fibrils, thinly viscid at first; cap tawny to chestnut-brown with
a paler (buff-colored) margin . C. castaneicolor (see C. cylindripes group, p. 430)
45. Not as above .46
46. Cap yellow to yellow-brown, tawny, or fulvous C. varius & others (see C. multiformis, p. 442)
46. Cap differently colored . 47
47. Cap reddish-brown to liver-brown or dull purplish; gills usually violet or lilac when young . .
. C. variicolor & others (see C. multiformis, p. 442)
47. Not as above; gills usually bluer or grayer . C. glaucopus group, p. 437
48. Odor distinctly fragrant and sweet (break open the flesh if there is any doubt) .49
48. Not as above (but odor may be distinctive) . 51
49. Odor aniselike; stalk with a basal bulb, at least when young C. odorifer(see C. percomis, p. 436)
49. Not as above . 50
50. Stalk and flesh yellow or yellowish .C. percomis, p. 436
50. Stalk and flesh white or whitish .C. luteoarmillatus (?) (see C. percomis, p. 436)
51. Odor strongly pungent(like green corn or corn silk); gills and cap yellowish when young, thecap
becoming browner or redder in age; stalk usually with darker fibrillose patches or zones; found
under conifers in the Pacific Northwest .C. superbus
51. Not as above . 52
CORTINARIUS 423

52. Gills yellow to yellow-orange or yellow-brown when young and stalk with an abrupt, usually
rimmed basal bulb (at least when young) . 53
52. Not as above . 54
53. Cap bright green to citrine-green when fresh (sometimes brownish at the center); common
under oak in California .C. sp. (unidentified) (see C. scaurus group, p. 440)
53. Cap not green . C. fulmineus & others, p. 441
54. Cap brownish to dingy-flesh colored, but usually with a patch or patches of white fibrillose
or felty veil material, only slightly viscid; stalk with a basal bulb; found under oak in California
.C. regalis, p. 443
54. Not as above . 55
55. Cap white or whitish (or at times ranging to buff or pale tan) . 56
55. Cap darker or brighter than above . 57
56. Stalk with a basal bulb; found under hardwoods in eastern North America .C. albidus
56. Not as above; stalk equal or club-shaped (thicker below) but lacking a prominent basal bulb;
often found under conifers . 138
57. Lower portion of stalk with concentric belts or transverse bands of colored (usually ochre)
veil tissue .C. crocolitus(see C. collinitus group, p. 431)
57. Not as above . 58
58. Cap radially corrugated or wrinkled, cinnamon to pinkish-cinnamon to cinnamon-brown;
found in the Pacific Northwest under conifers ..C. corrugis
58. Cap not radially corrugated . 59
59. V eil copious, usually leaving remnants on cap margin and/ ora fibrillose white sheath or patches
on the stalk (and often a slight ring or annulus) .... C. turmalis{see C. multiformis, p. 442)
59. Not as above; veil not so copious (but may leave a few remnants) .60
60. Stalk 1-4 cm thick and very firm, usually stout, more or less equal or narrowed slightly at the
base .C. crassus (see C. balteatus, p. 433)
60. Stalk with a basal bulb (at least when young) or not as thick and stout as above.
...C. multiformis & others, p. 442
61. Fresh fruiting body at least partially purplish, bluish, lilac, lavender-gray, lavender-white,
etc. (check immature gills, stalk apex, flesh, and cap) .62
61. Fresh fruiting body completely lacking violet, bluish, or pale lavender shades .87
62. Lower part of stalk with reddish bands, patches, or “bracelets” formed by the universal veil
or stalk with a fiery orange to red base .. .. 63
62. Not as above (but upper stalk may have a ring of rusty hairs from the cortina) . 64
63. Stalk 3-7 mm thick at apex, with reddish to vinaceous patches, fibrils, or “bracelets”; cap 2-5
cm broad .C. boulderensis (see C. armillatus, p. 448)
63. Usually larger than above; stalk with a bright orange or red base .C. rubripes, p. 449
64. Stalk typically with a persistent, somewhat membranous annulus (ring) formed by the veil,
usually club-shaped or bulbous; gills broad, well-spaced, deep purplish becoming dark
cinnamon-brown; cap deep purplish to brownish to copper-brown, often fibrillose or hoary;
found mainly under hardwoods in eastern North America C. torvus (see C. evernius, p. 450)
64. Not as above . 65
65. Cap hygrophanous (fading markedly as it loses moisture) and smooth (bald) or at most with a
few whitish veil remnants near the margin . 66
65. Cap smooth, silky, fibrillose, hairy, or scaly, but if smooth then not markedly hygrophanous 72
66. Fruiting body small (cap 1-3 cm broad; stalk less than 5 mm thick) .
.C. pulchellus & C. subflexipes (see C. obtusus group, p. 452)
66. Fruiting body medium-sized (larger than above). 67
67. Fresh fruiting body violet-tinged only at apex of stalk . . . C. brunneus (see C. laniger, p. 451)
67. Fresh fruiting body showing more violet than above (but violet tones may fade) . 68
68. Stalk with conspicuous remnants from the universal veil . 69
68. Stalk with inconspicuous veil remnants (or lacking them) .
. C. impennis & C. adustus(see C. evernius, p. 450)
Cortinariuspholideus occurs mostly with birch. Note prominent scales on cap and stalk.

69. Universal veil remnants on stalk yellowish to buff . C. subpurpureus (see C. evernius, p. 450)
69. Not as above . 70
70. Stalk more or less club-shaped (thicker at base) .C. lucorum (see C. evernius, p. 450)
70. Stalk equal or tapering downward . 71
71. Stalk often with a narrow, flaring annulus (ring); cap pale when fresh .
.C. urbicus (see C. evernius, p. 450)
71. Not as above .C. evernius & others, p. 450
72. Flesh bluish or violet but staining wine-red to wine-brown when cut or bruised, at least in base of
stalk; cap bluish or bluish-gray to grayish-lavender becoming deep purplish-brown in age;
cap surface usually fibrillose or with flattened scales; gills dark bluish or bluish-violet when
young; stalk with a basal bulb which often disappears in age; not common .C. cyanites
72. Not as above; flesh not staining wine-red when bruised; common. 73
73. Cap distinctly hairy, scaly, or fibrillose-scaly . 74
73. Cap smooth or silky or very slightly scaly (but may have universal veil remnants on it) ... 78
74. Stalk with only a tinge of violet or bluish-gray and soon fading to whitish; cap brownish to buff
beneath a white to grayish-white fibrillose-hairy layer (cap lacking violet or bluish shades);
found under conifers .C. plumiger (see C. evernius, p. 450)
74. Not as above . 75
75. Entire fruiting body deep violet (sometimes nearly black) when fresh (the gills may be rusty-
dusted in age) .C. violaceus, p. 446
75. Not as above; fruiting body paler, browner, or with only a tinge of violet . 76
76. Cap and stalk with brown to cinnamon-brown scales; stalk not radically bulbous (see above
photo); fairly common under hardwoods (especially birch) in eastern North America, rarely
in the Pacific Northwest .C. pholideus (see C. squamulosus, p. 445)
76. Not with above features; stalk not usually conspicuously scaly . 77
77. Stalk radically bulbous; cap with brown to dark chocolate-brown scales; found under hard¬
woods in eastern North America, rare (or absent) in West .C. squamulosus, p. 445
77. Not as above; stalk not radically bulbous and/or habitat different . 78
78. Flesh in stalk rusty-brown to cinnamon-, tawny-, or yellow-brown (often marbled); associated
with conifers . C. traganus, p. 447
78. Not as above (flesh in stalk usually white to buff or tinged violet) . 79

424
Cortinarius bolaris favors beech and other hardwoods in eastern North America. Note distinctive
red fibrils or scales on the cap and stalk.

79. Stalk with buff to tan, ochre, or brownish-yellow patches, zones, or “bracelets” formed by
the universal veil .C. anomalus & C. caninus (see C. alboviolaceus, p. 447)
79. Not as above; universal veil remnants differently colored (if present) . 80
80. Odor distinctive: either sweetish (like overripe pears), spermatic, or very unpleasant.81
80. Not as above; odor mild, radishlike, or merely fungal . 83
81. Odor sweetish, resembling overripe pears .C. fragrans (see C. traganus, p. 447)
81. Odor unpleasant or spermatic . 82
82. Odor usually spermatic; fruiting body rather small, the cap often umbonate; spore print dull
brown .(see Inocybe, p. 455)
82. Odor unpleasant but not spermatic; fruiting body medium-sized; spore print rusty-brown to
cinnamon-brown .C. camphoratus & others (see C. traganus, p. 447)
83. Stalk very thick (2-5 cm thick at apex) and firm; cap viscid when moist (but soon dry and often
shiny) .C. balteatus, p. 433
83. Stalk typically 2 cm thick or less at apex and/ or cap not viscid when young . 84
84. Cap and lower stalk viscid when very young, often with brown or olive-brown patches or spots
(of dried slime) when older; found under conifers; rare .
.C. griseoviolaceus (see C. cylindripes group, p. 430)
84. Cap and stalk not viscid (or cap only very slightly so in wet weather) and not spotted as above
(but may develop ochre, tan, buff, or brownish discolorations in age); found under both
hardwoods and conifers; common . 85
85. Cap predominantly pale violet to lilac-white or silvery-violet or pale bluish-violet (but buff to
ochre stains may develop in age) . 86
85. Cap differently colored (either grayish-buff or soon becoming brown to ochre throughout)
. C. subpulchrifolius & C. malachius (see C. alboviolaceus, p. 447)
86. Lower portion of stalk sheathed with conspicuous white universal veil tissue (fibrils); stalk often
rather long and club-shaped (thicker below but not abruptly bulbous) C. alboviolaceus, p. 447
86. Stalk often thick and stout with a prominent basal bulb and/or lacking conspicuous white
universal veil tissue .C. argentatus & C. subargentatus (see C. alboviolaceus, p. 447)
87. Growing in sand (but associated with pine); gills rusty-orange to brownish-orange when young
...C. aureifolius (see C. cinnamomeus group, p. 453)
87. Not with above combination of characteristics . 88
88. Cap olive-brown to olive-yellow to yellow-brown with small darker (often blackish) hairs
or fibrillose scales, especially toward center . C. cotoneus group, p. 445
88. Not as above . 89
89. Cap and stalk with reddish scales, fibrils, and/or streaks on a paler (whitish to yellowish)
background; found under hardwoods, especially beech, in eastern North America (see above
photo) . C. bolaris (see C. squamulosus, p. 445)
89. Not as above (but cap and stalk can be entirely reddish or the stalk can have reddish universal
veil remnants) ... 90
425
 426 CORTINARIACEAE

90 Stalk white to brownish with 1 -4 reddish to reddish-brown bands or belts of universal veil tissue
over the lower portion or stalk with a bright red to orange base or a sheath of vermilhon fibrils
(universal veil remnants) .^1
90. Not as above (but stalk may be entirely red or orange or the stalk may have a single zone of rusty
hairs formed by the cortina, usually near apex) .94
91. Cap 1 -3 cm broad; stalk wholly or partially sheathed by vermillion fibrils .
.C. miniatopus (see C. rubripes, p. 449)
91. Not as above; cap usually larger . 92
92. Stalk base red to bright orange; associated with hardwoods, especially oak C. rubripes, p. 449
92. Stalk with 1-4 reddish bands or “bracelets”; associated with conifers or birch . 93
93. Typically associated with birch .C. armillatus, p. 448
93. Typically associated with conifers (especially in the Pacific Northwest) or with hardwoods
other than birch .C. haematochelis & C. subtestaceus (see C. armillatus, p. 448)
94. Stalk 6-18 cm long and 0.5-2 cm thick at apex, usually whitish and club-shaped (thicker at base);
gills pale yellow when young; cap creamy to pale buff to yellowish or tawny-brown (often
darkest at center), silky or minutely scaly; found under hardwoods (especially beech) in eastern
North America . C. flavifolius
94. Not as above . 95
95. Growing on lawns or buried wood; stalk typically 3-5 mm thick and often with a small annulus
(ring); cap typically less than 4 cm broad, reddish-brown to cinnamon, fading to more or less
buff as it loses moisture; found in the Pacific Northwest .
.(see Galerina, Tubaria, & Allies, p. 399)
95. Not as above . 96
96. Stalk yellow to bright yellow but the gills whitish to brown (i.e., not brightly colored) when
young; cap smooth and hygrophanous, fading to yellowish as it loses moisture . 97
96. Not as above; stalk not yellow (but may have yellowish veil remnants), or if stalk yellow then
immature gills usually brightly colored also or cap minutely scaly . 98
97. Gills whitish when young; cap brown to tan when moist .C. renidens
97. Gills brownish or cinnamon when young; cap bronzy-brown to deep olive-brown when moist
. C. angulosus
98. Gills colorful (yellow, greenish, orange, red, dark red, or bright rusty-orange) when young;
stalk often colorful also; cap usually not hygrophanous . 99
98. Gills dull (whitish, grayish, brownish, or dull cinnamon) when young; stalk usually dull also
(brownish, gray, white, etc.); cap usually hygrophanous . 110
99. Immature gills orange-red to rusty-orange; cap hygrophanous (dark rust-red but fading as it
loses moisture); stalk usually paler than cap and gills; found only on the west coast in mixed
woods and under conifers.C. californicus (see C. phoeniceus var. occidentalis, p. 454)
99. Not as above (if colored as above, then cap not hygrophanous or found elsewhere) .... 100
100. Immature gills red to dark red . 101
100. Immature gills some shade of yellow, olive, orange, or bright cinnamon . 104
101. Entire fruiting body orange-red to red, dark red, or rust-red . 102
101. At least the stalk (and sometimes cap) yellowish or ochre to yellow-brown . 103
102. bruiting body cinnabar-red to rusty-red or dark red; found mainly in eastern North America,
especially under hardwoods .C. cinnabarinus & others (see C. sanguineus, p. 454)
102. Fruiting body blood-red to dark red; found mainly under conifers in northern and western
North America . C. sanguineus, p. 454
103. Cap red to dark red or reddish-brown .C. phoeniceus var. occidentalis, p. 454
103. Cap ochre to brown or yellowish .
. C. semisanguineus (see C. phoeniceus var. occidentalis, p. 454)
104. Cap orange-red to bright rusty-red to coppery-red (or occasionally orange-brown); immature
gills bright yellow to greenish-yellow or sometimes orangish; associated mainly with conifers
and willow; not common .C. uliginosus
104. Not as above; cap not reddish . 105
105. Cap tawny to rusty-orange and covered with numerous minute, erect scales .
.. rainierensis (see C. gentilis, p. 444)
105. Not as above; cap smooth to fibrillose or with a few scales only . 106
CORTINARIUS 427

106. Cap smooth and hygrophanous, orange-brown to yellow-brown or ochre when moist, fading
as it loses moisture; gills ochre-yellow to cinnamon-brown or brown (not particularly bright);
stalk deep orange-brown or cinnamon-colored, often with a few yellow veil remnants when
young; fruiting body small (stalk less than 5 mm thick) .C. gentilis, p. 444
106. Not as above; cap smooth to fibrillose or slightly scaly, not hygrophanous; fruiting body
variously colored but often brighter or more olive than above (especially the stalk) .... 107
107. Immature gills yellow to olive-yellow or greenish . 108
107. Immature gills saffron (orange-yellow) to orange or rusty-orange . 109
108. Stalk club-shaped and at least 2.5 cm thick at base; fruiting body bright yellow becoming
oranger or browner in age . C. callisteus (see C. cinnamomeus group, p. 453)
108. Not as above; stalk more slender and/or equal; fruiting body sometimes with olive tones
(but sometimes without) . C. cinnamomeus group & others, p. 453
109. Cap 3-9 cm broad; stalk usually 0.5-1.5 cm thick; gills ochre to tawny or orangish-cinnamon
when young; occurrence in North America questionable .
. C. orellanus & C. speciosissimus (see C. gentilis, p. 444)
109. Not as above; usually smaller or more slender; gills sometimes brightly colored (saffron,
orange, etc.); common in North America .
.C. croceofolius & others (see C. cinnamomeus group, p. 453)
110. Cap and stalk whitish to grayish or pale tan, but the flesh staining yellow when bruised and
finally turning (or aging) reddish; widely distributed, but especially common along the west
coast under conifers . C. rubicundulus (-C. pseudobolaris)
110. Not as above . Ill
111. Fruiting body small and rather fragile; cap sometimes umbonate, averaging 1 -4 (5) cm broad;
stalk averaging 2-6 mm thick (examine several specimens if unsure!) . 112
111. Fruiting body medium-sized or larger; cap not umbonate or only very broadly so, averaging
4-15 cm broad; stalk averaging 0.5-3 cm thick (check several specimens if unsure) . 121
112. Stalk usually with yellow or yellowish veil remnants, at least when young (check several speci¬
mens); gills ochre-yellow to cinnamon-brown when young .C. gentilis, p. 444
112. Not as above . 113
113. Gills staining black when bruised . C. washingtonensis (see C. obtusus group, p. 452)
113. Not as above . 114
114. Cap with a prominent black umbo (otherwise slightly paler) .
.C. nigrocuspidatus (see C. obtusus group, p. 452)
114. Not as above (but cap may be entirely blackish) . 115
115. Cap very dark when moist (deep vinaceous-brown to blackish-brown or black), but often paler
as it loses moisture . 116
115. Cap paler than above when moist . 119
116. Stalk with conspicuous white universal veil material, at least when young and fresh ... 117
116. Stalk with only slight universal veil remnants (if any) . 118
117. Odor geranium-like (at least when flesh is crushed); found in bogs or other wet places .
.C. paleaceus (see C. obtusus group, p. 452)
117. Not as above .C. stemmatus (see C. obtusus group, p. 452)
118. Cap blackish-brown when moist; gills and stalk also very dark; found under conifers .
. C. uraceus
118. Cap dark vinaceous-brown when moist; gills tawny-yellowish to pale brownish when young
. C. decipiens (see C. obtusus group, p. 452)
119. Cap (and sometimes stalk) minutely scaly .
.C. psammocephalus & C. incisus{see C. obtusus group, p. 452)
119. Cap more or less smooth . 120
120. Cap typically with an acute (sharp) umbo C. acutus & others (see C. obtusus group, p. 452)
120. Cap typically with an obtuse (blunt) umbo or with none at all C. obtusus group & others, p.452
121. Fruiting body (including stalk) rusty-orange to yellowish or yellow-brown; cap minutely scaly
and not hygrophanous; not definitely known to occur in North America .
. C. orellanus & C. speciosissimus (see C. gentilis, p. 444)
121. Not as above; very common in North America . 122
428 CORTINARIACEAE

122. Cap brownish to dingy flesh-colored, but often with a whitish patch of universal veil tissue;
stalk thick (1.5-3.5 cm at apex), with a large, abrupt, often rimmed basal bulb; fairly common
in California under oak . ^ regalis, p. 443
122. Not as above . 123
123. Cap covered with cinnamon-brown to chocolate-brown scales, not markedly hygrophanous;
either stalk also with scales or stalk radically bulbous; common under hardwoods in eastern
North America, rare in the West . 124
123. Not as above . 125
124. Stalk radically bulbous and smooth to fibrillose or only slightly scaly C. squamulosus, p. 445
124. Stalk equal or slightly thicker below, with brown scales like those on cap .
.C. pholideus (see C. squamulosus, p. 445)
125. Cap broadly cone-shaped and finely silky-scaly from the universal veil when young; stalk
equal, sheathed with white universal veil remnants, soon becoming hollow; cap dark to pale
cinnamon .C. hemitrichus
125. Not as above (but may be similar in some respects, such as color). 126
126. Cap blackish-brown when moist; gills and stalk also very dark; found under conifers .
. C. uraceus
126. Not as above; usually paler or brighter in color . 127
127. Stalk typically tapered downward or spindle-shaped (swollen near ground and tapered below),
the base often rooting . 128
127. Stalk typically equal to club-shaped or bulbous . 130
128. Cap cinnamon-brown when moist ..C. duracinus (see C. laniger, p. 451)
128. Cap vinaceous-brown to cocoa-colored when moist . 129
129. Stalk with conspicuous white universal veil remnants C. damascenus (see C. laniger, p. 451)
129. Universal veil remnants on stalk absent or evanescent .
.C. privignus & C. cacao-color (see C. laniger, p. 451)
130. Cap silky to fibrillose or fibrillose-scaly and white to silvery-gray or pale brownish-gray or
cap brown beneath a whitish to grayish fibrillose layer. 131
130. Cap darker, at least when moist, and usually more or less smooth or minutely scurfy (but cap
may fade to buff or paler as it loses moisture) . 133
131. Cap brown to buff beneath a fibrillose-scaly layer . ... C. plumiger (see C. evernius, p. 450)
131. Cap paler than above and silky to fibrillose (but sometimes becoming pale brownish-gray in
age) . 132
132. Cap silky-fibrillose, white to silvery, sometimes becoming pale brownish in age .
.C. pinetorum (see C. laniger, p. 451)
132. Cap whitish or silvery, usually with a faint hint of lavender or violet (examine several specimens
if unsure), silky; flesh and stalk apex often with a violet tinge also .
. C. alboviolaceus & others, p. 447
133. Cap yellow-brown when moist, fading to pale yellow-orange or orange-buffasit loses moisture
.C. armeniacus (see C. laniger, p. 451)
133. Cap redder or darker than above, at least when moist . 134
134. Gills widely spaced; cap smooth or scurfy .C. distans (see C. laniger, p. 451)
134. Not as above . 135
135. Stalk more or less equal .C. biformis& C. dilulus (see C. laniger, p. 451)
135. Stalk usually club-shaped or swollen below, at least when young (but sometimes equal, so
examine several specimens if unsure) . 136
136. Stalk with conspicuous universal veil remnants, at least when young . 137
136. Stalk with only slight traces of the universal veil, if any (but may have remnants from the
cortina) . C. triformis & others (see C. laniger, p. 451)
137. Cap dark or dull brown when moist; stalk similarly colored beneath whitish to brownish veil
remnants, the apex sometimes vinaceous- or violet-tinged C. brunneus (see C. laniger, p. 451)
137. Cap cinnamon to reddish-brown or chestnut-brown when moist; stalk similarly colored be¬
neath whitish veil remnants, not vinaceous- or violet-tinged at apex C. laniger & others, p. 451
138. Cap surface or cuticle bitter-tasting .C. crystallinus (see C. vibratilis, p. 429)
138. Not as above .C. lustratus & others
Cortinarius vibratilis. Note small size, slimy cap, and viscid stalk that is swollen at base.

Cortinarius vibratilis (Bitter Cortinarius)

CAP 1. 5-5 cm broad, broadly bell-shaped to convex to nearly plane; surface smooth, viscid
or slimy when moist, yellowish to tawny, fulvous, orange-brown, or yellow-orange (or in
one form brown) when fresh, fading as it ages or dries (sometimes to pale tan). Flesh thin,
whitish; odor mild, taste bitter (especially cap surface). GILLS close, notched to adnate
or slightly decurrent, at first whitish, soon becoming dull ochre or tawny and finally cinna¬
mon-brown. STALK 3-7 cm long, 3-10 mm thick at apex, usually clearly thickened below
(club-shaped) but sometimes equal; viscid or slimy when moist, but in age sometimes viscid
only at the base; white, smooth or nearly so. UNIVERSAL VEIL slimy, colorless.
CORTINA scanty, sometimes leaving a few hairs on stalk which trap falling spores.
SPORE PRINT rusty-brown; spores 7-9 * 4-5 microns, elliptical, roughened.
HABITAT: Solitary to widely scattered or in small groups in forest humus; widely distri¬
buted and fairly common, but rarely fruiting in large numbers. It usually occurs with
conifers, but in our area I have collected it under manzanita, madrone, and huckleberry,
in December.
EDIBILITY: Inedible due to the bitter taste. (If Russula brevipes is “better kicked than
picked,” then C. vibratilis is “better kicked than licked!”)
COMMENTS: The small size, viscid tawny cap, and viscid white stalk plus the bitter taste
—which can usually be detected by pressing your tongue to the surface of the cap—are
the hallmarks of this petite Cortinarius. It typifies a group of species within the subgenus
Myxacium which have a club-shaped (thickened) stalk, at least when young. Other species:
C. crystallinus is similar in size, shape, and bitter taste but is whitish to buff or pale tan
overall and has a non-viscid stalk. See also C. citrinifolius (under C. percomis).

Cortinarius mucosus (Slimy Cortinarius)

CAP 4-10 cm broad, convex to plane or broadly umbonate, or even uplifted in age; surface
smooth and very slimy when moist, chestnut- to reddish-brown or bright orange-brown,
often paler (tawny or ochre) in age. Flesh whitish; odor and taste mild. GILLS close,
usually more or less adnexed, whitish to grayish when very young, becoming pale ochra-

429
 CORTINARIACEAE
430

ceous or tawny, then cinnamon-brown from ripening spores. STALK 4-15 cm long, 1-2
(2.5) cm thick, more or less equal, slimy or viscid when moist, white or whitish but. some¬
times rusty-stained from spores; not breaking up into conspicuous scaly belts. UNI¬
VERSAL VEIL whitish, slimy, disappearing or leaving indistinct remains on stalk.
CORTINA usually forming a superior, hairy-fibrillose zone on stalk which is stained rusty-
brown by spores. SPORE PRINT rusty-brown; spores 11-18 x 5-7.5 microns, elongated-
elliptical, roughened.
HABITAT: Widely scattered to gregarious in mixed woods and under conifers (parti¬
cularly pine and spruce); widely distributed, but especially common in the West in the late
summer and fall, and in the southern United States. I have seen large fruitings in Wash¬
ington, New Mexico, Arizona, and northern California, but not in our area. Cortinarius-
classifier Joe Ammirati says it has been found under birch and willow in the Brooks Range
in northern Alaska; it also occurs with hemlock.
EDIBILITY: Better eschewed than stewed. It is too slippery to be worthwhile, and its
edibility is unknown.
COMMENTS: The smooth, viscid, tawny to orange-brown or reddish-brown cap and
viscid, whitish, equal stalk are the distinguishing features of this “slimy sucker.” The cap
color is reminiscent of C. collinitus, but the stalk lacks the scaly belts characteristic of that
species, and the taste is not bitter nor the stalk club-shaped as in C. vibratilis. Phaeocollybia
species are somewhat similar, but lack a cortina and have a long, rooting stalk. C. turmalis
(see comments under C. multiformis) is also similar, but has a dry stalk.

Cortinarius cylindripes group (Slimy Purple Cortinarius)

CAP 3-12 cm broad, convex or broadly bell-shaped becoming broadly umbonate to plane
or with an uplifted margin; surface smooth or in age sometimes wrinkled, viscid to ex¬
tremely slimy when moist, at first lavender or violet, but fading quickly or slowly (often
from center outward) to yellowish, brownish-ochre, tawny, brown, or paler. Flesh
rather thin, violet-tinged becoming whitish; taste mild. GILLS at first lavender or grayish-
lavender, soon grayish or grayish-cinnamon with whitish, finely scalloped edges, finally
rusty-cinnamon; adnate to adnexed or notched, close. STALK 5-15 cm long, 1-2.5 (3) cm
thick, equal or at times spindle-shaped when young (swollen slightly in the middle); viscid
or slimy throughout or at least over lower portion; violet to silvery-violet or lavender in part
or throughout, but often fading somewhat (like cap) in age; smooth or with whitish to
violet scales above. UNIVERSAL VEIL slimy, violet, forming a slime sheath on stalk.
CORTINA obscured by slime, evanescent or leaving a few hairs on stalk. SPOREPRINT
dark cinnamon-brown; spores 12-17 * 5.5-8 microns, elliptical, roughened.
HABITAT: Scattered to densely gregarious in woods; widely distributed and locally
abundant (along with similar species) in the late fall and winter. Like C. balteatus, it tends to
fruit with ericaceous shrubs and trees, e.g., manzanita, madrone, and/ or huckleberry.
EDIBILITY: Better eyed than fried; the snotlike universal veil will deter most.
COMMENTS: This species may not be the most distinctive or desirable of our local
Cortinarii, but it unquestionably qualifies as the slimiest. The yellowish to brownish or
lavender cap, slimy universal veil, and viscid, more or less equal, violet or lavender stalk are
the principal diagnostic characters of C. cylindripes, but there are numerous look-alikes,
including: C. stillatitius, with only slightly roughened spores; C. salor, with smaller, round
spores; C. pseudosalor, similarly colored in age, but without a purplish cap when young;
C. splendidus, with a small (3-5 cm) liver-brown to tawny cap, also found with conifers;
C. iodes(-C. he lio tropic us) and C. iodioides, with a purple cap thatdoesn’t fade much plus
a club-shaped to spindle-shaped stalk when young (the latter also has bitter-tasting slime);
Cortinarius cylindripes group includes a number of purple or partially purple species with a slimy
cap and stalk. The stalk is equal or slightly spindle-shaped and not conspicuously banded.

C. griseoviolaceus, a small violet-tinged conifer-lover whose cap and stalk are only thinly
viscid (if at all); and C. griseoluridus, with a club-shaped stalk when young and a violet
to olive or ochre cap, found under conifers in the Rocky Mountains. All of these have
viscid, violet-tinged stalks that lack the distinctive belts characteristic of C. collinitus,
and none have caps as dark or as wrinkled as those of C. vanduzerensis and C. elatior.
Other violet-tinged Myxaciums include: C. castaneicolor, found under northern conifers,
with a whitish to tawny, club-shaped stalk, pale violet gills when very young, and a buff
to tawny to chestnut-brown (darker at the center) cap; and C. delibutus and C. sphaero-
sporus (two very similar if not identical species), with a bright yellow to ochre or straw-
colored cap, violet-tinged gills at first, and an equal to club-shaped whitish stalk that
often has yellowish veil patches (or slime) below and often has a violet-tinged apex.

Cortinarius collinitus group Color Plate 104

(Belted Slimy Cortinarius)
CAP 3-10 cm broad, bell-shaped or convex becoming broadly umbonate to plane; surface
smooth, slimy or viscid, color variable: yellowish to orange-yellow, orange-brown, tawny,
ochre, fulvous, or even reddish-brown, the margin often paler or yellower (or in one form
bluish-violet) and sometimes striate. Flesh firm, whitish to yellowish-buff. GILLS close,
adnate to adnexed or notched, pallid or pale grayish (or violet-tinged in one form) when
young, becoming brown and finally rusty-brown in age. STALK 5-15 cm long, 0.5-2 cm
thick, equal or tapering downward, usually rooting somewhat in the humus; viscid or slimy
when moist, color variable but usually whitish above, and lower portion usually breaking
up into irregular whitish, yellowish, ochre, and/or rusty-brown (or in one form violet or
bluish-violet) bands, patches, or scaly rings (but these sometimes obscure). UNIVERSAL
VEIL fibrillose beneath a layer of slime, leaving bands or patches on stalk. CORTINA
pallid, usually forming a ring of hairs near top of stalk which turns rusty-brown from falling
spores. SPORE PRINT rusty-brown; spores 10-15 x 6-8.5 microns, elliptical, roughened.
HABITAT: Solitary, scattered, or in groups under both hardwoods and conifers;
widespread in the northern hemisphere and fairly common in our area in the late fall and
winter under tanoak and madrone at higher elevations in the coastal mountains (often
mingling with C. cotoneus and C. infractus). It is also abundant under aspen in the southern
Rocky Mountains and the Southwest and with conifers in the Pacific Northwest.
EDIBILITY: Better neglected than collected. In spite of the slime it is said to be edible,
but it is a species “complex,” and some of the variants are poorly known or untested.
COMMENTS: This species “complex” is relatively easy to recognize by virtue of the slimy-
viscid cap and belted or scaly, viscid stalk. It is a very confused group taxonomically,
however, and names like C. trivialis and C. mucifluus have been used to describe some of

431
432 CORTINARIACEAE

the variants which differ in color and spore size. Our local version, which might be C.
trivialis, does not show violet on the stem, and neither does the common aspen-loving
form. C. elatior is a similar species that is not so strikingly banded. It has a dark brown to
olive-brown or yellow-brown cap that is often radially wrinkled or grooved at the margin
in age, violet or bluish-tinged gills when young, and a tapered, rooting stalk that is also
bluish- or violet-toned (and slimy). It favors conifers and is fairly common in northern
California and the Pacific Northwest. Other species: C. pallidifolius has pallid gills
when young and a viscid club-shaped stalk with ochre-brown patches or zones; it is also
found under conifers in the Pacific Northwest, especially fir. C. crocolitus (-C. trium-
phans?) and close relatives have a yellowish to brown viscid cap and a dry stalk marked
with yellow-brown to ochre belts or patches (see photo on p. 419); they favor hardwoods.

Cortinarius vanduzerensis
CAP 4-10 cm broad, somewhat conical becoming broadly conical or convex; surface
smooth, very slimy when moist, deep chestnut-brown to nearly black when young, be¬
coming paler chesnut and finally cinnamon-brown in age, often wrinkled radially or
corrugated at maturity, especially toward margin. Flesh pallid to cinnamon-buff; odor
mild. GILLS close, adnate or adnexed, pallid or buff becoming pale brown and finally
dull cinnamon-brown. STALK 8-20 cm long, 1-2 cm thick, equal or often tapered slightly
toward the base, often deeply rooted, viscid to very slimy, bluish-violet to violet to dark
lavender above, paler below, often fading in age. UNIVERSAL VEIL slimy, sheathing at
least the lower half of the stalk and occasionally breaking up into concentric zones.
CORTINA forming a ring of hairs near top of stalk or disappearing. SPORE PRINT
rusty-brown; spores 11-15 * 7-9 microns, elliptical, roughened.
HABITAT: Solitary, scattered, or in groups under conifers; known only from northern
California and the Pacific Northwest, where it is fairly common in the fall and early winter.
I have found it several times under Sitka spruce.
EDIBILITY: Unknown, but much too slippery to be of value.
COMMENTS: The dark slimy cap, slimy violet or bluish-violet stalk, and pallid to brown
gills when young form a distinctive combination of features. It closely resembles C. elatior
(see comments under the C. collinitus group) in shape, size, and the wrinkled cap at
maturity, but does not have the bluish-violet immature gills of that species. The stalk
often roots deeply in the ground, but the presence of a veil distinguishes it from Phaeo-
collybia. The stalk occasionally shows concentric ring-zones, but not to the extent of
C. collinitus. (Violet-stalked forms of the latteralsohaveabrighterorpalercap.)Forother
similar “Myxaciums” with paler caps, see C. cylindripes group. Also see photo on p. 433.

Cortinarius ponderosus (Ponderous Cortinarius)

CAP 10-35 cm broad, convex becoming nearly plane in old age; surface viscid to slimy
when moist, brown to cinnamon or russet at the center, with small, flattened, spotlike,
russet to rusty-brown or brown scales; margin usually yellow or ochre-yellow to yellow-
olive or tawny, remaining inrolled for a long time. Flesh very thick and firm, whitish or
buff; odor mild to somewhat spermatic (Inocybe-like). GILLS close, slightly decurrent
to adnate or sometimes adnexed, lilac-tinged when young(at least near margin), becoming
dingy ochre and finally rusty-brown as spores mature; often with rusty or russet spots or
stains in age. STALK 8-20 cm long, (3) 4-7 cm thick, equal or slightly enlarged below or
tapered at the base; solid, very firm and hard; viscid below when moist; apex pallid or
colored like gills, lower portion with yellow-ochre to rusty-brown or russet stains and/ or
fibrils or fibrillose patches. UNIVERSAL VEIL slimy, yellow, disappearing or leaving
Left: Cortinarius vanduzerensis has a dark slimy cap and long, violet-tinged stalk. Right: Cortinarius
ponderosus (or something very similar—see comments below). This species may be the largest of
all the Cortinarii, but this is a rather small specimen.

slime on stalk. CORTINA often leaving a hairy zone on upper stalk. SPORE PRINT
rusty-brown; spores 8.5-11 * 5-6 microns, elliptical, roughened.

HABITAT: Solitary to scattered or gregarious or in arcs on ground in mixed woods (oak,

ponderosa pine, etc.) in northern California and southern Oregon in late fall and winter;
not common, but often prolific when it fruits. A very similar if not identical species occurs
under oak and madrone (see comments).
EDIBILITY: Better chucked than plucked. It is certainly large enough to be tempting,
but its edibility has not been determined, and several Cortinarii are very poisonous.
COMMENTS: This massive Cortinarius can almost be identified by its size alone. The
stalk is exceptionally hard and thick and the fruiting bodies develop gradually over a con¬
siderable length of time. An equally ponderous but possibly distinct Cortinarius occurs
in our area. It differs in several minor respects: the stalk is dry or only slightly viscid (tacky),
the intermediate color phase of the gills is grayish rather than ochre, the cap is viscid but not
usually slimy (and the slime on the cap, when present, is not yellow), and the upper half of
the stalk is slightly different in color. In size, shape, and overall aspect, however, it is
remarkably similar (see photograph). Whether it is just a variation of C. ponderosus ora
distinct species is uncertain, but collectors should have no trouble recognizing it. For other
large species without a pronounced basal bulb, see C. balteatus.

Cortinarius balteatus
CAP (3) 6-20 (25) cm broad, broadly convex with an inrolled margin becoming plane in
age; surface viscid when moist but soon dry and often shiny, smooth, without scales but
sometimes appearing streaked or blotched; dull violet or lavender to lavender-gray when
young (but sometimes whitish while still covered by humus), soon becoming tawny to
tawny-olive or brown from the center outward, the margin usually remaining lavender
until old age. Flesh thick, firm, whitish to grayish to buff or slightly violet-tinted under the
cuticle; odor mild. GILLS adnate to adnexed or notched, close, whitish or tinged violet
to violet-gray when young, gradually becoming brown as spores mature. STALK 5-12 cm
long, (1) 2-4 (5) cm thick, equal above, often slightly narrowed at base (but sometimes
thicker); firm, solid, dry, pallid or lavender becoming brownish-stained in age, especially
below (or staining yellow in one form). CORTINA disappearing or leaving hairs on stalk
which trap falling spores.-SPORE PRINT dull cinnamon-brown; spores 9-11 * 5-6
microns, elliptical, roughened.

433
Cortinarius balteatus is a large firm purple-tinged species. Note how the stalk lacks a basal bulb

HABITAT: Scattered to densely gregarious on ground under hardwoods and sometimes

conifers, usually with members of the Ericaceae (huckleberry, manzanita, madrone, etc.)
also present; widely distributed. In our area it is sometimes common in the fall and early
winter under madrone. Usually only one crop is produced but the fruiting bodies develop
and decay slowly, and so may be found over a period of several weeks. I have also seen it in
the Oregon Cascades (near Crater Lake) in both the spring and fall.
EDIBILITY: Unknown. As Alexander Smith points out, it is firm and meaty enough to
warrant experimentation—but in light of the recent rash of poisonings caused by species
of Cortinarius, who wants to be the one to try it?
COMMENTS: This cumbersome Cortinarius can be recognized by its violet-tinged
gills when young, violet-tinted cap margin, and hard thick stem which is not abruptly
bulbous. The cap is only thinly viscid and may not seem so at all in dry weather. In wet
weather our form is often full of tiny springtail beetles (columbalids). Other cumbersome,
viscid-capped Cortinarii without an abrupt basal bulb include: C. largus, very similar but
with violet-tinged flesh and an ochre to reddish-brown cap in age; C. balteato-cumatilis,
also quite similar but with an odor “like overripe pears” (Stuntz); and C. crassus, with a
yellow-brown to russet or reddish-brown cap, short thick stalk, and pallid to buff gills
which gradually darken to cinnamon. The latter is fairly common under western conifers,
especially at higher elevations (I have seen it in the Rockies).

Cortinarius subfoetidus
CAP 2.5-10 cm broad, broadly umbonate to convex or plane in age; surface viscid orslimy
when moist, smooth or appearing fibrillose, bright lavender or violet or at times bluish-
lavender, in age sometimes fading at the center to buff or paler. Flesh tinged cap color, paler
in age; odor distinctly fragrant, but with a fetid (nauseating) component. GILLS close,
adnexed or notched to adnate, at first lilac or violet, but becoming pale to dull brown and
finally rusty-brown in age. STALK 5-8 cm long, 0.7-2 cm thick, more or less equal, colored
like cap or paler (often fading), the apex often whitish; not viscid. UNIVERSAL VEIL
fibrillose, forming a pale lavender to violet sheath over lower portion of stalk. CORTINA
transient, sometimes leaving a few hairs at stalk apex which trap falling spores. SPORE
PRINT rusty-brown; spores 7-10 * 5-5.5 microns, elliptical, roughened.

434
CORTINARIUS 435

HABITAT: Solitary to widely scattered or in small groups in duff and moss under conifers;
known only from the Pacific Northwest, fruiting primarily in the fall, especially with
hemlock and/ or fir. I have seen it in northern Idaho, where, according to Alexander Smith,
it is more common than anywhere else.
EDIBILITY: Better wasted than tasted.
COMMENTS: The sickeningly sweet odor (which has been likened to that of overripe
pears) and bright lavender to violet color combine to make this one of the easiest of all
Cortinarii to recognize. Alas, the casual hunter may not get a chance to recognize it
because it is not a very common species. The viscid cap distinguishes it from C. traganus,
C. camphoratus and other “Sericeocybes,” while the stalk is neither viscid as in the C.
cylindripes group nor bulbous as in C. olympianus and other purplish “Bulbopodiums.”
Although uncommon, it attracts more than its share of attention because of its beautiful
color. C. balteato-cumatilis (see comments under C. balteatus) in similar in odor and
color but is much more robust (stalk typically at least 2 cm thick).

Cortinarius infractus (Sooty-Olive Cortinarius)

CAP 4-13 cm broad, obtuse or convex becoming broadly umbonate or plane; surface
viscid when moist, dark olive to sooty-olive becoming slightly browner (dingy yellow-
brown) in age or sometimes mottled with tawny shades; margin sometimes wavy and/or
faintly zoned. Flesh thick, firm, whitish or tinged violet or ochre-buff; taste bitter (at least of
the skin). GILLS notched to adnexed or adnate, close, dark olive to sooty-olive or olive-
brown, or in some forms tinged violet at first, becoming dark cinnamon-brown in old age.
STALK 3-13 cm long, 0.5-2.5 cm thick, equal or more often enlarged slightly below; solid,
fibrillose, dry, whitish or tinged cap color (or in some forms tinged violet at apex), and often
dingy olive-brown toward the base. CORTINA pale grayish, usually forming a hairy
median or superior zone on stalk which turns rusty-brown from falling spores. SPORE
PRINT rusty-brown; spores 7-9 * 5-7 microns, elliptical to nearly round, roughened.
HABIT AT: Scattered to gregarious on ground under both hardwoods and conifers; widely
distributed and common. In our area it fruits in the fall and winter, usually under tanoak
and madrone (often accompanied by C. collinitus, C. cotoneus, and Entoloma species).
EDIBILITY: Better overlooked than cooked—it is probably bitter-tasting.

Cortinarius infractus is a common sooty-olive species with a viscid cap when moist. Note collapsed
hairs (cortina) on stalk. These are fairly mature specimens; young ones have smaller caps.

\ , '♦ "V
436 CORTINARIACEAE

COMMENTS: A distinctive Cortinarius by virtue of the sooty-olive color of its cap and
gills. The cap is viscid when moist, but there is no abrupt bulb at the base of the stalk as in
the C. scaurus group. A number of varieties have been recognized, some of which show
violet in the stalk apex and/ or immature gills. The common version in our area, however, is
violet-less. C. immixtus is a similar species with brighter (yellowish-olive-green) immature
gills, a mild taste, and larger spores; it has been reported from the Pacific Northwest.

Cortinariuspercomis (Fragrant Cortinarius)

CAP 4-8 cm broad, convex to broadly convex with an inrolled margin when young, ex¬
panding in age; surface smooth, viscid when moist, bright to dull yellow to bright ochre
or at times tinged fulvous or greenish-yellow. Flesh yellow; odor strongly aromatic and
sweet (see comments); taste mild to slightly unpleasant. GILLS close, adnexed or notched
to adnate, pale yellow to sulfur-yellow when young, becoming browner (dingy olive-brown
to rusty-cinnamon) as spores mature. STALK 4-8 cm long, 0.7-1.5 (2.5) cm thick at apex,
equal or more often club-shaped (thicker below), but lacking a sharply-defined bulb;
bright yellow to yellowish-white, dry, fibrillose, solid, firm. UNIVERSAL VEIL sulfur-
yellow, fibrillose, often leaving remnants (fibrils) on stalk. CORTINA usually collapsing to
form a ring of hairs on upper stalk which turn rusty-brown from falling spores. SPORE
PRINT rusty-brown; spores 10-13 x 5-6.5 microns, elliptical, roughened.
HABITAT: Solitary, scattered, or in small groups on ground under conifers; widely
distributed. It is fairly frequent in western North America in the late summer and fall, but
seldom in quantity. In our area a similar fragrant species (see comments) occurs with oak.
EDIBILITY: Unknown. I have cautiously sampled our local variety (see comments), and
found its flavor to be a distinct disappointment (it didn’t live up to the odor).
COMMENTS: The strong, remarkably sweet odor (best detected by breaking open the
flesh) and yellow color are the outstanding characteristics of this attractive Cortinarius.
The odor is difficult to describe but is reminiscent—to different people—of citrinella,
toilet bowl cleaner, or papaya. In our area C. luteoarmillatus (?), with a similar odor, is
common in the fall and winter under live oak. It has a pale ochre to pale tan, buff, or pale
yellowish cap, white flesh, and a white stalk with or without yellowish universal veil zones.
C. citrinifolius looks and smells like C. percomis, but has a viscid stalk; it occurs in the
Pacific Northwest. C. odorifer has an aniselike odor and a rimmed basal bulb (at least when
young) and a greenish-yellow to cinnamon to vinaceous-brown cap. C. osmophorus is
a fragrant European species that closely mimics our local variety in color, but it also has
a rimmed bulb. None of these are worth eating.

Cortinarius luteoarmillatus (or a very similar species—see above comments) is common in California
under oak. Note how stalk is club-shaped (thicker below) but not bulbous. Sweet odor is distinctive.
CORTINARIUS 437

Cortinarius mutabilis group (Purple-Staining Cortinarius)

CAP 4-10 cm broad, convex becoming plane or undulating; surface smooth, viscid when
moist, at first violet or grayish-violet, but usually suffused in age with buff, ochre, or brown,
especially toward the center (margin often remaining violet). Flesh thick, pallid or pale
violet becoming dark purple when bruised or cut and then rubbed (sometimes staining
slowly); odor mild. GILLS adnate or more often adnexed or notched, close, dull violet
or grayish-lilac when young, slowly becoming pale brownish as spores mature; turning
deep lilac or purple when bruised. STALK 4-10 cm ong, 1-2 cm thick at apex, thicker or
swollen at base but without an abrupt bulb; pallid to pale violet or colored like cap, staining
deep lilac or purple when bruised. STALK 4-10 cm long, 1-2 cm thick at apex, thicker or
swollen at base but without an abrupt bulb; pallid to pale violet or colored like cap, staining
which traps falling spores. SPORE PRINT rusty-brown; spores 7-9 * 4.5-5.5 microns,
elliptical, roughened.
HABITAT: Scattered to gregarious or in small clumps on ground or very rotten wood
under conifers, western North America; especially common at higher elevations in the late
summer and fall. I have seen it in the Rocky M ountains and Cascades, but not on the coast.
EDIBILITY: Unknown, like most of us.
COMMENTS: This species “complex” is variable in a number of respects, but can be
recognized by the purple-staining gills and stalk surface. The flesh also stains purple, but
the reaction seems to vary somewhat in brightness and duration. Other species: C. pur-
purascens is a purple-staining species with a purple-brown to reddish-brown, often
variegated cap and more sharply defined basal bulb on the stem; C. occidentalis stains
purple, but has a bluish cap that becomes grayish in age; C. subpur pur ascens has purple-
staining gills, but the flesh does not stain. All of these grow mainly with conifers.

Cortinarius glaucopus group

CAP 4-12(17) cm broad, convex becoming plane or with margin slightly uplifted; surface
smooth, viscid or tacky when moist, color extremely variable: rich greenish-brown to
olive-black, greenish-gray, steel-gray, bluish-gray, etc., often streaked or mottled with
yellowish or ochre fibrils and often becoming fulvous, cinnamon, or rusty-colored in age
(typically from the center outward); margin sometimes yellowish-olive and often wavy.
Flesh firm, thick, pallid to grayish or tinged violet-blue, becoming ochraceous in age; also
ochraceous in base of stalk. GILLS close, adnate to adnexed or notched, bluish-violet to
bluish-gray or sometimes gray when young, darkening to rusty-brown as spores mature.
STALK 4-10 cm long, 1-3 cm thick at apex, more or less equal above, but with a bulb at
base, the bulb abrupt and rimmed when young but often oblique or poorly defined in age;
color variable but usually violet, blue, or green at apex and variously colored (often paler)
below, often browner in age; dry, solid, firm. CORTINA pallid or pale bluish-violet,
usually leaving hairs on stalk which are stained rusty-brown by falling spores. SPORE
PRINT rusty-brown; spores 6-10 x 4-5.5 microns, elliptical, slightly roughened.
HABITAT: Scattered to densely gregarious or in small clumps or sometimes in troops,
associated with both hardwoods and conifers; widely distributed but particularly common
in western North America. Our members of this “complex” fruit prolifically in the fall and
winter and are the most common “Bulbopodiums” of our live oak woodlands. In the Rocky
Mountains and Pacific Northwest, however, it favors conifers, especially spruce.
EDIBILITY: Better drowned and beaten than browned and eaten. Many members of
this group have not been adequately tested and some are apparently bitter.
COMMENTS: This species “complex” is extremely variable in color, but can be told in the
button stage by its conspicuous bulb and blue-gray to bluish-violet gills. In age it becomes
Cortinarius glaucopus group is common under both hardwoods and conifers. In the button stage
(not shown here) the basal bulb is usually quite large and abrupt, but becomes less pronounced as
the stalk elongates.

cinnamon- or rusty-colored and is easily confused with dozens of other Cortinarii.

Younger specimens can resemble the blewit (Clitocybe nuda), but have a cortina and rusty-
brown spores. C. pseudoarquatus is a similar species with a more buff or brownish cap
and significantly larger spores. For other bluish-violet “Bulbopodiums” (most of which
are more violet than blue and/ or have differently colored caps), see comments under the
C. sodagnitus group and C. olympianus.

Cortinarius sodagnitus group

CAP 3-7 (10) cm broad, convex to plane; surface smooth, viscid when moist, at first bright
violet or lilac overall, but soon becoming brown to ochraceous or buff from the center
outward (or becoming somewhat silvery in dry weather), the margin retaining violet tones
well into maturity. Flesh thick, firm, whitish with a violet tinge at top of stalk and ochre tint
in basal bulb; not staining appreciably; taste bitter (at least of the skin). GILLS violet when
young, becoming duller and suffused with brown or cinnamon-brown, but the edges
usually remaining violet until old age; close, adnate to adnexed or notched. STALK 2.5-8
cm long, 0.5-2 cm thick, with a small but sharply defined, rimmed bulb at the base (1-3 cm
broad); violet like the cap or pallid when young, discoloring brownish or rusty from the
base upward, but usually retaining distinct violet zone at apex even in old age; solid, not
viscid. CORTINA violet-tinged, leaving a median to superior zone of hairs on stalk, the
hairs often rusty-stained by spores. SPORE PRINT rusty-brown; spores 10-12 * 5.5-6.5
microns, elliptical, roughened. Cap surface staining red or wine-red in KOH or NaOH.
HABITAT: Solitary, scattered, or gregarious on ground under hardwoods; distribution
uncertain. This species “complex” is quite common in our live oak woodlands in the fall
and winter. It was originally described from Europe, where it favors beech.

EDIBILITY: Better tossed than sauced—it is probably bitter.

COMMENTS: This beautiful member of the subgenus Bulbopodium can be told by its
rather small but well-defined, abrupt, rimmed bulb at the base of the stem (see photo at
top of p. 439), by its violet colors when young, and by the tendency of the cap margin and
stem apex to remain violet even in old age. Fresh specimens mimic the blewit (Clitocybe
nuda) in size, shape, and color, but have a cortina and rusty-brown spores. Closely related
species include: C. dibaphus, with a slightly pinker or more lilac cap and gills which turn
brown more quickly (but otherwise very similar, if not the same); and C. caesiostramineus,
with a slightly grayer cap. There are dozens of other purple or violet “Bulbopodiums”—
see comments under C. olympianus for some of them.

438
Cortinarius sodagnitus group. Note small but very prominent basal bulb. Common in California
under oak.

Cortinarius olympianus
CAP 3-7 (10) cm broad, convex becoming plane, the margin inrolled at first; surface
smooth, slimy or viscid when moist, pale violet or lilac, sometimes fading to lilac-white,
or sometimes tinged yellowish at center. Flesh fairly thick, white to grayish. GILLS pale
lilac to pinkish-lilac when young, becoming browner in age but usually retaining a lilac
tinge for a long time; notched or adnexed to adnate, close, not staining when bruised.
STALK 3-7 cm long, 0.7-1.2 cm thick at apex, solid, firm, not viscid, with a distinct basal
bulb that is rimmed, at least when young; lilac or pale violet above, often brownish-stained
at base. CORTINA scanty, lilac-white, often leaving a few hairs on stalk which trap falling
spores. SPORE PRINT rusty-brown; spores 8-10 x 5-6 microns, elliptical, slightly
roughened.
HABITAT: Scattered to gregarious on ground under conifers, fairly common in the
Pacific Northwest in the late summer and fall, especially where there is fir, spruce, and/ or
hemlock. I have seen large fruitings in Mt. Rainier National Park in late September.
EDIBILITY: Unknown.
COMMENTS: This is one of several viscid-capped Cortinarii with a rimmed bulb and
overall lilac to violet-white color. There are several similarly colored species that favor
hardwoods and are especially prominent in eastern North America. These include:
C. michiganensis and C. caesiocyaneus, both larger with conspicuous bulbs and pale violet
to pale bluish-violet overall color (when fresh), the latter with a whitish universal veil;
C. velicopia, similar to the previous two species but discoloring ochre to buff or tan on the
cap and basal bulb with age (often rather quickly); C. aggregatus, with darker violet gills
when young and only a small, oblique bulb at the stem base; C. caerulescens, with a darker
blue or violet-blue cap when young; and C. cyanites (see couplet #72 of the key to
Cortinarius). There are also several species that are very similar in size, shape, and color
but do not have viscid caps, e.g., C. argentatus and C. subargentatus (see comments under
C. alboviolaceus and photo on p. 448).

Cortinarius cedretorum Color Plate 105

CAP 5-15 cm broad, convex with an inrolled margin when young, becoming broadly con¬
vex or plane; surface smooth, viscid or slimy when moist, at first yellow but darkening
slowly to cinnamon-brown, orange-brown, or reddish-brown from the center outward,
the margin often remaining yellowish. Flesh thick, firm, pallid to pale yellow with distinct
lavender or violet areas near cap cuticle and in stalk; often ochraceous-yellow in base of

439
440 CORTINARIACEAE

stalk; odor mild. GILLS close, adnate to adnexed, yellow or sometimes greenish-yellow
when young, dull cinnamon-brown in age, but often with an intermediate lavender to dull
purple phase. STALK 4-12 cm long, 1.5-3 cm thick at apex, with a conspicuous, abrupt
basal bulb which is rimmed, at least when young and is 3-7 cm broad; solid, not viscid,
color variable: pale lavender or yellow at apex or throughout, becoming dingy or often
rusty-stained in age, the bulb usually yellow. CORTINA pale yellow to greenish-yellow,
usually leaving hairs on the stalk which trap falling spores. SPORE PRINT rusty-brown;
spores 11-14 * 6.5-8 microns, elliptical, roughened.
HABITAT: Solitary to scattered or gregarious on ground in woods, mainly in western
North America. It is sometimes common in our area under oak in the late fall and winter; in
the Pacific Northwest it favors conifers and in the Southwest I’ve seen it in mixed woods of
aspen, spruce, and fir.
EDIBILITY: Unknown.
COMMENTS: The yellow gills and viscid yellow cap when young plus the beautiful violet-
tinged flesh (see color plate!) and bulbous stalk form a distinctive set of characteristics.
In our local form the purplish phase of the gills is often transient and difficult to detect.
C. aureofulvus is a similar oak-loving species which may show violet in the flesh but not
in the gills. C. atkinsonianus shows the same transient violet gill phase as C. cedretorum,
but has an olive-yellow to yellowish to deep reddish-brown cap and grows with hard¬
woods in eastern North America. C. montanus has violet-tinged gills with olive or yel¬
lowish-olive edges when young, and a rusty-brown to olive-brown, yellow-brown, or varie¬
gated/ streaked cap. It is often common under old-growth conifers in western mountains,
particularly the Cascades.

Cortinarius scaurus group

CAP 3-10 cm broad, convex to broadly convex with the margin inrolled somewhatatfirst,
expanding in age; surface smooth, viscid when moist, color variable: usually olive-green
to dark olive-brown, but in some varieties brownish at the center and greenish to greenish-
yellow at the margin, and in others bright grass- or citrine-green, often with a browner
center or brownish spots and fibrils. Flesh pallid to yellowish, thick, firm; odor and taste
usually mild. GILLS close, notched or adnexed to adnate, typically olive to greenish-
yellow (in some forms with a fleeting initial lavender or bluish-violet stage), but in some
forms yellow when young, all forms becoming browner or rustier as spores mature.
STALK 4-10 cm long, 0.5-1.5 (2) cm thick at apex, equal above, the base with a more or less
rimmed, abrupt bulb when young that may become oblique or obscure in age; usually
greenish like the cap or yellower or even pallid (in age), the apex sometimes bluish, the
bulb often sulfur-yellow; solid, not viscid. CORTINA olive to yellow, usually leaving
hairs on stalk which turn rusty-brown from falling spores. SPORE PRINT rusty-brown;
spores 9-13 * 6-7.5 microns, elliptical, roughened.
HABITAT: Solitary to scattered or gregarious on ground in woods; widely distributed.
Some members of this group (see comments) favor conifers and are fairly common in the
Pacific Northwest; others grow under oak and other hardwoods, including one uniden¬
tified variety that is common in our area in the fall and winter with live oak(see comments).
EDIBILITY: Due to difficulty in identification, this entire group should be avoided. The
group as a whole is probably edible, but I wouldn’t bet my life on it—and that’s just what
you do when testing a Cortinarius!
COMMENTS: The above description has been broadened to include a number of bright
green to greenish-yellow species with a viscid cap and well-defined bulb (at least when
CORTINARIUS 441

young). Mycologists prefer to refer to them collectively as the C. scaurus group (when
they refer to them at all, which is seldom), for they are a confusing lot, both in the field
and in the literature. They are readily distinguished as a group, however, by their color.
The “true” C. scaurus of Europe is said to have a spotted cap, slender stalk (less than 1 cm
thick), and grow with conifers. Some of the other species in the group include: C. herpe-
ticus, very similar but with a thicker stalk and often with violet-tinged immature gills and
flesh and a whitish basal bulb; C. prasinus, with an olive-colored cap, olive to olive-yellow
gills, and yellowish or olive-tinged stalk, fairly common in the Pacific Northwest under
conifers; C. virentophyllus, with a deep green cap that fades to yellow as it ages, green gills,
and a pale bluish stem, found under hardwoods in eastern North America; C.flavovirens,
with a farinaceous odor, greenish-yellow gills, and yellow stalk, reported from southern
California; C. orichalceus and C. rufo-olivaceus, with a reddish to reddish-brown cap (at
least at center), and greenish to yellowish stalk (the latter species is quite large and robust,
the former is smaller and can have an oranger cap); and finally, a distinctive but
unidentified species which is sometimes common under live oak in California. It has a
bright citrine-green cap (sometimes shaded with brown at the center), yellow gills when
young, and a yellowish stalk with a bulb at the base. Whether it is one of several similar
European species, or is unnamed and endemic to California is unclear, but it is definitely a
member of the C. scaurus group, which should satisfy all but the most hardcore Cortinarius
-categorizers. See also C. montanus (under C. cedretorum).

Cortinarius fulmineus
CAP 5-15 cm broad, convex becoming plane, the margin at first incurved; surface viscid
when moist, entirely yellow or with only the margin yellow and the center fulvous to ochre
to orange-brown, sometimes with small spotlike scales. Flesh thick, firm, yellow to
yellowish-white or buff; odor and taste mild or radishlike. GILLS close, adnate or
notched, yellow to yellow-brown or ochre, becoming ochre-cinnamon and finally rusty-
brown as spores mature. STALK 3-7 cm long, 1-3 cm thick at apex, often rather short and
stout, with a large rimmed bulb at the base; solid, firm, not viscid; yellowish-white or
yellow, sometimes becoming ochraceous in age. CORTINA pallid or yellowish, often
leaving hairs on stalk which turn rusty-brown from falling spores. SPORE PRINT rusty-
brown; spores 8-10 * 4.5-6 microns, elliptical, roughened.
HABIT AT: Solitary to widely scattered or gregarious on ground under hardwoods; widely
distributed. It is not uncommon in our area in the fall and winter in mixed woods and
under live oak.

EDIBILITY: Unknown.
COMMENTS: This species typifies a number of yellowish to rusty-orange Cortinarii with
a viscid cap, yellowish to orange gills, and a distinctly rimmed bulb when young. The
rimmed bulb places it in the colorful subgenus Bulbopodium, and the yellowish or pallid
flesh separates it from another “Bulbopodium,” C. cedretorum. Other species: C.fulgens
is similar but brighter orange in color; C. elegantior is also similar but has larger spores
(12-16 microns long), while C. elegantioides has even larger spores and a decidedly bitter
taste. There are also several beautiful “Bulbopodiums” with a bright yellow cap (or yellow-
margined with a redder, browner, or oranger center) and lilac to violet gills at first. These
include: C. metarius, common under spruce and fir in western North America, with a
fairly broad basal bulb; C. calochrous, widespread under both hardwoods and conifers
and very similar to C. metarius, but with a smaller bulb and slightly shorter spores; C.
citrinipedes, cap yellowish with a darker brown margin; and C. cyanopus (-C. amoene-
lens), widespread, with an ochre cap, bitter-tasting cap cuticle, and deep violet gills when
young.
442 CORTINARIACEAE

Cortinarius multiformis
CAP 4-12 cm broad, convex becoming plane or broadly umbonate, the margin at first
inrolled; surface smooth, viscid when moist, ochre to ochre-buff to tawny oryellow-brown
or occasionally more reddish or fulvous and sometimes becoming rustier in age, often with
a whitish silky bloom when young. Flesh thick, firm, pallid; odor mild or slightly pungent.
GILLS adnate to adnexed or notched, close, at first whitish, then tan or watery brown,
finally rusty-cinnamon. STALK 4-10 cm long, 1-2 (2.5) cm thick at apex, usually with a
bulb at the base that is abrupt when young but often obscure or even absent in age; dry,
solid, firm, white or discoloring tan or ochre. UNIVERSAL VEIL disappearing or leaving
a thin whitish film on cap. CORTINA scanty, white, disappearing or leaving a few hairs
on stalk which trap falling spores. SPORE PRINT cinnamon-brown; spores 7-11 x 4-6
microns, elliptical, minutely roughened.
HABITAT: Solitary to scattered or gregarious on ground in woods (mainly under coni¬
fers); widely distributed. It is common throughout much of the West, but infrequent or
absent in our area. I have seen large fruitings under spruce near Santa Fe, New Mexico.
EDIBILITY: Unknown. It is eaten in Europe, but I can find no information on the North
American version, and there are too many similar species for me to recommend it.
COMMENTS: The viscid, ochre to tan cap and pallid to tan immature gills serve to dis¬
tinguish this species from most other Cortinarii. The cap and gill color suggest a Hebeloma,
but the spores are cinnamon-brown and the odor is not radishlike. The hoary white film on
young caps is reminiscent of the gypsy mushroom (Rozites caperata), but the veil is not
membranous. There are many similar Cortinarii with non-viscid caps that sometimes
mingle with C. multiformis (see C. laniger). Other viscid-capped species with an equal to
club-shaped (but not abruptly bulbous) stalk include: C. turmalis, with a tawny to fulvous
or yellow-brown to darker brown cap and a well-developed white veil that usually leaves
a distinct fibrillose sheath and/or ring on the stalk; C. cliduchus and C. lalus, without
such a copious veil but otherwise similar to C. turmalis (the latter with a paler cap); C.
crassus, a stout, thick-stemmed species (see comments under C. balteatus); C. varius,
a widespread species with a fulvous to bright yellow-brown or ochre cap and violet gills
when young plus a white stalk; C. variicolor, with a purplish to brownish cap, lilac gills
when young, and a fibrillose, whitish or lilac-tinged stalk; and C. claricolor, with bluish-
gray to brownish gills, a yellower cap, and more copious universal veil. All but the last
species favor conifers and occur fairly commonly in the Pacific Northwest and/ or Rocky
Mountains.

Cortinarius magnivelatus
CAP 3-10 cm broad, convex becoming plane or irregularly undulating; surface moist or
dry but not viscid, smooth or fibrillose, entirely white or white at margin and ochre toward
the center, often stained buff or ochraceous in age or in bruised areas; margin incurved.
Flesh thick, firm, white to buff; odor mild. GILLS close, slightly decurrent to adnate or
adnexed, whitish when very young, becoming buff and then tawny-brown to brown; often
forked near the stalk. STALK 4-6 cm long, 1-3 cm thick at apex, often (but not always)
enlarged below, colored like cap or whiter, solid. VEIL thick, tough, membranous, elastic,
whitish, typically covering the gills through maturity or shredding radially but not
normally detaching from the cap and stalk. SPORE PRINT rusty-brown; spores 9-14
x 6-8 microns, elliptical, roughened.
HABITAT: Solitary to gregarious under mountain conifers (especially fir and pine),
usually buried in the duff; fairly common throughout the higher mountains of California,
particularly in the late spring and summer, but probably more widespread.
EDIBILITY: Unknown.
Two conifer-loving Cortinarii with persistent veils. Cortinarius magnivelatus (two at right) has a
tough, whitish veil that is reminiscent of a rubber band. Cortinarius verrucisporus (three at left)
is yellower and has a short stalk and thinner veil that tends to tear radially (see comments below).

COMMENTS: The whitish color, underground growth habit, and tough, thick, persistent
veil distinguish this Cortinarius from most others. Presumably the latter features are adap¬
tations to the pendulum-like changes of weather that occur in our western mountains.
Many Cortinarii (particularly “Bulbopodiums”) tend to fruit under the duff in dry
weather, but they do not have the persistent membranous veil of this species. That the veil
does not break suggests that this species is on its way to becoming “gastroid.” The spores,
however, are forcibly discharged (hence a spore print is obtainable), unlike the truly
gastroid genus Thaxterogaster (see p. 734). The membranous nature of the veil can
lead to confusion with other brown-spored genera, but the overall aspect is clearly “cor-
tinarioid.” Several other semi-gastroid Cortinarii occur in western North America. Like
C. magnivelatus, they have a persistent veil and tend to grow under the duff, although
the mature caps may surface if conditions are wet enough. These species include: C.
wiebeae, discovered in the Cascades, similarly colored but larger with thin, fragile gills
that are brown or rusty (not whitish) when young; C. verrucisporus, fairly common in
the Sierra Nevada, with a yellow to yellow-brown or rusty-stained cap and a thinner
yellowish veil that stretches from the stalk or stalk base to the cap margin and usually tears
radially (see above photo), plus broad gills, a frequently underdeveloped stalk, and very
coarsely warted spores; C. bigelowii, fairly common in Idaho, with a pale yellow-brown
to yellowish cap, whitish veil, and a short, fat bulbous stalk; and C. velatus of the Sierra
Nevada, easily distinguished by its purplish to lavender-tinged cap and somewhat thinner,
semi-cobwebby (but persistent) veil. Other semi-gastroid species undoubtedly occur in the
West but have yet to be described.

Cortinarius regalis
CAP (4) 6-15 cm broad, convex with an inrolled margin, becoming broadly umbonate to
plane or with margin slightly uplifted in age; surface dry or very slightly tacky, usually with
a large patch or patches of white fibrillose universal veil tissue at first; background
brownish to dingy flesh-colored to dull brown with a vinaceous tinge, often appearing
somewhat streaked. Flesh thick, pallid or tinged vinaceous or cap color; odor mild to some¬
what musty or sometimes fruity. GILLS usually adnexed or notched, pallid in button
stage, becoming grayish-brown to dull watery-brown and eventually dark brown. STALK
6-16 cm long, (1) 2-3.5 cm thick at apex, with a large, abrupt and/ or rimmed bulb at base
(3-6 cm thick); dry, solid, firm pallid or tinged cap color, fibrillose. UNIVERSAL VEIL

443
444 CORTINARIACEAE

whitish, felty-fibrillose, often leaving remnants on cap and fibrils or hairs on stalk which
become rusty-stained. CORTINA white, disappearing or also leaving hairs on stalk.
SPORE PRINT dull rusty-brown; spores 7-11 * 5-6 microns, elliptical, finely roughened.
HABITAT: Solitary, scattered, or in groups on ground in woods; known only from Cali¬
fornia, where it usually grows with oak. I find it regularly in the fall and winter.
EDIBILITY: Unknown.

COMMENTS: The fairly large size, thick bulbous stalk, absence of distinct violet hues,
and whitish veil material on the cap combine to distinguish this Cortinarius. The bulb is
typical of the subgenus Bulbopodium, but the cap is not truly viscid, making this species
something of an anomaly. Viscid-capped species with conspicuous whitish veil remnants
on the cap and a prominent bulb at the base of the stalk include: C. calyptratus, medium¬
sized, with violet gills when young, occurring under conifers in northern California; and
C. calyptrodermus, larger, with deep violet gills when young, occurring under hardwoods
in eastern North America. Another viscid-capped species with purplish (or purplish-blue)
gills when young, C. volvatus, has a universal veil which forms a whitish sheath or “volva”
on the basal bulb, but does not normally leave conspicuous remnants on the cap.

Cortinarius gentilis (Deadly Cortinarius)

CAP 1 -5 cm broad, conical or bell-shaped at first, expanding somewhat in age but usually
retaining an umbo; surface smooth, not viscid but somewhat hygrophanous, tawny to
ochre to orange-brown or rusty-yellow, fading in age or as it dries. Flesh thin, yellowish.
GILLS fairly well-spaced, adnexed toadnate, ochre-yellow to cinnamon-brown becoming
browner in age. STALK 3-10 cm long, 3-5 (7) mm thick, more or less equal, colored like
the cap or more cinnamon-colored; often (but not always) with traces of the yellow veil
when young; not viscid. UNIVERSAL VEIL yellow, disappearing or leavinga few patches
or hairs on stalk. CORTINA also yellow, disappearing or leaving hairs on stalk. SPORE
PRINT rusty-brown; spores 7-9 * 5.5-7 microns, elliptical, minutely roughened.
HABITAT: Scattered to gregarious orin troops in moss or duff under conifers; widely dis¬
tributed. It is sometimes abundant in the summer and fall in the Rocky Mountains and
Pacific Northwest, but is not likely to be collected by the average mushroom hunter.
EDIBILITY: DEADLY POISONOUS! Fortunately, it is seldom eaten because of its
small size. Along with several other Cortinarii in the subgenus Leprocybe (notably C.
orellanus and C. speciosissimus), it contains toxins which destroy the liver. Symptoms
are greatly delayed (it may be two weeks before they appear!), making the culprit difficult
to pinpoint.
COMMENTS: This species would be just another “LBM” were it not for the fact that it
is deadly poisonous. The yellow-brown to dull orange-brown overall color, somewhat
hygrophanous cap, small size, yellow veil, and rusty-brown spores form a distinctly
undistinguished set of distinguishing features. It is most likely to be confused with the
C. cinnamomeus group, which is more brightly colored, does not have a hygrophanous
cap, and contains different pigments (anthraquinones). Related species thought to be
deadly poisonous include C. rainierensis, C. orellanus, and C. speciosissimus. The latter
two species have caused a rash of deaths in Europe. They are somewhat larger (cap 3-9 cm
broad and tawny-brown to orange-brown, fulvous, or cinnamon-colored) than C. gen¬
tilis; C. speciosissimus has an umbonate cap, C. orellanus has an un-umbonate cap
and a yellowish stem. They may well occur in North America, but their presence has not
yet been definitely established. C. rainierensis, on the other hand, was originally described
from the Pacific Northwest (under conifers). It is rusty-orange to tawny or brownish-
orange, and its cap is covered with small, more or less erect or protruding scales.
CORTINARIUS 445

Cortinarius cotoneus group (Scaly Cortinarius)

CAP 3-10 cm broad, obtuse to convex becoming plane; surface dry, covered with small,
dark, fibrillose scales and appearing blackish to olive-brown at the center where the scales
are densest, and greenish to olive-brown, olive-yellow, or yellow-brown toward the margin
(sometimes paler overall in age). Flesh brownish to pale olive-yellow; odor often somewhat
radishlike. GILLS adnate to adnexed or notched, rusty-yellow to olive-yellow to dull
yellowish or yellow-brown, becoming dull orange to dark cinnamon-brown in age.
ST ALK 4-15 cm long, 0.8-3 cm thick, equal or thicker at base; dry, solid, pale olive-yellow
or yellowish above, tawny to olive or colored like cap below, often dingier or browner
throughout in age. UNIVERSAL VEIL yellow or olive-yellow, often leaving hairy fibrils
or patches on lower stalk. CORTINA disappearing or leaving a zone of hairs on upper
stalk which trap falling spores. SPORE PRINT rusty-brown; spores 7-9 * 5-7.5 microns,
elliptical to nearly round, roughened.
HABITAT: Solitary to scattered or gregarious on ground in woods, widely distributed.
Our representative of this group is fairly common in tanoak-madrone woods in the late fall
and winter, often mingling with C. collinitus and C. infractus. In the Pacific Northwest,
however, it is common under conifers.
EDIBILITY: U nknown, but possibly very dangerous! It belongs to the same group (sub¬
genus Leprocybe) as the deadly poisonous C. gentilis, C. orellanus, and C. speciosissimus.
COMMENTS: The dry cap with small blackish scales on an olive-yellow to yellow-brown
background distinguishes this Cortinarius “complex” from most others. It is one of several
distinctive Cortinarii with a hairy and/ or scaly cap. Others include: C. clandestinus, prac¬
tically identical but with pallid gills when very young; the deep violet C. violaceus( see
description); and a striking trio of eastern and northern hardwood-lovers: C. bolaris,
C. pholideus, and C. squamulosus (see the latter species for more details).

Cortinarius squamulosus Color Plate 108

(Bulbous Scaly Cortinarius)
CAP 4-10 cm broad, convex to broadly umbonate or plane; surface dry, fibrillose, soon
breaking up into dense fibrillose scales, brown tinged with purple at first, usually more
or less chocolate-brown in age, the background paler (yellower or more cinnamon).
Flesh rather thick, whitish to grayish or pinkish-tinged; odor often somewhat spicy, espe¬
cially in age. GILLS sometimes adnate at first but usually deeply notched by maturity,
close, dark purplish or purplish-brown soon becoming brown to cinnamon-brown or
chocolate-brown. STALK 6-15 cm long, 1-2 cm thick at apex, with a conspicuous basal
bulb 3-6 cm broad; smooth to fibrillose or slightly scaly, at first purplish-tinged, becoming
more or less cap-colored, often with a median bandlike ring (from universal veil?). COR¬
TINA whitish to brownish, disappearing or leaving a few hairs on upper stalk. SPORE
PRINT dark rusty-brown; spores6-8 * 5-7 microns, elliptical to nearly round, roughened.
HABITAT: Solitary to gregarious under hardwoods (especially oak), often in low wet
woods; not uncommon in the late summer and fall in northeasern North America. I have
seen impressive fruitings in Minnesota and Wisconsin, but have yet to find it on the west
coast. One source reports it from California but does not mention the habitat.
EDIBILITY: “Consistency very pleasant and flavor fairly good,” says Mcllvaine, who ate
almost anything. However, several relatives are either poisonous or haven’t been tested.
COMMENTS: Like C. violaceus, this species is one of the few truly distinctive Cor¬
tinarii (i.e., one that beginners can recognize in the field), but like that species it is not
particularly common. The dry, purplish-brown to brown, densely scaly cap plus the
446 CORTINARIACEAE

radically bulbous stem are its main features. Another scaly-capped species, C. pholideus,
is common under birch in the Northeast, often on or near rotten logs, and also occurs
rarely in the Pacific Northwest. Its stalk is equal or only slightly thicker at the base and
is decorated with distinctive brown to cinnamon-brown scales like those on the cap (see
photo on p. 424) and the fruiting body is slightly more cinnamon-colored overall than
C. squamulosus. It might be mistaken for a Pholiota because of the brown scales on the
stem and tendency to grow on or near rotting logs, but the dry scaly cap, violet-tinged
stalk apex (when young), and roughened, cinnamon-brown spores distinguish it. Still
another distinctive hardwood-loving easterner, C. bolaris, is quite different: its cap and
stalk feature reddish scales, fibrils, and/ or streaks on a whitish to yellowish background
(see photo on p. 425 and couplet #89 of the key to Cortinarius).

Cortinarius violaceus (Violet Cortinarius) Color Plate 109

CAP 3.5-12 (15) cm broad, convex becoming broadly convex, broadly umbonate, or
plane; surface dry, densely covered with minute erect, tufted hairs or small scales, giving it
a rough, somewhat velvety appearance; deep violet to nearly black, often with a metallic
luster in age; margin often somewhat paler and fringed or ragged. Flesh thick, deep violet
becoming grayish-violet; odor mild or cedarlike. GILLS adnate becoming adnexed or
notched, fairly well-spaced, deep violet or colored like cap, then dusted with cinnamon-
brown spores. STALK 6-18 cm long, 1-2.5 cm thick at apex, equal or more often thicker
below, dry, fibrillose or woolly, deep violet, solid, firm. CORTINA violet, soon disap¬
pearing or leaving a few indistinct hairs near top of stalk which may catch falling spores.
SPORE PRINT rusty-brown; spores 13-17 * 7-10 microns, broadly elliptical to oblong,
roughened. Cystidia present on both the faces and edges of the gills.
HABITAT: Solitary or in twos and threes under conifers, sometimes next to rotting logs;
widely distributed but quite rare except in certain localities, such as the old-growth
coniferous forests of Mt. Rainier and Olympic national parks in Washington. In our area
it shows up occasionally in the late fall and winter in mixed woods; in northern California
it is sometimes quite common under Sitka spruce; in Europe it is said to favor hardwoods.
EDIBILITY: Edible, but not choice. Its principal appeal is its beauty.
COMMENTS: There are dozens upon dozens of violet Cortinarii, but none are as deeply
colored as this one. Its hue alone makes it as unique as it is unforgettable—the color
photograph hardly does it justice. The cap is dry and rough due to the presence of many
small scales or tufted fibrils, another distinctive feature. The color is at times so deep that it
borders on black, making it difficult to pick out in the shade of the forest. Its only rivals
for color are some of the deep blue or violet Leptonias (see L. carnea and L. nigroviolacea),
but they lack a cortina and have pinkish spores. Several colorful Cortinarii make wonder¬
ful dyes (e.g., C. sanguineus, C. semisanguineus), but C. violaceus, despite its intense
color, is not one of them.
It is ironic to note that C. violaceus, the species chosen to typify the immense genus
Cortinarius, is perhaps the least typical of the 1000+ species! Besides its unique color it is
practically the only species to possess sterile cells (cystidia) on both the edges and faces
of the gills. This is the kind of difference to which the “splitters” (see p. 10) owe a living, and
C. violaceus might indeed have its own little genus by now, were it not that under the
existing rules of the International Code of Botanical Nomenclature, the “new” genus
would have to be Cortinarius (since C. violaceus is the type species or “model” of Cor¬
tinarius), and all of the other 1000+ Cortinarii would have to be placed in a new genus
(or genera), thus compounding the confusion that already exists. Noteventhemostardent
“splitter” wants to be responsible for a mess like that! Perhaps as compensation, C. vio¬
laceus is itself split by some “splitters” into two species: C. violaceus, with oblong-elliptical
spores, and C. hercynicus, with rounder spores.
CORTINARIUS 447

Cortinarius traganus (Lilac Conifer Cortinarius) Color Plate 106

CAP 4-13 cm broad, obtuse or convex becoming plane or broadly umbonate; surface
smooth, dry, finely silky or fibrillose, violet to lilac, sometimes with white wedge-shaped
sectors, occasionally also with rusty or ochraceous stains; margin often hung with veil
remnants. Flesh rusty-brown to tawny- or yellow-brown (often marbled) in the stalk,
usually paler in the cap and yellower in the stalk base; odor often faintly pungent or sweet.
GILLS pale cinnamon or ochre-buff becoming cinnamon- or rusty-brown in age; fairly
well-spaced, adnexed to adnate. STALK 5-12 cm long, 1-3 (5) cm thick, usually enlarged
below, solid, dry, finely fibrillose, lilac or purplish, sometimes also with white areas.
CORTINA pale lilac, leaving hairs on upper stalk which turn rusty-brown from falling
spores. SPORE PRINT rusty-brown; spores 7-10 x 5-6 microns, elliptical, roughened.
HABITAT: Solitary, scattered, or gregarious (sometimes in clumps) under conifers;
widely distributed. It is a common late summer and fall mushroom in the mossy, old-
growth forests of the Pacific Northwest and northern California, but does not seem to
extend south into our area. I have often found it growing with the gypsy mushroom,
Rozites caperata.
EDIBILITY: Not firmly established (like so many of us!), but poisonous according to one
source and merely “indigestible” according to another.
COMMENTS: This characteristic inhabitant of northern coniferous forests is easily
recognized by its beautiful lilac or lavender color, rusty-brown gills and spores, and rusty
to tawny-brown flesh in the stalk. The latter feature distinguishes it from most other
members of the subgenus Sericeocybe of Cortinarius (see C. alboviolaceus), and the
sweetish odor (“like overripe pears”) that is frequently present is also distinctive. The cap
is not viscid as in C. olympianus, C. sodagnitus, and others, and the presence of a cortina
separates it from the edible blewit (Clitocybe nuda). Some authorities consider the Ameri¬
can variant to be a distinct species, C. pyriodorus, while reserving the name C. traganus
for the European version, which is said to smell “like goats.” Other species: C.fragrans
also smells like overripe pears, but has a slightly paler cap and whitish or lavender-tinged
flesh in the stalk. C. caesiifolius has a buff to brownish cap, bluish gills when young, and
a fleeting, disagreeable odor. C. camphoratus has a lavender to lilac-white cap (fading to
buff), violet or bluish-lilac gills and stalk when young, and a powerful, remarkably raunchy
odor of rotting meat or vegetables. The stench alone distinguishes it from other lilac
Cortinarii, although a more deeply colored species, C. amethystinus, is said to smell “like
burnt hair.” C. camphoratus grows almost exclusively with conifers and is widely distri¬
buted. I have found it in northern California under Sitka spruce, in November.

Cortinarius alboviolaceus (Silvery-Violet Cortinarius)

CAP 3 -8 cm broad, obtusely bell-shaped becoming convex or broadly umbonate to nearly
plane; surface dry, silky-shining, pale violet soon becoming pale silvery-violet, lilac-white,
or even whitish. Flesh pallid to pale violet; odor mild. GILLS adnate or adnexed or
notched, fairly close, pale violet to purple-gray, then eventually cinnamon-brown as spores
ripen. STALK 4-12 cm long, 0.5-1 (1.5) cm thick at apex, usually club-shaped or enlarged
at base, dry, silky, violet or pale violet above, clothed with whitish silky fibrils below (but
pale violet underneath the fibrils). UNIVERSAL VEIL white, silky, usually forming a
thin, soft, silky sheath over lower half of stalk. CORTINA white, evanescent or leaving
hairs at top of stalk which trap falling spores. SPORE PRINT rusty-brown; spores 7-10
x 4-6 microns, elliptical, minutely roughened.
HABIT AT: Scattered to gregarious or in small clumps in forest humus, associated mainly
with hardwoods; widely distributed. It turns up occasionally in our area in the late fall and
winter under tanoak and other trees, but I have never seen it in large numbers.
EDIBILITY: Unknown.
Left: Cortinarius alboviolaceus. Note the rather long stalk sheathed with copious universal veil
remnants. Right: Cortinarius argentatus (see comments under C. alboviolaceus) is also pale violet
or bluish-violet but has a thicker, stouter stalk with a bulb at the base.

COMMENTS: The beautiful silvery-violet to lilac-white overall color plus the dry silky
cap, mild odor, and whitish universal veil sheath on the stem distinguish this attractive
mushroom from its close relatives in the subgenus Sericeocybe (a closely-knit group of
dry-capped Cortinarii with violet to pale lilac pigments). The flesh is never rusty-brown or
tawny as in C. traganus, and the gills are violet-tinged when young. Inocybe lilacina is
somewhat similar but smaller and more umbonate, with dull brown spores and a spermatic
odor. Similar species include: C. argentatus, lilac-blue to violet-gray but with a thicker,
stouter, prominently bulbous stalk (see above photo) that lacks a silky white sheath and
with a radishlike odor and tendency to turn ochre in age, found mostly under hardwoods
(especially eastern); C. subargentatus, favoring hardwoods, very similar to C. argen¬
tatus, but lacking the odor; C. subpulchrifolius, with a grayish-buff cap that often
develops ochraceous or rusty stains in age, dull purplish immature gills, and a dull purplish,
equal or club-shaped stalk that is sheathed by veil remnants, found under hardwoods in
eastern North America; C. malachius, favoring conifers, violet-tinged at first but its cap
soon becoming ochre or brownish; and C. caninus and C. anomalus, growing under both
hardwoods and conifers, the former with a violet-tinged cap that becomes reddish-brown
and buff to tan universal veil zones on the stalk, the latter with a violet-tinged to gray or
brown cap, distinctly violet flesh and violet stalk apex, and ochre veil zones on the stalk.
For more odoriferous and/ or brightly colored “Sericeocybes,” see C. traganus, and for
violet or violet-tinged hygrophanous Cortinarii, see C. evernius.

Cortinarius armillatus (Bracelet Cortinarius)

CAP 5-13 cm broad, obtuse to broadly bell-shaped or convex, becoming nearly plane or
broadly umbonate; surface smooth or sometimes with small scales in age, not viscid and
only slightly hygrophanous (if at all); dull tawny or yellow-brown to rusty-brown, orange-
brown, or reddish-brown; margin sometimes hung with veil remnants. Flesh thick, pallid
or brownish; odor usually radishlike. GILLS fairly well-spaced, broad, adnatetoadnexed
or notched, pale or dull cinnamon becoming rusty-brown as spores mature. ST ALK 7-15
cm long, 1 -2.5 cm thick at apex, club-shaped (thicker below), dry, whitish to brownish with
one or more (usually 2-3) dull red bands or “bracelets” below the cortina. UNIVERSAL
VEIL fibrillose, forming reddish bands on stalk. CORTINA whitish and copious, often
leaving hairs on upper stalk that turn rusty-brown from falling spores. SPORE PRINT
rusty-brown; spores (7) 9-13 * 5.5-7.5 microns, elliptical, roughened.

448
Left: Cortinarius armillatus, a birch-lover with reddish bracelets on stalk. Right: Cortinarius boulder-
ensis (see comments below) has reddish or vinaceous bracelets but is smaller and purple-tinged.

HABITAT: Solitary or more often scattered or in groups on ground in woods; associated

primarily if not exclusively with birch and found throughout the range of birch (therefore
absent in California). It is especially common in northeastern North America in the late
summer and fall, but I have also seen it in northern Idaho and British Columbia.
EDIBILITY: A good edible—as I can personally attest. It is one of the most distinctive
of the Cortinarii, but you should be very cautious nevertheless!
COMMENTS: The hallmarks of this common eastern species are the reddish bracelets
on the stalk, yellow-brown to cinnamon-brown cap, and association with birch. In the
Pacific Northwest a slightly smaller conifer-lover, C. haematochelis, occurs. It also has
reddish bracelet(s) on the stem, but its spores are much smaller and nearly round. A third
species with reddish (or vinaceous) fibrillose patches or “bracelets,” C. boulderensis, is
even smaller (cap 2-5 cm broad, stalk 3-7 mm thick at apex), with a violet-brown to reddish-
brown or brownish cap and violet-tinged immature gills and stalk apex. It also grows under
conifers in the Pacific Northwest, but I have found something very similar—if not the
same—in our area in mixed woods (see photo above). C. subtestaceus is a similar but
smaller easterner with reddish-brown to brown “bracelets.” Also see C. anomalus
(under C. alboviolaceus) and C. crocolitus (under the C. collinitus group), both of which
have more ochraceous-colored bands or patches on the stalk.

Cortinarius rubripes (Red-Footed Cortinarius)

CAP 4-12 cm broad, obtuse to convex becoming plane or broadly umbonate; surface
hygrophanous but not viscid, watery cinnamon to reddish-brown fading to tawny, ochra-
ceous, or ochre-buff, sometimes tinged with pink or sometimes with concentric zones;
finely silky-fibrillose in age, the margin sometimes tinged violet when young. Flesh thin,
reddish-brown or pallid; odor mild. GILLS fairly well-spaced, adnate to adnexed or
notched and seceding; often tinged violet in button stage, but soon tawny to cinnamon, and
finally cinnamon-brown. STALK 4-9 cm long, 0.5-1.5 cm thick at apex, enlarged below,
dry, pallid or brownish above, the base and mycelium fiery orange to red-orange or peach-
colored. UNIVERSAL VEIL red-orange, leaving remnants at base of stalk. CORTINA
disappearing or leaving a few hairs on upper stalk. SPORE PRINT rusty-brown; spores
7-10 x 5-6 microns, elliptical, roughened.
HABITAT: Solitary to scattered or gregarious under hardwoods; originally described
from Michigan, but it—or something very much like it—also grows in our oak-tanoak-

449
450 CORTINARIACEAE

madrone woods in December and January. In eastern North America it is sometimes

abundant after late summer rains.
EDIBILITY: Unknown
COMMENTS: The bright orange to fiery red stalk base and overall brown to reddish-
brown color combine to distinguish this Cortinarius from most others. A European
species, C. builliardi, is probably the same, in which case that would be the “correct” name
for our species. C. armillatus and C. haematochelis are somewhat similar, but normally
show one to four dull reddish bands on the stalk rather than having a bright red-orange
base. C. miniatopus is a small species (cap 1-3 cm broad) with orangish gills when young
and a slender stalk that is wholly or partially sheathed by Vermillion fibrils (densest at the
base); it is not uncommon under conifers in the Pacific Northwest.

Cortinarius evernius
CAP 3-10 cm broad, conical to bell-shaped at first (and scarcely wider than the stalk), then
becoming convex or obtusely umbonate to plane; surface smooth, markedly hygro-
phanous: violet or brown with a purple tinge when moist, quickly fading to vinaceous or
reddish-brown or paler as it loses moisture; margin at first with whitish silkiness. Flesh thin,
at first violet or violet-tinged, but fadingas it dries or ages. GILLS at first violet with whitish
edges, but quickly fading to brown and then darkening to cinnamon-brown; adnate or
becoming notched, well-spaced. STALK 7-15 (20) cm long, 0.8-2 cm thick, equal or
narrowed toward base, usually rather long and often extending deep into the humus or
moss; not viscid; pale to deep violet when fresh (darker below), but often covered with veil
material at first, and fading as it dries. UNIVERSAL VEIL whitish or violet-tinged,
forming fibrillose patches or zones on stalk which may disappear in age. CORTINA
whitish, often disappearing. SPORE PRINT rusty-brown; spores 8-10 * 5-6 microns,
elliptical, slightly roughened.
HABITAT: Solitary, scattered, or in groups under conifers, often in moss; widely distri¬
buted but mainly northern. It is not a common species, but several difficult-to-distinguish
look-alikes (see comments) are quite ubiquitous, including at least two species in our area.
EDIBILITY: Unknown. Do not experiment!
COMMENTS: This species has been included to represent a large number of medium¬
sized, hygrophanous Cortinarii in the subgenus Telamonia that are violet or violet-tinged
when fresh and moist. They are very difficult to identify and some are so markedly hygro¬
phanous that they lose their original color in an hour or two—or in the time it takes to bring
them home! Other “Telamonias” showing violet shades when fresh and moist include:
C. lucorum, a fairly common conifer-lover that differs from C. evernius only in having
a more or less club-shaped stem; C. saturninus, with a brown cap and more or less equal
stalk that is violet above and whitish below; C. brunneus, which sometimes shows violet
at the stalk apex (see comments under C. laniger) and has an equal to club-shaped stalk;
C. subpurpureus, with yellow or yellowish-buff veil patches on the lower portion of the
stalk; C. adustus and C. impennis, with little or no veil remnants on the stalk; and C.
plumiger, with a grayish-violet to watery violet or grayish-blue-tinged stalk that quickly
fades to whitish and a cap that is brownish to buff beneath a white to grayish-white,
fibrillose-hairy layer. All of these are partial to conifers, but another species, C. torvus,
favors hardwoods and is the most distinctive of the lot. It has broad, widely-spaced gills
that are deep purplish when young, a brownish to deep purplish cap, sweetish odor, and
club-shaped or bulbous stalk that often has a well-formed annulus (ring)—an unusual
feature for a Cortinarius! It is fairly common in eastern North America but I can find no
mention of it being found in the West. Another species that tends to form an annulus,
C. urbicus, does occur in the West, but it has a much paler cap than C. torvus.
CORTINARIUS 451

Cortinarius laniger (Brown Cortinarius)

CAP 3-12 cm broad, broadly bell-shaped or convex becoming broadly convex or some¬
times plane; surface smooth, hygrophanous but not viscid, dark reddish-brown or chest¬
nut-brown to reddish-brown or cinnamon-brown when moist, fading as it loses moisture
to pale reddish-brown or tan; often covered with a white silky coating when very young,
especially at margin. Flesh whitish or colored like cap. GILLS fairly close or rather well¬
spaced, adnate to adnexed or notched, pale rusty-brown becoming a beautiful cinnamon-
brown. STALK 3-10 cm long, 1-3 cm thick, enlarged at base (at least when young), dry,
solid, firm; cinnamon to brown (like cap) beneath a coating or patches of fibrillose whitish
veil remnants. UNIVERSAL VEIL fibrillose or slightly cottony, white, usually leaving a
whitish silky coating or patches and zones of white fibrils on stalk and sometimes even a
slight annulus (ring). CORTINA disappearing or leaving a few hairs on upper stalk.
SPORE PRINT rusty-brown; spores 7-10 * 5-6 microns, elliptical, roughened.
HABITAT: Scattered to gregarious on ground under conifers; widely distributed. I have
seen large fruitings under spruce in the Rocky Mountains, and have also found it in the
Pacific Northwest. In our area similar species (see comments) are common under oak.
EDIBILITY: Unknown. Do not experiment!
COMMENTS: This medium-sized brown Cortinarius with the hygrophanous cap is one
of a multitude of medium-sized brown Cortinarii with hygrophanous caps that belong to
the baffling subgenus Telamonia. C. laniger is best recognized by its cinnamon-brown
color and club-shaped stalk that is covered by whitish veil remnants, and by the complete
absence of violet shades on the fruiting body. Similar “BUM’s” (Boring Ubiquitous
Mushrooms) are legion. Some have an equal to club-shaped or bulbous stalk like that of
C. laniger. These include: C. bulbosus, with a paler or duller brown, swollen stalk and
slightly smaller spores, found under both hardwoods and conifers; C. bivelus, especially
common in the southern Rockies, with a cinnamon-brown to tawny-brown cap and larger
spores; C. triformis, with a strongly hygrophanous cap (dark reddish-brown when moist,
yellow-brown or tan as it dries out), but with a transient veil that leaves only slight traces
(if any) on the stalk, found under both hardwoods and conifers but favoring oak in Cali¬
fornia; C. distans, with minute branlike scales on the cap and widely spaced gills, found
under oak, ponderosa pine, etc.; C. armeniacus, with a swollen or bulbous stalk and deep
yellow-brown cap that fades to orange-buff or pale yellow-orange as it loses moisture;
C. brunneus, a widespread nondescript conifer-lover with an equal to club-shaped, dull
brown stalk whose apex sometimes has a slight violet or vinaceous tinge; C. dilutus and C.
biformis, both smaller with a more or less equal stalk and deep reddish-brown (moist) to
buff (dry) cap, favoring northern conifers (the former with nearly round spores); and C.
pinetorum, a fungal feature of western coniferous forests with a fibrillose, white to silvery-
gray to pale brownish-gray cap that may be slightly tacky when wet, plus brownish gills and
a whitish, more or less club-shaped stalk. Other species have a stalk which is spindle-shaped
and/ or tapered below (or occasionally equal). These include: C. privignus and C. cacao-
color, with a dark or dull vinaceous-brown to cocoa-colored cap when moist (the former
developing a silky whitish sheen as it loses moisture) and stalk which lacks persistent veil
remnants; C. damascenus, similar to the previous two species but with an equal or tapered
stalk that is covered with a persistent coating of white veil fibrils; and C. duracinus, which
has a whitish stalk and a cinnamon-brown cap that fades to buff or pale tan as it dries out.
The above species are but a few of the hundreds of brownish “Telamonias.” As already
pointed out, recognition of these fungi is very difficult and none should be eaten. For
medium-sized “Telamonias” with a violet tinge when moist, see C. evernius, and for
smaller species, see the C. obtusus group.
452 CORTINARIACEAE

Cortinarius obtusus group (Little Brown Cortinarius)

CAP 1-4 cm broad, bell-shaped to conical at first, expanding in age but usually retaining
a blunt umbo; surface smooth, hygrophanous but not viscid, brown to reddish-brown to
tawny-brown and faintly striate when moist, fading to tan or paler as it loses moisture.
Flesh thin, brownish; odor usually radish- or iodine-like (but often faint). GILLS light
to dull brown becoming cinnamon-brown in age, close, adnate to adnexed. STALK
4-8 cm long, 2-6 mm thick, equal or often slightly thicker in the middle and tapered below;
dry, hollow in age, fragile; colored like the cap or often yellower or paler, whitish or pallid
as it loses moisture. UNIVERSAL VEIL and CORTINA scanty, pallid or whitish, disap¬
pearing or leaving a few fibrils on stalk. SPORE PRINT brown to rusty-brown; spores
8-10 x 4-5.5 microns, elliptical, roughened.
HABITAT: Scattered to densely gregarious on ground under conifers, especially pine
and spruce; widely distributed. This species and its numerous look-alikes (see comments)
are common in most regions, including ours.
EDIBILITY: Unknown. Do not experiment!
COMMENTS: The above description encompasses several small boring, brownish
Cortinarii (“LBM’s”) sometimes referred to as the C. obtusus-C. acutus complex. (C.
acutus is slightly slimmer, smaller, and paler than C. obtusus, with a more acute umbo;
another species, C. scandens, is bluntly umbonate like C. obtusus, but has smaller spores.)
The “true” C. obtusus and C. acutus both have rather pale stems (usually paler than the
cap or even whitish). Because of their small size, they might be confused with Inocybes
or Galerinas. The former, however, have duller brown spores while the latter usually have
a yellower fruiting body and typically grow in moss or on wood. Other smalTTelamonias”
are legion, especially under northern or mountain conifers. Even experts have difficulty
differentiating them from each other and there is absolutely no reason for the amateur to
even try. Nevertheless, I will mention a few of them for the categorically-inclined: C.
subcuspidatus and C.fasciatus resemble C. acutus, but the first has yellowish veil material
on the stalk, or if not, can be told by its dark cinnamon-brown gills when young, and the
latter has a darker stalk and tends to grow in clusters. C. stemmatus is a slightly larger
species with a blackish-brown cap that fades to reddish-brown or reddish-cinnamon as it
loses moisture plus a mild odor and whitish veil material on the stalk (see photograph);
it is common under conifers (especially pine in our area). C. nigrocuspidatus can usually
be told by its dark cap with a prominent black umbo, and C. incisus by its minutely scaly,
brownish to reddish-brown cap and white veil remnants on the stalk. C. psammocephalus
has tawny-brown fibrillose scales or patches on both the cap and stalk. C. paleaceus has
a blackish-brown cap that fades to dingy brown in age, whitish veil remnants on the cap

Cortinarius stemmatus (see comments under C. obtusus) is one of dozens of small, brownish, difficult-
to-demystify Cortinarii. The cap is blackish-brown when moist but turns reddish-brown as it loses
moisture. Note umbonate cap.
CORTINARIUS 453

margin and stalk (at least when young), and a geranium-like odor; it grows in mossy bogs
or other wet places under spruce. C. decipiens also has a dark cap (vinaceous-brown or
darker when moist), but it lacks white veil remnants on the stalk, while C. washingtonensis
can easily be told by its black-staining gills. All of the above species are especially common
under northern conifers, but some may also occur with hardwoods. Finally, there are a
number of small “Telamonias” that are distinctly violet or violet-tinged when fresh and
moist, including: C. pulchellus, violet overall when young and especially fond of alder;
and C. subflexipes, which has a rusty-brown to ochraceous cap but shows violet in the
gills and stalk apex when young. For larger “Telamonias,” see C. laniger and C. evernius.

Cortinarius cinnamomeus group

CAP 1.5-6 cm broad, obtusely conical becoming convex to more or less plane or often with
an umbo; surface dry, finely fibrillose, yellowish to olive-yellow to tawny, ochre-buff,
yellow-brown, or olive-brown, becoming more cinnamon-colored in age. Flesh thin,
yellowish or olive-yellow. GILLS adnate to adnexed or notched, close, yellow to olive-
yellow becoming tawny, then rustier in age as spores ripen. STALK 2.5-10 cm long, 3-5
(10) mm thick, more or less equal, often curved, dry, finely fibrillose or with small fibrillose
scales, yellow to yellow-brown or ochraceous, often browner toward base and duller in age.
CORTINA yellow or yellow-olive, disappearing or leaving inconspicuous hairs on upper
stalk. SPORE PRINT rusty-brown; spores 6-9.5 * 4-5 microns, elliptical, minutely
roughened.
HABITAT: Widely scattered to gregarious on ground under conifers, especially pine;
widely distributed, and common in our area from late fall through early spring.
EDIBILITY: To be avoided! It is said to be edible, but some very similar species (see C.
gentilis) are deadly poisonous!
COMMENTS: Also known as Dermocybe cinnamomea, this contrary little Cortinarius
and its numerous look-alikes are not cinnamon-colored at all. Rather, the dominant colors
are yellow, yellow-olive, and ochre. The color plus the slender stem, dry cap, and evan¬
escent fibrillose veil (cortina) are characteristic of a large complex of species in the
subgenus Dermocybe which are best left to the specialist. It may actually be that the“true”
C. cinnamomeus does not occur here, but a number of very similar species certainly do,
including: C. cinnamomeo-luteus, said to be very common in the Pacific Northwest;
C. humboldtensis{from northern California) and C. olivaceopictus, the latter with reddish
to orange fibrils on the stalk; C. raphanoides, olive-yellow, with a radish odor; and C.
thiersii, with larger spores, especially common under mountain conifers. Another group of

Cortinarius cinnamomeus group is common under conifers, but identifying the various species
in the “complex” is very difficult. Some forms have sharply conical caps (right), others do not (left).
454 CORTINARIACEAE

species, typified by C. croceofolius, have saffron to rusty-yellow or orange gills when

young and a browner or more cinnamon-colored cap. They are just as common as the C.
cinnamomeus group, and their ranks include: C. aurantiobasis, with larger spores; C. zakii,
with deep orange gills when young and stalk coated with brown to vinaceous-brown fibrils;
and C. aureifolius, a very distinctive species that usually grows in sand (often partially
buried!) under pines, and has orange to brownish-orange(or in one form, yellowish) gills, a
cinnamon-brown to dark reddish-brown cap, and larger spores. Finally, there is C. callis-
teus, a bright yellow to orange-ochre species with a thick club-shaped stem (at least 1 cm
thick at apex and 2.5 cm or more thick at base). None of these species should be eaten.
(Note: some authorities have a concept of C. cinnamomeus which more closely fits the
C. croceofolius of this book, and vice-versa, at least insofar as gill color is concerned.)

Cortinarius phoeniceus var. occidentalis

CAP 2.5 -8 cm broad, convex becoming plane or broadly umbonate; surface dry, finely
fibrillose or silky, maroon-red or dark red to reddish-brown; margin sometimes lobed.
Flesh thin, reddish near cuticle, olive-brownish near gills; odor mild or slightly radishlike.
GILLS deep red, vinaceous-red, or blood-red, becoming rustier as spores ripen; adnate to
adnexed or notched. STALK 3-10 cm long, 0.4-1.2 cm thick, more or less equal, dry,
fibrillose, yellowish or ochre or sometimes brownish in old age. CORTINA scanty, yel¬
lowish, disappearing or leaving a few hairs on upper stalk. SPORE PRINT rusty-brown;
spores 6-8 * 4-5.5 microns, elliptical, minutely roughened.
HABITAT: Solitary to scattered or in groups under conifers in the Pacific Northwest and
California; sometimes common in the fall and early winter. In our area it also grows with
madrone and/ or huckleberry, but never in large numbers.
EDIBILITY: Not recommended, but an excellent choice for dyeing (not dying!).
COMMENTS: The beautiful deep red gills and maroon-red to reddish-brown cap are set
off nicely by the yellowish stem, making this Cortinarius (or Dermocybe) relatively easy to
identify. Other species: C. semisanguineus also has red gills, but has a slender yellow stem
and yellowish to ochre or olive-brown cap; it occurs in California, but is more common in
eastern North America and the Pacific Northwest. Another species, C. californicus, has a
smooth, hygrophanous cap that is dark rusty-red when moist but fades as it loses moisture.
It has orange-red to rusty-orange or “burnt sienna” gills, and a rather long (7-15 cm) stalk
that is paler than the cap (dull orange or even paler, often with orange fibrils from the
cortina). It is known only from west coast and occurs in a variety of habitats, but seldom in
large numbers. For completely red species, see C. sanguineus.

Cortinarius sanguineus (Blood-Red Cortinarius)

CAP (1) 2-5 cm broad, obtusely bell-shaped becoming convex to plane or broadly umbo¬
nate; surface dry, silky or finely fibrillose, colored evenly deep carmine-red to blood-red
or deep red. Flesh thin, blood-red to reddish-purple; odor mild or faintly pleasant. GILLS
close, adnate to adnexed, blood-red to dark blood-red, but in age dusted with cinnamon-
colored spores. STALK 4-10 cm long, 3-5 (8) mm thick, equal or slightly thicker below,
same color as cap or slightly darker, dry, base with yellowish to yellow-orange or some¬
times pinkish downy hairs. CORTINA red or at least tinged red, often scanty and disap¬
pearing or leaving a few hairs on upper stalk. SPORE PRINT rusty- to reddish-brown;
spores 6-9 * 4-6 microns, elliptical, minutely roughened.
HABITAT: Solitary to widely scattered or in small groups, mainly under conifers and
often in beds of moss; widely distributed but generally rare. In favorable years it fruits in
the fall under spruce and fir along the northern California coast, but never in large num¬
bers. I have found it as far south as Mendocino County in California.
CORTINARIUS 455

EDIBILITY: Unknown. Do not experiment! Like many species in the subgenus Dermo-
cybe, it can be used to dye wool or yarn, yielding a brilliant array of pinks, reds, and purples.
COMMENTS: Also known as Dermocybe sanguined, this beautiful mushroom is easily
recognized by its blood-red to dark red color, small size, and evanescent cortina. It might
be mistaken for a brightly colored Hygrocybe, but the spores are not white and the gills
are not waxy. There is no yellow in the capand/ orstalkasin C.phoeniceusv ar. occidentalis
and C. semisanguineus, and the cap is not hygrophanous as in C. californicus (see
comments under C. phoeniceus var. occidentalis). Other completely red Dermocybes
include: C. puniceus, said to differ by its ochraceous to golden-brown cortina, dark red to
purple-red color, and preference for hardwoods; and C. cinnabarinus, also favoring
hardwoods (but not limited to them), with a slightly larger, cinnabar-red to rusty-red
or dark red fruiting body and larger spores. The latter species has often been reported from
eastern North America, but according to Cortinarius-experts Alexander Smith and
J oseph Ammirati, American material differs slightly from the European version. (At least
one common hardwood-lover has been recognized as a “new” species, C. marylandensis.)

INOCYBE
Small to medium-sized, typically terrestrial mushrooms. CAP often umbonate or conical; surface
typically dry and silky, woolly, scaly, or radially fibrillose; margin often splitting at maturity; odor
of flesh often distinctive (usually unpleasant). GILLS adnate to adnexed or free, usually brown at
maturity (but often with whitish edges). STALK central, often rather slender; apex often powdery
or with small flakes. VEIL absent, or if present then fibrillose or cobwebby and usually evanescent.
VOLVA absent. SPORE PRINT brown. Spores smooth, warty, angular, or nodulose, lacking an
apical germ pore. Gills with cystidia, at least on edges. Cap cuticle filamentous.

INOCYBE is a large, listless, and lackluster assemblage of mundane, malodorous brown

mushrooms that are of little interest to the average mushroom hunter except that many are
poisonous. The best means of recognizing an Inocybe is by its characteristically silky,
fibrillose, minutely scaly, and/or woolly cap which is often umbonate and seldom viscid.
The spore color is some shade of brown, and is generally duller than that of Cortinarius. In
addition, most Inocybes have a noticeable odor—occasionally sweet or fruity as in /.
pyriodora, but more often unpleasant (pungent, spermatic, fishy, or like fresh green corn
but not often radishlike as in Hebeloma).
Like Cortinarius, I nocy bes are largely terrestrial and mycorrhizal and are a maj or fungal
facet of temperate forests. U nlike Cortinarius, they are not the least bit colorful. They come
in an endless, senseless procession of boring browns, yucky yellows, gratuitous grays, and
wishy-washy whites, with only I. lilacina (among the common species) deviating from the
norm. Almost without exception they not only qualify as “LBM’s” (p. 32), but as
“BUM’s” (“Boring Ubiquitous Mushrooms”), and it is necessary to know the size and
shape of the spores and cystidia before an accurate identification can be made. Even then,
unraveling them is a trying and tedious task whose futility is only exceeded by its
pointlessness, and underscored by the sad fact that most Inocybes are poisonous. I would
venture to guess, in fact, that Inocybe contains a higher percentage of poisonous species
than any other major mushroom genus, including Amanital The toxin is muscarine,
which can be fatal in large amounts (see p. 894). It is therefore advisable to avoid all
Inocybes, even those rumored to be edible.
Inocybe is a very large genus—almost as immense, in fact, as Cortinarius. (The late
Daniel Stuntz listed over 400 North American species in section Inocybium alone, i.e.,
those with smooth spores and cystidia on the faces of the gills!) A mere nine Inocybes are
described here, which for most intents and purposes are eight too many! Several other
species are keyed out.
456 CORTINARIACEAE

Key to Inocybe
l. Odor fragrant or fruity (though sometimes sickening, as in rotten fruit) .2
1. Odor spermatic, pungent, fishy, like green corn, mild, etc., but not fragrant or fruity . 5
2. Cap olive to olive-brown at center./. corydalina (see I. pyriodora, p. 459)
2. Not as above . 3
3. Either cap conical with reddish-brown to wine-red radiating fibrils or cap with a whitish
patch of universal veil tissue and odor with a green corn component . 7
3. Not as above . 4
4. Odor of sweet peas .I. suaveolens (see I. pyriodora, p. 459)
4. Odor spicy or fruity or otherwise (but not of sweet peas) .I. pyriodora & others, p. 459
5. Stalk base or lower portion dull bluish-green, olive, or dull greenish . . I. calamistrata, p. 462
5. Not as above . 6
6. Cap grayish-lilac to grayish-brown or pinkish-brown with a smooth white or creamy, umbonate
center (or in one form cap entirely whitish and stalk pinkish); stalk typically pruinose(granular-
dandruffy) most of its length, with a bulb at base .I. albodisca (see I. maculata, p. 458)
6. Not as above (but may have some of above features) . 7
7. Cap small (2-5 cm broad) and yellowish to brown, but with a patch of whitish universal veil
tissue at the center (or sometimes overall), at least in most specimens; stalk white when young
and not shaggy or woolly ./. lanatodisca & others (see I. maculata, p. 458)
7. Not as above . 8
8. Cap 3-8 cm broad, at first covered with a whitish universal veil that leaves patches of tissue near
margin; stalk 5-10 mm thick, also sheathed with woolly remnants of the veil, dingy brownish
beneath the remnants; widespread, but especially common in the Rockies . . . /. leucoblema
8. Not as above . 9
9. Stalk smooth or fibrillose or occasionally with a few small fibrillose patches or scales .... 10
9. Stalk distinctly scaly (i.e., with numerous small scales similar in color to those on cap) ... 25
10. Some part of fruiting body lilac to violet when fresh (but may fade or discolor in age).
. /. lilacina& others, p. 461
10. Not as above (but stalk or cap may be pink, carmine, vinaceous, etc.) . 11
11. Cap white, smooth to very finely silky and not discoloring appreciably in age or when bruised;
stalk not pruinose (granular-dandruffy) or pruinose only at apex.I. geophylla, p. 460
11. Not as above (but cap may be white initially and then discolor in age, or cap may have white veil
remnants) . 12
12. Cap and stalk whitish at first, but usually developing pink, red, or orange tones as it ages; very
common, especially under conifers .I. pudica, p. 460
12. Not as above . 13
13. Cap conical when young, often with a pointed umbo in age, the surface radially fibrillose (and
often splitting in age) but not woolly or scaly, creamy to straw-colored, yellow-brown or brown
(but not reddish); gills sometimes with an olive or greenish tinge . 14
13. Not as above . 15
14. Odor resembling that of fresh green corn; cap creamy to straw-colored .... I. sororia, p. 457
14. Odor spermatic; cap usually somewhat darker (yellow-brown to brown), at least at the center
.I.fastigiata & others (see /. sororia, p. 457)
15. Cap brownish with a very prominent gray to dark brown or black “nipple” (umbo) at center;
widely distributed .I. fuscodisca
15. Not as above . 16
16. Cap with reddish-brown to carmine or wine-red fibrils, more or less conical when young; stalk
whitish at least at apex; found mainly with oak (in California) ./. jurana, p. 458
16. Not as above . 17
17. Stalk pale pink to salmon-pink; cap reddish-brown, at least at center .
.I. laetior&I. oblectabilis (see I. jurana, p. 458)
17. Not as above . 18
INOCYBE 457

18. Stalk entirely pruinose (minutely granular-scurfy-dandruffy) and discoloring dark gray to dark
brown or vinaceous-brown from the base upward in age .
./. picrosma, I. leucomelaena, & others
18. Not as above . 19
19. Fruiting body fairly small (cap typically up to 4 cm broad; stalk less than 5 mm thick) ... 20
19. Fruiting body medium-sized (cap usually at least 3 cm broad and stalk at least 5 mm thick 23
20. Cap with loosely-arranged or torn-up fibrils that give it a shaggy or ragged appearance, espe¬
cially at margin; color more or less dark brown./. lacera (see I. lanuginosa, p. 462)
20. Not as above; cap usually woolly or scaly or differently colored . 21
21. Cap conical to bell-shaped, the surface often splitting radially but not scaly; stalk with a basal
bulb./. mixtilis (see I. sororia, below)
21. Not as above; cap usually scaly or woolly. 22
22. Cap dark brown to blackish, with small erect scales; often growing on rotten wood; spores
warty./. lanuginosa, p. 462
22. Not as above; usually terrestrial .I.flocculosa & many others (see /. lanuginosa, p. 462)
23. Cap yellow-brown to ochre or dull cinnamon (sometimes darker at center), fibrillose or in age
becoming somewhat scaly; stalk similarly colored at first but becoming brown to smoky-brown
from the base upward; common under conifers in the Pacific Northwest.I. olympiana
23. Not as above; cap usually brown to dark brown or chestnut-brown (but sometimes overlaid
with whitish veil remnants) . 24
24. Stalk with a large turniplike bulb at base; cap usually silky-fibrillose or smooth; spores angular
.I. nap ip es (see I. sororia, below)
24. Stalk equal or with small bulb; cap sometimes scaly; spores smooth I. maculata & others, p. 458
25. Cap relatively large (5-12 cm broad); stalk fairly thick (at least 5 mm); both cap and stalk with
dark brown scales (those on stalk arranged in more or less concentric rings) .
./. sp. (unidentified) (see I. calamistrata, p. 462)
25. Not as above; smaller . 26
26. Cap and stalk yellowish to yellow-brown to cinnamon.
.I. terrigena & I. caesariata (see I. calamistrata, p. 462)
26. Cap and stalk with darker brown scales ./. hystrix(see I. calamistrata, p. 462)

Inocybe sororia (Corn Silk Inocybe)

CAP 2-8 (10) cm broad when expanded, sharply conical or bell-shaped when young, often
expanding in age but retaining a prominent umbo, the margin usually uplifted or splitting
in old age; surface dry, silky, radially fibrillose, creamy to pale yellowish, honey-yellow,
or straw-colored, often somewhat browner at the center or in age. Flesh thin; odor usually
strongly pungent, like freshly husked or green corn. GILLS close or crowded, pallid
becoming yellowish, then olive-yellow and finally brownish-gold or brown, the edges
usually paler; adnate to adnexed, often seceding to free. STALK 3-14 cm long, 2-5 (10) mm
thick, equal or with a slightly enlarged base, white or tinged cap color, fibrillose and often
scurfy. VEIL absent. SPORE PRINT brown; spores 10-13 (17)x 5-8 microns, elliptical or
bean-shaped, smooth. Cystidia on gills thin-walled.
HABITAT: Solitary, scattered, or in small groups on ground in woods; widely distributed.
In our area it is common in the fall and winter under live oak and pine; farther north it
favors conifers. I have seen large fruitings locally as well as in Oregon.
EDIBILITY: Poisonous! It contains high concentrations of muscarine.
COMMENTS: A common and prominent Inocybe, easily recognized by its sharply
conical to umbonate, creamy to yellowish, fibrillose cap and odor of fresh green corn. The
stem is often quite long and slender. I. fastigiata is a very similar, widespread species with a
strongly spermatic odor and a slightly darker (yellow-brown to brownish-ochre) cap, at
least at the center. Other species: I. cookei has a silky yellowish cap, small but prominent
bulb at the base of the stem, slight odor, and smaller spores; /. napipes has a blunter
Inocybe sororia. Left: Young, relatively robust specimens. Right: A slender mature specimen. Note
the conical or umbonate cap. Green corn odor is also distinctive.

brownish cap, a cortina (cobwebby veil) when young, a turniplike bulb at the base of the
stem, and warty (nodulose) spores; /. mixtilis also has warty spores, but is smaller than
/. napipes and lacks a cortina. The latter two species are fairly common under conifers in
the Pacific Northwest, but I have not seen them locally. All of the above are poisonous.

Inocybe jurana (Reddish Inocybe)

CAP 2 -8 cm broad when expanded, conical or bell-shaped, expanding in age but usually
retaining an umbo; surface dry, radially fibrillose, buff or brownish with darker brown
fibrils, but soon flushed carmine, reddish-brown, or wine-red (vinaceous); center some¬
times with small scales, margin often splitting in age. Flesh tinged pinkish or vinaceous,
thin; odor mild to somewhat fruity or unpleasant. GILLS pallid becoming grayish-brown
or dull brown, close, adnate to adnexed or even free. STALK 2-8 cm long, 4-10 (15) mm
thick, equal or slightly enlarged at base, dry, white or flushed cap color below. VEIL absent.
SPORE PRINT brown; spores 9-15 * 5-8 microns, elliptical or bean-shaped, smooth.
Cystidia on gills thin-walled.
HABITAT: Solitary, scattered, or in small groups on ground in woods and at their edges;
widely distributed. In our area I have found it twice under live oak in the winter.
EDIBILITY: Not recommended. According to some sources it is edible, but why tempt
fate? Several similar species are poisonous.
COMMENTS: The deep reddish or vinaceous-toned cap is the hallmark of this illustrious
Inocybe. In other respects it rather resembles I. sororia, but does not smell like fresh
green corn. Other species: /. laetior has a bright salmon-pink stem and dark reddish-brown
centered cap with a yellowish margin; it occurs under conifers in the Pacific N orthwest and
is one of numerous Inocybes “discovered” and named by Daniel Stuntz. /. oblectabilis
has a pale pink stalk and reddish-brown cap.

Inocybe maculata (Brown Inocybe)

CAP 2-8 cm broad, conical or bell-shaped, becoming convex or plane with an obtuse
umbo; surface dry, radially fibrillose, dark brown to chestnut-brown, at first with whitish
down (at least at center) which may break up into small scales or wear away; margin often
lobed, splitting in age. Flesh thin, whitish; odor mild or slightly aromatic. GILLS pale

458
INOCYBE 459

grayish becoming grayish-brown to olive-brown or dull brown, close, adnate to adnexed

or seceding. STALK 3-10 cm long, 0.5-1.2 cm thick, equal or with an enlarged base,
fibrillose-striate, white or flushed cap color (brownish). VEIL absent except for whitish
down on cap. SPORE PRINT brown; spores 9-12 * 4.5-6.5 microns, elliptical or bean¬
shaped, smooth. Cystidia on gills thin-walled.
HABITAT: Solitary or in small groups in woods and under trees; widely distributed.
Occasional in our area in the fall and winter.
EDIBILITY: Poisonous; it contains muscarine.
COMMENTS: Any medium-sized to large Inocybe with a dark brown fibrillose or hairy
cap and smooth spores can be referred here and conveniently forgotten about. I have
encountered forms with an obnoxious odor, others which were mild and still others with a
slightly aromatic or truffle-like smell. Undoubtedly more than one species is involved.
Another fairly large species, /. serotina, has a brown cap with a whitish to yellowish-buff
center and a bulb at the base of the stem. /. napipes{set comments under /. sororia) also has
a basal bulb and can be quite large. There are also several smaller species that feature
white universal veil tissue at the center of the cap. These include: I. lanatodisca, cap brown
to yellow-brown with a moldy-looking patch of whitish fibrils at the center which may
disperse in age, a whitish stalk (at least when young), and cystidia only on the edges of the
gills plus a pungent to sweetish odor, fairly common in our area under oak but also
occurring with conifers; and /. sindortia, similar to I. lanatodisca, but with cystidia on
the faces of the gills as well as the edges. Another species, I. albodisca, has a lilac-gray
to pinkish-brown (or sometimes whitish) cap withasmoothwhitishumbonate centeranda
basal bulb on the stalk; it is common in northern latitudes under both hardwoods and
conifers. None of the above species should be eaten.

Inocybepyriodora (Fragrant Inocybe)

CAP 2-7 cm broad, bell-shaped or obtusely conical becoming broadly umbonate; surface
dry, at first silky-smooth but often fibrillose or fibrillose-scaly (especially at center) in age
(or with torn-up appearance); white, the fibrils becoming dingy ochre to yellow-brown or
brownish, sometimes pinkish- or reddish-stained in age. Flesh thin, white, slowly staining
pinkish or reddish when cut; odor fragrant, usually spicy (cinnamon- or matsutake-like)
but sometimes like overripe pears or unpleasant in old age. GILLS adnate to adnexed or
notched, close, whitish becoming dull cinnamon, brown, or reddish-tinged. STALK 4-10
cm long, 3-15 mm thick, more or less equal, white throughout or colored like cap except for
very apex, sometimes developing pinkish stains; smooth or finely fibrillose. VEIL fibril¬
lose or cobwebby, evanescent. SPORE PRINT brown; spores 7-10 (12) * 4.5-7.5 microns,
elliptical to bean-shaped, smooth. Cystidia on gills with slightly thickened walls.
HABITAT: Solitary, scattered, or in small groups in woods or at their edges, underbrush,
along trails, etc., favoring hardwoods but also occurring with conifers; widely distributed.
It is not uncommon in our area in the fall and winter in mixed woods and under oak.
EDIBILITY: To be avoided—despite the enticing odor it is probably poisonous.
COMMENTS: This is one of several nondescript Inocybes with a spicy or fruity fragrance.
The odor is often reminiscent of the matsutake (Armillariaponderosa), but is sometimes
fruity. Other fragrant species include: /. bongardii, cap scaly, odor fruity, spores larger;
I. corydalina, cap whitish with brownish fibrils and an olive to olive-brown center and
a strongly pungent or sickeningly sweet odor (like rotting fruit); I. godeyi, a reddening
species with a slight fruity odor and abrupt bulb at the base of the stem;/, cookei, which
has a slight fruity odor and abrupt basal bulb but does not redden; I. hirtella, with an
almond extract or cyanide odor when fresh; and /. suaveolens, which smells like sweet
peas and has nodulose-angular spores and is fairly common in the Pacific Northwest.
Inocybe pudica is a whitish species that develops reddish to salmon-orange tones as it ages. It is
especially common under conifers.

Inocybe pudica (Blushing Inocybe)

CAP 2-6 (8) cm broad, conical or bell-shaped when young, becoming convex to nearly
plane at maturity, but usually retaining an umbo; surface dry or tacky, silky-fibrillose to
nearly smooth, white at first but developing pinkish to reddish or orange stains as it ages.
Flesh thin, white; odor unpleasant or spermatic. GILLS pallid or flushed pinkish or
orange, becoming grayish-brown or dull brown as spores mature, close, adnate to
adnexed, notched, or free. STALK 4-8 cm long, 0.4-1 cm thick, equal or enlarged at base,
smooth or silky-fibrillose, firm; white, but discoloring like the cap. VEIL fibrillose or
cobwebby, whitish, evanescent. SPORE PRINT brown; spores 7-10 x 4-6 microns, ellip¬
tical or bean-shaped, smooth. Gills typically with both thick- and thin-walled cystidia.
HABITAT: Scattered or in groups or troops on ground under conifers; widespread,
but especially common on the west coast. In our area it is often abundant in the late fall,
winter, and early spring under pine and Douglas-fir. It seems to favor brushy areas or
immature second-growth stands.
EDIBILITY: Poisonous; like most Inocybes, it contains muscarine.
COMMENTS: This is one of the most common Inocybes of the western United States
as well as one of the most easily recognized. The tendency of the white fruiting body to
“blush” pink, red, or orange plus the growth under conifers and relatively smooth cap are
the principal fieldmarks. The cap may be slightly tacky or slippery in wet weather, but is
not truly viscid. /. pyriodora is somewhat similar but has a fragrant or spicy odor.

Inocybe geophylla (Little White Inocybe)

CAP 1 -3 (4) cm broad, conical to bell-shaped, expanding in age to plane but often retaining
an umbo; surface dry, white or sometimes with a slight yellow tinge in age; smooth to silky
or finely fibrillose; margin sometimes uplifted or split in age. Flesh thin, white; odor disa¬
greeable (spermatic). GILLS adnate to adnexed or notched, close, at first pallid, then
grayish, finally dull brown. STALK 2-6 cm long, 2-5 mm thick, equal or slightly thicker
at base, firm, white or grayish-white, finely fibrillose. VEIL fibrillose, whitish, evanescent
or leaving a slight hairy zone on stalk. SPORE PRINT brown; spores 8-10 x 5-6 microns,
elliptical, smooth. Cystidia on gills thick-walled.
HABITAT: Scattered to gregarious on ground (or occasionally very rotten wood) in
woods; widely distributed and common. It is abundant in our area throughout the
mushroom season, especially under live oak, pine, and Douglas-fir.
EDIBILITY: Poisonous; it contains muscarine.

460
Inocybe geophylla is small but common. Note silky white cap and brown gills; odor is spermatic.

COMMENTS: The small size, white umbonate cap, and dull brown gills at maturity
characterize this ubiquitous little Inocybe. It might be mistaken at first glance for a small
waxy cap (Camarophyllus), Mycena, or Alboleptonia, but the spores and mature gills
are brown. It also resembles I. pudica, but does not “blush.”

Inocybe lilacina (Lilac Inocybe)

CAP 1.5-4 (5) cm broad, obtusely conical or bell-shaped, expanding in age but often
retaining an umbo; surface dry, silky-fibrillose to nearly smooth, pale to deep lilac, but
often with pinkish, gray, or brownish tones (or ochre at center), and often paler or whiter in
age. Flesh thin, white or tinged lilac; odor disagreeable (spermatic). GILLS adnate to
adnexed or notched, close, pallid or tinged lilac, then grayish to dull brown, the edges often
whitish. STALK 2.5-6 cm long, 0.3-1 cm thick, equal or slightly thicker at base, finely
fibrillose, firm, colored more or less like cap or with white areas; base whitish. VEIL
fibrillose, evanescent or leaving a very slight hairy zone on stalk. SPORE PRINT brown;
spores 7-9 * 4-5 microns, elliptical, smooth. Cystidia on gills thick-walled.
HABITAT: Solitary, scattered, or in groups on ground in woods; widely distributed. In our
area it can be found in the late fall and winter under pine and Douglas-fir, but is not as
numerous as its close relative I. geophylla.
EDIBILITY: Poisonous; like most Inocybes it contains muscarine.
COMMENTS: Also known as /. geophylla var. lilacina, this species is about the only
common Inocybe that is the least bit colorful, although faded caps can be so pale that they
resemble I. geophylla. The umbonate cap, relatively small size, and brown spores distin¬
guish it from the blewit (Clitocybe nuda) and most other purple or lilac mushrooms.
Several species of Cortinarius resemble it, but are generally larger and have brighter (rusty-
brown) spores. There are also several other Inocybes which are violet or violet-tinged
when fresh. I. obscurioides, for instance, is one of several species with a violet stalk or stalk
apex and a brownish cap that typically shows violet only at the margin (if at all).

Inocybe lilacina is lilac-tinged when fresh. Note umbonate cap of most specimens.
462 CORTINARIACEAE

Inocybe lanuginosa (Woolly Inocybe)

CAP 1.5-3 (4) cm broad, convex to bell-shaped becoming umbonate, plane, or with an
uplifted margin; surface dry, densely woolly or scaly, dark brown to brownish, the scales
small and often erect. Flesh thin, watery brownish; odor mild to slightly unpleasant
(spermatic). GILLS adnate, adnexed, or notched, fairly close, pallid becoming grayish,
then dull brown or cinnamon-brown. STALK 2-5 (8) cm long, 2-4 (7) mm thick, more or
less equal, fibrillose or fibrillose-scaly, colored more or less like cap except for paler apex.
VEIL cobwebby, pallid, evanescent. SPORE PRINT brown; spores 8-10.5 * 5-7 microns,
elliptical with blunt warts (nodulose). Cystidia on gills typically thin-walled.
HABITAT: Solitary or in small groups on very rotten wood or in forest humus; widely
distributed. It favors conifers but also grows with hardwoods; I have not seen it in our
area, but there are dozens of similar species.
EDIBILITY: Unknown, but do not experiment—it may very well contain muscarine!
COMMENTS: There are innumerable little brown Inocybes with hairy or minutely scaly
caps. They are differentiated largely on microscopic characters such as the shape and size
of their cystidia and spores. Most have an unpleasant or spermatic odor, and none should
be eaten. This one is distinctive because it often grows on very rotten wood—highly
unusual behavior for an Inocybe! Other brownish species—all typically terrestrial—
include: I. lacera, widespread in a variety of habitats but especially common under aspen,
with a torn-up or raggedly scaly cap and long, nearly cylindrical, sometimes warty spores;
/. flocculosa, one of dozens of species with a hairy cap, smooth (bald) stalk, and smooth
spores; and I. agardhii, one of several species with a woolly-scaly cap that hang out with
alder and willow. For still other species, see the key to Inocybe.

Inocybe calamistrata (Scaly Inocybe)

CAP 1-4 cm broad, bell-shaped to convex, expanding only slightly in age; surface dry,
breaking up into densely-arranged scales, dark brown to coffee-brown; margin not typi¬
cally splitting. Flesh thin; odor spermatic or fishy. GILLS adnate to adnexed or free, close,
brown to dull cinnamon-brown or colored like cap, the edges whitish. STALK 4-10 cm
long, 3-5 mm thick, equal or tapering upward, firm, covered with recurved scales (like cap)
which may be obliterated or wear away in age; brown to dark brown (like cap), the base or
lower portion dingy greenish-blue to olive-green (both inside and out). VEIL disappearing
(not forming a distinct ring on stalk). SPORE PRINT brown; spores 9-13 x 4.5-6.5
microns, elliptical-oblong, smooth.

HABITAT: Solitary, scattered, or in small groups on ground under conifers or some¬

times hardwoods; widely distributed, but not common. I have seen it in several localities,
but have yet to find it in our area.
EDIBILITY: Unknown. A recent study revealed the presence of psilocybin.

COMMENTS: The blue-green to olive-green stem base and scaly convex cap which does
not split radially in age are the main features of this uninteresting agaric. It is included here
as a representative of those Inocybes with scales on both the cap and the stem. Others
include: /. hystrix, with small, brown to dark brown scales on both the cap and stalk; I.
terrigena, larger and fleshier (in fact, somewhat reminiscent of a Pholiota), with golden-
brown to cinnamon scales on the cap and stalk and sometimes a slight annulus (ring), wide¬
spread but especially common under aspen in the southern Rocky Mountains;/, caesar-
iata, somewhat similar in color to I. terrigena but with smaller scales, fairly common in
eastern North America; and finally, a very robust, unidentified local species with a dark
brown scaly cap and several zones of concentrically-arranged dark brown scales on the
stem (cap 5-12 cm broad, stalk 0.5-1.5 cm thick!). None of these have the blue-green or
olive-green stem base of I. calamistrata.
 463

HEBELOMA
Small to medium-sized, terrestrial mushrooms. CAP typically viscid when wet and more or less
smooth; white to buff, tan, or some shade of brown. Flesh often with a radishlike odor. GILLS
adnate to adnexed or notched, usually brown at maturity; often with whitish edges. STALK fleshy,
central; apex often powdery or with small flakes. VEIL usually absent or if present thencobwebby-
fibrillose and evanescent. VOLVA absent. SPORE PRINT brown (or rarely reddish-brown).
Spores typically elliptical, smooth or roughened, lacking a germ pore. Cystidia typically present on
gill edges, often conspicuous. Cap cuticle filamentous.

THIS is yet another faceless and featureless collection of brownish mushrooms. Those
that are too large to qualify as “LBM’s” most certainly fall into the category of “BUM’s”
(“Boring Ubiquitous Mushrooms”). The stalk is fleshy and the gills attached but not
decurrent, giving the fruiting body the stature of a Tricholoma or Entoloma. The odor
is usually mild or radishlike, not spermatic as in Inocybe, and the cap is typically smooth
and viscid rather than dry and fibrillose or scaly. Hebeloma is closely related to Cortinarius,
but that genus has rusty-brown spores and a cobwebby veil (cortina). Some Hebelomas
have a cortina, but can be distinguished by their duller spore color and/or the presence of
sterile cells (cystidia) on the edges of the gills, plus the frequently powdered or flaky stem
apex. Agrocybe is also similar, but has a cellular cap cuticle and usually grows in grass,
dung, wood or wood chips, or cultivated soil. Hebelomas, in contrast, are largely mycor-
rhizal. As a result, they are found in forests or at their edges or on tree-studded lawns and
cemeteries. (However, H. syriense feeds on decaying corpses and is said to have led to
the discovery of at least one crime!)
About 200 species of Hebeloma occur in North America, but none are exceptionally
distinctive or colorful. Whites, browns, tans, and buffs predominate, making identification
on the basis of color hopeless. More minute measures of individuality must be taken into
account, such as the size and shape of the cystidia and spores. Many mushroom-hunters,
however, prefer to recognize only two varieties—the big brown ones and the not-so-big
brown ones! Hebelomas should not be eaten, as some are definitely poisonous and the
group as a whole is poorly known. Three widespread species are described here.

Key to Hebeloma
1. Spore print reddish to pinkish-brown or pinkish-cinnamon . 2
1. Not as above; spore print dull brown or at times rusty-brown . 3
2. Cap whitish when fresh, but often aging ochre, gray, or brownish; spores elliptical {not angular)
.H. sarcophyllum (see H. crustuliniforme, p. 464)
2. Not as above; spores more or less angular . (seeEntoloma, p. 242)
3. Odor spermatic; cap often (but not always) umbonate and rather small . (see Inocybe, p. 455)
3. Not as above . 4
4. Veil present when young, disappearing or leaving remnants on the cap margin and/or a slight
ring (annulus) or fibrillose sheath on the stalk . 5
4. Veil absent or rudimentary. 6
5. Lower portion of stalk sheathed with conspicuous veil remnants; cap often with veil remnants
also, especially near margin .H. strophosum (see H. mesophaeum group, p. 465)
5. Stalk not conspicuously sheathed with veil remnants .H. mesophaeum group, p. 465
6. Cap small (typically less than 4 cm broad) and white to grayish, often hairy at margin; gills
adnate to decurrent and peeling easily from cap . (see Clitocybe& Allies, p. 148)
6. Not as above . 7
7. Growing on or near corpses, mainly in eastern North America; cap brown to reddish-brown .
.H. syriense
7. Growing in woods, near trees, etc., but not on corpses; cap variously colored (including brown
or reddish-brown); common and widespread . 8
464 CORTINARIACEAE

8. Cap typically brown to cinnamon-brown or dark reddish-brown (but sometimes paler brown
and often shaded with gray or overlaid with a pallid sheen); stalk usually with small protruding
scales.H. sinapizans group & others, p. 465
8. Cap white to buff, pale brown, tan, crust-colored, or yellow-brown; stalk often with a dandruffy
or scurfy apex but not normally with scales . 9
9. Odor sweet .H. sacchariolens (see H. crustuliniforme, below)
9. Odor typically mild or radishlike, etc., but not sweet . 10
10. Cap viscid or slimy when wet; stalk apex usually dandruffy (with small white flakes); gills often
with white edges; odor usually (but not always) radishlike; found in woods or on lawns near
trees . H. crustuliniforme & others, below
10. Not as above; cap only slightly viscid; found in wood chips, gardens, etc. (see Agrocybe, p. 467)

Hebeloma crustuliniforme (Poison Pie)

CAP 3-11 cm broad, convex or broadly convex with an inrolled margin, becoming plane
to obtusely umbonate or with an uplifted margin in age; surface viscid when moist, smooth,
whitish to buff, pale tan, or crust-brown (usually darker toward center and paler at
margin); margin naked. Flesh thick, white; odor distinctly radishlike. GILLS crowded
(often appearing to slightly overlap one another), adnate or notched, pallid when young,
becoming watery brown and finally dull brown; edges white and minutely scalloped, often
beaded with water droplets in wet weather and brown-spotted when dry. ST ALK 4-13 cm
long, 0.5-1.5 (2) cm thick, usually equal except for an enlarged base; solid, fibrillose; white
or tinged cap color; apex powdered or with flakes or granules; base sometimes with white
mycelial threads. VEIL absent. SPORE PRINT brown; spores 9-13 * 5.5-7.5 microns,
elliptical or almond-shaped, smooth or minutely roughened. Cystidia abundant on edges
of gills.
HABITAT: Solitary, scattered, or in groups or troops on ground in woods or at their
edges and on lawns or cemeteries near trees; widely distributed and common. In our area
it is usually abundant from late fall through early spring under pine, and to a lesser extent,
oak. It is by far our most common Hebeloma.
EDIBILITY: Poisonous—it causes mild to severe gastrointestinal distress.
COMMENTS: In spite of my general disdain for Hebelomas, I must admit this is a most
attractive mushroom when fresh. The smooth, viscid, pallid to pie-colored cap, attached

Hebeloma crustuliniforme is a common poisonous species with a whitish to pale tan cap, brown
gills (at least at maturity), and radishlike odor. For photo of young buttons, see p. 466.
1. A “toadstool,” Lactarius atroviridis (see comments under
L. olivaceoumbrinus, p. 70).

2. Lactarius deliciosus, p. 68; edible but not necessarily delicious.

 3. Close-up of the “bleeding” latex
of Lactarius rubrilacteus, p. 68.

4. Lactarius indigo (Indigo Milk Cap), p. 69.

5. Lactarius alnicola (Golden Milk Cap), p. 71, has a

very peppery taste.
6. Lactarius rubrilacteus (also known as L. sanguifluus or the Bleeding Milk
Cap), p. 68.

7. Lactarius torminosus (Bearded Milk Cap), p. 73; common under birch.

8. Lactarius corrugis, an edible eastern milk cap (see
comments under L. volemus, p. 78).

9. Lactarius fallax (Velvety Milk Cap), p. 77.

10. Lactarius fragilis (Candy Cap), p. 80; smells like

maple syrup when cooked or dried.
11. Russula fragrantissima
group, p. 92; can be fragrant
or foul-smelling.

12. Russula cyanoxantha,

p. 94; like many Russulas,
its color is extremely variable.

13. Russula rosacea (Rosy

Russula), p. 99.
14. Russula xerampelina (Shrimp Russula), p. 102; as beautiful as it is delicious!

15. Hygrophorus speciosus, p. 126; a beautiful larch- and pine-loving waxy cap
 ISiP

16. Hygrophorus pudorinus, p. 124, favors spruce.

17. Close-up of the golden-flaked stalk of

Hygrophorus chrysodon, p. 119.

18. Hygrocybe psittacina ( = Hygrophorus

psittacinus), p. 118, is dark green or bright
green when fresh but soon fades.
Ray Gipson

19. Hygrocybe conica (= Hygrophorus conicus

or Witch’s Hat), p. 116, blackens when bruised
or handled.
20. Hygrocybe coccinea (= Hygrophorus coccineus), p. 114.

21. Hygrocybe punicea (= Hygrophorus puniceus), p. 114

 22. Hygrocybe (= Hygrophorus) flavescens (Golden Waxy Cap), p. 115.

24. Omphalina
luteicolor (see
comments under
O. ericetorum, p.
223) grows in
groups on rotting
conifers.
23. Hygrocybe ( = Hygrophorus) calyptraeformis
(Salmon Waxy Cap), p. 117; pointed pink or
salmon-colored cap is distinctive.
 25. Aspens and cottonwoods (p. 42) supply Pleurotus
and Flammulina in the fall and winter, morels in the
spring, Leccinum in the summer.

26. Flammulina velutipes

H
. -
(Velvet Foot), p. 220. jsjfej'.1.'.!
mfifeg
Dan Harper

27. Pleurotus ostreatus

(Oyster Mushroom), p. 134,
growing on bush lupine by
the ocean.
28. Pleurotus ostreatus (Oyster Mushroom), p. 134.
29. Laccaria laccata (Lackluster Laccaria), p. 172.

31. Leucopaxillus amarus (Bitter

Brown Leucopaxillus), p. 168.
32. Clitocybe (= Lepista) nuda (Blewit), p. 153. Some forms have only a slight
lilac tinge; others are perfectly purple.

33. Tricholoma flavovirens (Man on Horseback), p. 179.

34. Clitocybe gibba, p. 157.

36. Marasmiellus candidus,

p. 206.
37. Armillariaponderosa (— Tricholoma magnivelare or White Matsutake), p. 191.

38. Marasmius oreades (Fairy Ring Mushroom), p. 208, a common suburban

lawn mushroom.
39. Close-up of a cluster of
Armillariella ( = Armillaria) mellea
group (Honey Mushroom), p. 196.

41. Omphalotus olivascens (Western Jack-O-Lantern Mushroom), p. 147.

42. Armillariella (= Armillaria) mellea group (Honey Mushroom), p. 196.

43. Armillaria
albolanaripes, p. 194.
 viv..
44. Xeromphalina campanella, 45. Marasmius plicatulus, p. 209;
p. 222; these specimens are one of our most beautiful
somewhat browner than normal. mushrooms.

46. Mycena haematopus (Bleeding Mycena), p. 231.

 47. Marasmius oreades (Fairy Ring Mushroom), p. 208.

48. Mycena murina group (Yet Another Mycena), p. 234, often

fruits by the hundreds under pines.

49. Collybia acervata, p. 215, grows in bundles under conifers.

50. Amanita phalloides (Death Cap), p. 269; don’t expect this deadly species
to be as green and pristine as these perfect specimens!

51. Entoloma madidum, p. 243.

52. Leptonia carnea,

p. 250, is dark blue
and often iridescent.
53. Amanita virosa (Destroying Angel)
(see comments under A. ocreata,
p. 271).

55. Amanita magniverrucata, p. 274, has exaggerated warts on the cap

56. Amanita aspera (also called A. francheti), p. 278; cap color ranges from
bright yellow to dark brown.

CD
Q_
03
X
c
03
O

57. Banana slugs (see p. 29)

prefer mushrooms to almost
anything but other banana
slugs. In this stage they are
commonly called “Poor
Man’s Peaches.”

58. The yellow-capped variety of Amanita

muscaria (Fly Agaric), p. 282.
 JPfarSiiUS 4^1

59. Amanita muscaria (Fly Agaric), p. 282; the red-capped variety.

60. Amanita caesarea group (Caesar’s Amanita),
p. 284, has bright yellow to yellow-orange gills.

61. Amanita calyptrata ( = A. calptroderma or Coccoli, Coccora), p. 284.

 62. Amanita calyptrata,
p. 284; the dark-capped
(fall) form.

63. Amanita calyptrata,

p. 284; the winter-spring
form with a pale yellow to
whitish cap.

64. Amanita velosa (Springtime Amanita), p. 286, often grows

at the edges of meadows when spring wildflowers are in bloom.
65. Amanita pachycolea
(Western Grisette),
p. 290; young
specimens with dark
66. Amanita pachycolea (Western Grisette),
brown caps.
p. 290; mature specimen with a paler cap.

67. Close-up of the glutinous veil in

Limacella illinita, p. 292.

68. Volvariella speciosa, p. 259, is

abundant in gardens and cultivated fields.
69. Lepiota rachodes (Shaggy Parasol), p. 297.

70. Lepiota lutea (= Leucocoprinus luteus),

p. 302, grows in lawns, gardens, flower pots.
71. Agaricus campestris (Meadow Mushroom or Field Mushroom), p. 318.

72. Agaricus californicus, p. 327, a mildly poisonous mushroom often

mistaken for the meadow mushroom.
73. Agaricus cupreobrunneus (Brown Field Mushroom), p. 319.

74. Agaricus fuscofibrillosus, p. 325, is

one of several species that “bleed”
(stain red) when cut.

75. Agaricus subrutilescens (Wine-Colored Agaricus),

p. 326; can be slender (as shown here) or robust.
76. Agaricus praeclaresquamosus ( = A. meleagris), p. 329; poisonous to many
people.

77. Agaricus augustus (The Prince), p. 337; it is also shown on the front cover.

Mr* J
78. Agaricus osecanus group (Giant Horse Mushroom), p. 333, differs
microsopically from the better known A. arvensis.

79. A Greedy Person (see p. 27) laden with giant horse mushrooms
(Agaricus osecanus group, p. 333). Turn to next page to see what happens
to Greedy People.

knhmwm
 80. Crack in the soil caused by
Agaricus bitorquis ( = A.
rodmani), p. 321, a delicious
edible mushroom that often
grows underground. This picture
is dedicated to those purists
who insist that mushrooms must
be photographed exactly as
they occur in nature.

81. What happens to Greedy People (see p. 27

and Color Plate 79).
 82. Agaricus subrufescens
(Almond Mushroom),
p. 336; smells and tastes
like almond extract.

83. Panaeolus solidipes

( = P phalaenarum),
p. 355.

84. A long-stemmed
form of Endoptychum
depressum, p. 730; it looks
like an Agaricus but lacks
gills.

Wr Bn i

v t 3
! ‘ ti
['* '> Jj ■ % '4 ** ✓ \
.Ak f *0 M

85. Coprinus comatus

(Shaggy Mane), p. 345.
 87. Coprinus plicatilis, p. 352; the
cap is oval when younger.

88. Psilocybe cyanescens, p. 371, stains blue when bruised.

89. Stropharia ambigua (Questionable Stropharia), p. 377.

90. Stropharia kauffmanii,

p. 380.

91. Stropharia aeruginosa

(Blue-Green Stropharia),
p. 380.
92. Naematoloma fasciculare (Sulfur Tuft), p. 382.

93. Naematoloma aurantiaca, p. 382.

 94. Naucoria vinicolor,
p. 404, a rare mushroom.

95. Pholiota aurivella group (Golden

Pholiota), p. 390; the caps are very
slimy when wet.

96. Pholiota squarrosa (Scaly Pholiota), p. 389.

 Xm m::
| 1v

97. Pholiota squarrosa (Scaly Pholiota), p. 389.

98. Pholiota squarrosoides (see comments under P squarrosa,

p. 389); youngsters are much scalier than these mature specimens
99. Gymnopilus spectabilis group, p. 410. The bark has been
stripped away to show the base of the cluster; it is not
normal for the caps to be so misshapen.

100. Phaeocollybia californica, p. 415, grows in troops or loose bundles.

rV.

Wan
101. Rozites caperata
(Gypsy Mushroom),
p. 412; note wrinkled
cap.

102. Agrocybe praecox group, p. 469.

103. Close-up of the collapsed

cortina (cobwebby veil) of a
Cortinarius (p. 417); the
rusty-brown color is caused by a
coating of spores.
104. Cortinarius
collinitus group,
p. 431; the cap is
slimy when wet.

105. A young Cortinarius cedretorum (p. 439),

sliced in half to show the cobwebby veil or
cortina.

106. Cortinarius traganus, p. 447, a common conifer-lover.

 mm

&S£jk

107. Little Brown Mushrooms 108. Cortinarius squamulosus,

(“LBM’s”), p. 32, on a wet stump. p. 445; note the bulbous base.

109. Cortinarius violaceus, p. 446, is deep purple to nearly black

110. Close-up of the forked gills of Hygrophoropsis
aurantiaca (False Chanterelle), p. 479.

111. Phylloporus rhodoxanthus (Gilled Bolete), p. 480; the

gills often stain blue or greenish when bruised.

, i. V
112. Gomphidius subroseus (Rosy Gomphidius), p. 483, is
associated with Douglas-fir.

113. Chroogomphus vinicolor (Pine Spike), p. 485; cap

ranges from grayish to orangish to wine-colored or reddish-
brown.

114. Suillus pungens (Pungent

Slippery Jack), p. 503, grows under
pine, often with pine spikes.
 117. Suillus ponderosus
(see comments under
S. caerulescens, p. 496).

118. Suillus luteus

(Slippery Jack),
p. 500.

119. Suillus fuscotomentosus (Poor Man’s

Slippery Jack), p. 504.

120. Close-up of the glandular-dotted stem

of Suillus fuscotomentosus, p. 504.
 121. Suillus subolivaceus (Slippery Jill), p. 499.

122. Suillus cavipes, p. 494, is common under larch; note hollow stem.

123. Fuscoboletinus
ochraceoroseus
(Rosy Larch
Bolete), p. 506,
may be hard to
pronounce but is
easy to see.
124. Suillus lakei, p. 495, is common under Douglas-fir.

126. Fuscoboletinus spectabilis (Bog Bolete) (see key to Fuscoboletinus on p. 506).

 127. Boletus mirabilis, p. 521; a mature
specimen growing on a buried hemlock log.

128. Boletus zelleri, p. 518.

129. Boletus mirabilis, p. 521; this specimen

is younger and darker-capped than #127.

130. Boletus rubripes (Red-Stemmed Bitter Bolete), p. 524;

young specimens.
131. Boletus flav ip or us, p. 522. Pores are brilliant yellow—this
photograph does not do them justice!

132. Boletus dryophilus,

p. 520, is often shorter and
stouter than shown here.

133. Boletus regius (Red-Capped Butter Bolete),

p. 526; unusually small specimens.
134. Boletus appendiculatus (Butter Bolete), p. 525; pores
normally stain blue when bruised.

135. Boletus erythropus, p. 526,

is one of many boletes that stain
blue when cut open.

136. Boletus subvelutipes (see comments

under B. erythropus, p. 526).
 •3
V4:
^

137. Boletus satanas (Satan’s Bolete), p. 527, a poisonous and bulbous oak-lover.

138. Boletus haematinus

(see comments under
B. pulcherrimus,
p. 528).

139. Boletus frostii, p. 528; note the red

cap and netted stalk.
140. Boletus aereus (Queen Bolete), p. 531. Cap is dark brown
beneath a whitish bloom when young, cinnamon-brown or paler
in age.

141. Boletus barrowsii (White King Bolete), p. 529.

142. A basket of Boletus edulis (King Bolete), p. 530.

143. Boletus edulis (King Bolete), p. 530. This red-capped variety is abundant in
the Rocky Mountains in late summer when many of the wildflowers bloom.
144. Boletus edulis (King Bolete), p. 530; the typical brown-capped form.
 145. Leccinum manzanitae
(Manzanita Bolete), p. 539.

146. Leccinum ponderosum (see comments

under L. manzanitae, p. 539).

147. Leccinum insigne (Aspen Bolete), p. 540

 148. Gyroporus castaneus, p. 510,
has a strong, nutty flavor.

150. Tylopilus felleus (see comments under

T. indecisus, p. 535), a bitter-tasting eastern bolete.

149. Tylopilus
gracilis (see
comments under
T. chromapes,
p. 533) is a
common eastern
bolete.

151. Polyporus tuberaster (Stone Fungus),

p. 563, fruits from an underground “tuber.
 ■■ ..

gf a **
1 W&L

152. Fistulina hepatica (Beefsteak Fungus), p. 553, with banana slugs and
chinquapin; this is a mature individual.

153. Phaeolus schweinitzii, p. 570; young, colorful specimens.

154. Laetiporus sulphureus (Sulfur Shelf), p. 572. These
juicy specimens are just emerging.

155. Laetiporus sulphureus,

p. 572. A mature 30-lb.
cluster.
 156. Ganoderma tsugae (see comments under
G. lucidum, p. 577) is one of several varnished
polypores.

157. Hydnellum
zonatum (see comments
under H. scrobiculatum,
p. 627) has a zoned cap
with spines on the
underside.

158. Trametes ( = Coriolus) versicolor

(Turkey Tail), p. 594; zoned caps are
lined underneath with tiny pores.

I f|K Hpf' \ W wk K.; ym fri pftfe |t|

JBL m M r ’ 'iKm 1 v \1 H'l

ft ' W
ft E * E'P’k -I
1 » ft \ f P® w Bf- f 11

w mw ft 1 k ® H
l \ |V: 1: ft»v' * ft
MV
| mm'A IV- ft ft . InE fU |v IjiS *
159. Close-up of the
spines of Hydnum
imbricatum, p. 619.
 161. Dentinum ( = Hydnum)
repandum (Hedgehog
Mushroom), p. 618. In coastal
California it can often be found
when the wild irises are in
bloom.

160. Hydnellum peckii

(Strawberries and Cream),
p. 627, exudes red droplets in
wet weather.

162. Dentinum repandum (Hedgehog Mushroom), p. 618, has spines

underneath the cap instead of gills or pores.
 Herb Saylor
163. Hericium abietis, p. 614, can weigh 50 lbs. or more!

164. Hericium ramosum, p. 615, is more delicate than the above species
165. Hydnum scabrosum group (Bitter Hedgehog), p. 620.

166. Clavariadelphus truncatus, p. 634, is a conifer-loving club fungus.

 168. Ramaria araiospora
(Red Coral Mushroom),
p. 655.

167. Clavaria purpurea

(Purple Fairy Club), p. 637.

169. Clavulina cinerea (Ashy

Coral Mushroom), p. 641.

170. Tremella mesenterica

(Witch’s Butter), p. 673.
171. Clavariadelphus
ligula, p. 633, often
grows in troops
under conifers.

172. Sparassis crispa ( = S.

radicata or Cauliflower
Mushroom), p. 657.

173. Tremella foliacea

(Brown Witch’s Butter),
p. 673.
174. Gomphus floccosus group (Scaly Chanterelle), p. 661; the caps are not
always as brightly colored as these.

175. Cantharellus cibarius (Chanterelle), p. 662. This clean, slender-stemmed,

yellow form is common in eastern North America.
 176. Gomphus clavatus
(Pig’s Ears), p. 661.

111. The gills of this eastern

chanterelle (Cantharellus
cibarius, p. 662) are unusually
thin, deep, and well-developed.

178. Cantharellus cibarius (Chanterelle), p. 662; the large orange form common
on the west coast.
179. Cantharellus subalbidus (White Chanterelle), p. 662, a common western species.

180. Cantharellus infundibuliformis

group (Funnel Chanterelle, Winter
Chanterelle), p. 665, a common
northern species; also called
C. tubaeformis.

181. Cantharellus cinnabarinus

(Red Chanterelle), p. 664, a
common eastern species.
182. Craterellus cornucopioides (Horn of Plenty), p. 666, blends in well with its
surroundings. The caps are black when moist, brown in dry weather.

183. Polyozellus multiplex (Blue Chanterelle), p. 668, a deep blue or violet-tinged

conifer-lover.
184. In coastal California, giant puffballs (Calvatia gigantea
group, p. 682) often grow on poppy-laced hillsides.

185. Pisolithus tinctorius, p. 712; a

young specimen sliced open to
show the numerous spore capsules.
 186. Calvatia booniana
(Western Giant Puffball),
p. 684; these rather small
specimens are much too
old to eat.

188. Calostoma
cinnabarina,
p. 718, a colorful
stalked puffball.

187. Battarrea
phalloides, p. 717,
usually grows in deserts
or sandy soil.

189. Pisolithus tinctorius (Dead

Man’s Foot), p. 712; common in
poor soil, along roads, in patios
and parking lots.
 190. Scleroderma citrinum (Common Earthball), p. 708.

Phil Sharp

191. Dictyophora indusiata (Basket Stinkhorn),

p. 770, is tropical, but a closely related species
occurs in eastern North America.

192. Lysurus mokusin (Lantern Stinkhorn), p. 776.

Like other stinkhorns, this species relies on flies for
spore dispersal.
 194. A stinkhorn “egg” (Phallus impudicus,
p. 768) sliced open to show the olive-colored
spore mass.

193. Phallus impudicus

(Stinkhorn), p. 768;
mature specimen.

195. Mutinus elegans

(Dog Stinkhorn) (see
comments under
M. caninus, p. 771).

196. Mutinus elegans

(see comments under
M. caninus, p. 771);
sometimes called
“Devil’s Dipstick.”
 198. A Clathrus archeri (p.
774) “egg” sliced open to
show the mucilaginous
spore mass.

197. Clathrus archeri (Octopus Stinkhorn),

p. 774; a mature specimen.

199. In Oregon, black morels (Morchella elata group, p. 790) are often found
with calypso orchids.
 200. Neolecta irregularis
(Irregular Earth Tongue),
p. 871.

201. The coastal California version of the white morel (Morchella

deliciosa, p. 789); ridges are whitish when very young.
202. Black morels (Morchella elata group, p. 790) are easily overlooked because
they look like fallen pine cones.

203. Morchella esculenta (Morel), p. 787.

204. Helvella compressa

(Compressed Elfin
Saddle), p. 811; a pale
specimen.
 206. Gyromitra esculenta
(False Morel, Brain
205. Verpa bohemica Mushroom), p. 801.
(Early Morel),
p. 793.

207. Peziza domiciliana, p. 822, a common indoor cup fungus.

210. Sarcoscypha coccinea (Scarlet Cup
Fungus), p. 836.

211. Caloscypha fulgens (Snowbank

Orange Peel Fungus), p. 837, is usually
greenish- or bluish-stained.
212. Truffles, anyone? These are Tuber separans, p. 859.

213. Pachyphloeus citrinus

(Berry Truffle), p. 856.

214. Microglossum viride (Green Earth Tongue), p. 870;

young specimens.
 216. Hypomyces lactifluorum
(Lobster Mushroom), p. 884.

217. “Too many mushrooms.” Overindulgence is a frequent

cause of so-called “mushroom poisoning” (see p. 888).
HEBELOMA 465

(usually notched) brown gills, absence of a veil, and radishlike odor (which is usually quite
pronounced but sometimes slight) are the important fieldmarks. You will undoubtedly
encounter several subtlely different species which more or less fit the above description,
but they can only be differentiated microscopically. Other species: H. hiemale is smaller
(cap about 3 or 4 cm broad), with only a slight radish odor and gills not beaded with drop¬
lets. H. sacchariolens is distinguished by its strong sweet or fruity odor. H. albidulum is
one of several species with a white or whitish cap and little or no odor. H. sarcophyllum
is an unusual but distinctive mushroom with a chalk-white cap that may become reddish-
gray or brownish-tinged in age, deep flesh-colored gills, and reddish-brown spores. It
resembles Pluteus pellitus, but has attached gills and grows on the ground. It could also
be mistaken for an Entoloma, but does not have angular spores. N one of the above species
should be eaten.

Hebeloma sinapizans group (Scaly-Stalked Hebeloma)

CAP 4-13 (20) cm broad, convex to broadly convex with an inrolled margin, becoming
plane or with an uplifted, often wavy margin; surface slightly viscid when moist, smooth,
brown to cinnamon, ochre-brown, pinkish-tan, or dark reddish-brown, but often shaded
with gray or overlaid with a pallid sheen toward margin, which is at first minutely cottony.
Flesh thick, whitish; odor usually distinctly radishlike. GILLS close, usually adnexed or
notched, pallid becoming pale brown, then dull brown or dull cinnamon, the edges
minutely serrated and often beaded with droplets in wet weather and brownish-dotted
when dry. STALK 4-13 cm long, 1 -3 cm thick, usually swollen at base, whitish, with distinct
pallid to brownish flakes or protruding scales; apex usually powdered with small white
granules; firm, solid. VEIL absent. SPORE PRINT dull brown; spores 10-13 * 6-8
microns, elliptical, obscurely roughened. Cystidia present on gill edges.
HABIT AT: Scattered to gregarious, sometimes in rings, on ground under both hardwoods
and conifers or near planted trees on lawns; widely distributed. This species “complex” is
fairly common in our coastal pine forests in the late fall, winter, and early spring. Elsewhere
it often grows with oak and a related species (see comments) is fond of aspen.
EDIBILITY: Poisonous—it causes nausea, vomiting, diarrhea, etc.
COMMENTS: This large, sturdy species or species “complex” is best recognized by its
brown to reddish-brown cap, thick stem adorned with small scales, absence of a veil, and
dull brown, usually notched gills. It is our largest Hebeloma, more robust than H. crustu-
liniforme, and not nearly as attractive. It might be mistaken for a dull colored Cortinarius
were it not for the absence of a cortina even in small buttons. It is sometimes found growing
with Tricholoma imbricatum, which has a brown cap, white to flesh-colored gills, and
white spores. Other species: H. insigne is a robust, scaly-stemmed species that occurs
commonly under aspen and conifers in the mountains of Colorado and New Mexico.

Hebeloma mesophaeum group (Veiled Hebeloma)

CAP 2-6.5 cm broad, convex or umbonate, expanding to broadly umbonate, plane, or
with margin uplifted in age; surface smooth, viscid when moist but soon dry, color variable
but usually dark brown to vinaceous-brown or reddish-brown at the center and paler
(grayish, pinkish, buff, or even whitish) at the margin, often fading in age to dingy brown
or tan and often adorned with thin filmy or fibrillose patches of veil material at or near the
margin, which is inrolled when young. Flesh watery brownish or whitish; odor and taste
typically radishlike (but in some variants mild and in others pungent and/or bitter).
GILLS close, adnate to adnexed or notched, whitish or grayish becoming brown, finally
dull brown; edges usually whitish. STALK 3-8 cm long, 3-8 (10) mm thick, more or less
equal, fibrillose, whitish to dingy brownish, slowly become dark brown from base upward;
Left: Hebeloma crustuliniforme (p. 464). Note brown gills and shape of buttons (bottom). Right:
Hebeloma mesophaeum group. The evanescent veil (not visible here) often leaves fibrils on stalk.

apex often mealy. VEIL cobwebby-fibrillose, white to grayish or buff-colored, thin,

disappearing or forming a slight fibrillose zone on stalk. SPORE PRINT brown; spores
7-11 x 5-7 microns, elliptical, minutely roughened. Cystidia present on gill edges.
HABITAT: Widely scattered to densely gregarious on ground in woods and near trees;
widely distributed. In the northern U nited States this species and its numerous look-alikes
are common under conifers, especially in cool weather (fall, spring). In our area they occur
only sporadically, but I have seen thousands of specimens in a cottonwood-willow
woodland adjacent to a small reservoir, in February and March.
EDIBILITY: To be avoided—several Hebelomas are poisonous and all are difficult to
identify.
COMMENTS: The presence of a cortina (cobwebby veil) in young specimens distin¬
guishes this species and its close relatives from the larger, more prominent members of the
genus such as H. crustuliniforme and H. sinapizans, but leads to confusion with Inocybe
and Cortinarius. The former, however, usually has a silky to fibrillose or scaly cap, while
the latter has a brighter (rusty-brown) spore print and lacks cystidia on the gill edges.
There are dozens of equally drab veiled Hebelomas which are best differentiated from
H. mesophaeum microscopically. (A recent monograph by Alexander Smith, Verna
Stucky Evenson, and Duane Mitchell recognizes nearly 100 veiled species in the western
United States alone!) Among them are: H. strophosum, very similar and widespread,
with a thicker, more persistent veil that forms a woolly sheath on the stem below the fibril¬
lose annulus (ring); and H. fastibile, one of several larger, fleshier species with a reddish-
brown cap with paler inrolled margin and a thicker (1-1.5 cm) stalk.
oo

BOLBITIACEAE spores

TWO microscopic features separate this rather small brown-spored family from the
Cortinariaceae: the cap cuticle is typically cellular (composed of round to pear-shaped
cells) and the spores are smooth and usually furnished with a large germ pore that gives
them a truncate (chopped-off) appearance. People not armed with a microscope are better
off learning the three genera in the Bolbitiaceae (Bolbitius, Agrocybe, and Conocybe)
individually, rather than trying to devise an unwieldy set of ifs, buts, and ands for
distinguishing them as a unit in the field. The Coprinaceae also have a cellular cap cuticle
and spores with a germ pore, but give a much darker(purple-brown to black) spore print.
Like the Coprinaceae, the Bolbitiaceae are mostly frail, saprophytic fungi. They grow in

466
BOLBITIACEAE 467

grass, dung, decaying wood, and humus and are among our most common suburban
mushrooms. In contrast, the bulk of the Cortinariaceae are mycorrhizal sylvan fungi.
Owing to their fragile consistency, the Bolbitiaceae have little food value. The three
common genera are keyed below. If your “LBM” does not key out persuasively, check
Galerina, Tubaria, & Allies (p. 399), and see comments on p. 32

Key to the Bolbitiaceae

1. Cap viscid when moist and conspicuously striate at maturity, at least at margin; fruiting body
soft, sometimes dissolving somewhat in wet weather . Bolbitius, p. 473
1. Not as above (cap may be viscid or striate but generally not both) ... 2
2. Cap typically conical to bell-shaped, at least when young; spore print ochre-brown to bright
rusty to cinnamon-brown . Conocybe, p. 470
2. Spore print dull brown to rich brown (“coffee-brown,” “cigar-brown,” “earth brown,” etc.), or
if brighter than cap convex to plane . 3
3. Gills yellowish to orangish; spore print more or less rusty-orange . . (see Gymnopilus, p. 407)
3. Not as above; spore print dull brown to rich brown .Agrocybe, below

AGROCYBE
Small to medium-sized, saprophytic mushrooms. CAP convex to plane or broadly umbonate,
smooth or cracked but not truly scaly, dry or slightly tacky, rarely striate. GILLS typically attached,
brown to rusty-brown at maturity. STALK central, thick or thin, not markedly fragile (usually
pliant). VEIL present or absent, sometimes forming an annulus (ring) on stalk. VOLVA absent.
SPORE PRINT brown. Spores smooth, usually with a germ pore. Cap cuticle typically cellular.

AGROCYBE is a difficult genus to characterize. The most common species are best
recognized by their smooth or cracked (but not scaly), convex to plane cap, brown spores,
and occurrence in grass, manure, wood chips, or cultivated ground. In some types a
membranous veil is present and in others it is not. Some species resemble Hebeloma and
Pholiota and have been keyed out with those genera (several Agrocybes were originally
placed in Pholiota); others resemble Stropharia, but can be distinguished by their browner
or brighter spore color.
Several Agrocybes are edible, but extreme care must be taken not to confuse them with
the metagrobolizing masses of nondescript brown-spored mushrooms—particularly the
poisonous Hebelomas. The two species described here are among our most common
urban and suburban fungi, but also grow in rural and wooded habitats.

Key to Agrocybe
1. Stalk 5-15 mm thick, with a prominent basal bulb; found in manure, greenhouses, mushroom
farms, etc.(see Conocybe, p. 470)
1. Not as above (but may grow in manure) . 2
2. Partial veil present and membranous or kleenexlike (check young specimens if unsure!), often
(but not always!) forming a ring on stalk or leaving pieces of tissue on cap margin.3
2. Not as above; partial veil absent or rudimentary and evanescent .8
3. Growing on ground in woods; cap dark brown to rusty-brown, often viscid or slimy when moist;
cap margin often striate in age, gills often decurrent A. erebia (see A. praecox group, p. 469)
3. Not as above . 4
4. Cap margin striate when moist; cap usually less than 5 cm broad and usually hygrophanous
(fading markedly as it dries) .(see Galerina, Tubaria, & Allies, p. 399)
4. Not as above . 5
5. Found in swamps and other wet places; cap typically 3 (4) cm broad or less; stalk 2-4 mm thick
.A. paludosa
5. Not as above . 6
6. Growing on hardwoods, often clustered; southern A. aegerita(see A. praecox group, p. 469)
6. Not as above . 7
468 BOLBITIACEAE

7. Cap olive-brown to yellow-brown, tan, creamy, or whitish; veil membranous or kleenexlike(not

fibrillose); stalk lacking scales; cap not normally viscid; found in grassy or cultivated
areas or on wood chips or sometimes in woods .A. praecox group, p. 469
7. Not as above . (seePholiota, p. 384)
8. Fresh cap dark brown to grayish-brown; in groups or clusters on dead hardwoods . A.firma
8. Not as above .*. 9
9. Cap medium-sized (5 cm or more); stalk usually 4 mm thick or more .
.A. sororia{see A. pediades group, below)
9. Cap small; stalk generally less than 5 mm thick . 10
10. Either cap translucent-striate when moist or cap hygrophanous (brown fading to buff or whitish
as it loses moisture) and gills adnate to decurrent . . (see Galerina, Tubaria, & Allies, p. 399)
10. Not as above . 11
11. Spore print rusty-brown or brighter; stalk white and/ or cap 3-6 cm broad; found on dung and
manure; not common .(see Conocybe, p. 470)
11. Not as above; found in grassy or open places, also in dung . 12
12. Cap wrinkled or pitted, white to pale tan; subtropical and tropical (on lawns, etc.) A. retigera
12. Not as above; widespread and common .A. pediades group, below

Agrocybe pediades group (Common Agrocybe)

CAP 1 -3 (4) cm broad, hemispherical (rounded) to convex, or sometimes broadly convex
to plane in age; surface dry or slightly viscid, smooth or sometimes fissured (with cracks)
in age, usually ochre to golden-brown or yellow-brown, but varying to yellowish-buff,
creamy, or even rusty-brown; margin not striate, but sometimes with whitish veil rem¬
nants. Flesh thin, pallid; odor mild or farinaceous. GILLS close, at first adnate but often
seceding; pallid, soon becoming brown to rusty-brown or cinnamon-brown. STALK 2-5
(7) cm long, 1.5-3 (6) mm thick, more or less equal, dry, pallid or buff to yellow-brown (often
paler at apex, darker below); often longitudinally striate. VEIL absent, or if present then
evanescent and fibrillose (not membranous) and either disappearing or leaving slight
remnants on the cap margin and/ or stalk. SPORE PRINT brown; spores 9-13 x 6.5-8
microns, elliptical, smooth, truncate from an apical germ pore.
HABITAT: Scattered to gregarious in grass and cultivated ground, dung or manure, or
in sand; widely distributed and common, flourishing in our area whenever conditions are
conducive. In warm weather it can often be found mingling with Marasmius oreades,

Agrocybe pediades group. A nondescript little agaric common in dung, lawns, and other grassy areas.
AGROCYBE 469

Panaeolus foenisecii, and Conocybe lactea; in cool weather it associates with Psilocybe
coprophila, Stropharia semiglobata, and the Panaeolus campanulatus group.
EDIBILITY: Edible, but not recommended; it is easily confused withpoisonous“LBM’s.”
COMMENTS: The yellowish cap, brown spores, absence of a ring, and small size typify
this commonplace, lawn-loving “LBM” and its close relatives. The cap is not conical as in
Conocybe and Panaeolus, and the spores are neither purple-brown nor purple-black
as in Stropharia, nor white as in Marasmius. Small Hebeloma species are similar, but
generally grow near trees or in the woods and have a filamentous cap cuticle. Other species:
A. pediades var. platysperma has larger spores and frequently has watery spots or streaks
on the cap; A. semiorbicularis (-Naucoria semiorbicularis) is said to have a slightly viscid
cap and broader spores, but is otherwise identical;^. arvalis(-A. tuberosa) is a rare species
that arises from a small, black, easily-overlooked “tuber” (sclerotium); A. amara has a
bitter taste and grows in dense groups in manure or greenhouses; A. sororia is a much larger
species (cap 5-15 cm broad, stalk 0.5-1.2 cm thick) with a tan to tawny cap and bitter taste.
It resembles A.praecox but lacks a veil. I have found it fruiting abundantly on wood chips
in a garden. For another photo of the A. pediades group, see p. 43.

Agrocybepraecox group Color Plate 102

(Spring Agrocybe; Early Agrocybe)
CAP (2) 3-10 cm or more broad, convex or broadly umbonate to plane, or at times withan
uplifted margin in age; surface dry or slightly tacky, smooth or in age sometimes cracked or
fissured; creamy to creamy-ochre, yellow-brown, tan, hazel-brown, or olive-brown;
margin often with veil remnants, not striate. Flesh white or pale ochre, soft; odor mild to
farinaceous; taste mild or more often somewhat bitter to farinaceous. GILLS close, broad,
adnate but often seceding; at first pallid, then light brown to grayish, finally dull brown or
ochre-brown. STALK 3-13 cm long, 0.3-1 or more cm thick, equal or tapered below (or
occasionally enlarged below), white or pallid, but often brownish-stained below in age;
fibrillose-striate, the base usually with white mycelial threads. VEIL membranous but thin;
forming a fragile, superior, skirtlike ring on stalk or leaving remnants on cap margin or
gills or disappearing entirely. SPORE PRINT rich brown (’’cigar-brown”); spores 8-12 x 5-
7 microns, elliptical, smooth, truncate from the apical germ pore. Cap cuticle cellular.
HABITAT: Solitary, scattered, or gregarious in grassy or cultivated areas, along roads, in
wood chips, gardens, woods, etc.; widely distributed, fruiting mainly in the spring. In our
area this species “complex” is among the very common early spring mushrooms mush¬
rooms (February-April), but occurs at other times also. The largest fruitings I’ve seen
were in iceplant along a freeway offramp and on a parkway mulched with wood chips. It
also occurs fairly commonly under mountain conifers in the spring.
EDIBILITY: Edible, but not recommended. The taste is mediocre at best, disgusting at
worst, and there are a number of variants in this group whose edibility is unknown.
COMMENTS: Originally called Pholiola praecox, this attractive fungus can be found
almost anywhere but is especially common in towns and along rural roads. The creamy to
brownish cap, membranous veil (check young specimens!), brown spores, and habitat are
the telltate traits. Both slender and fairly robust forms occur, and both are quite attractive
in their prime. Specimens with a well-formed ring are reminiscent of Agaricus, but the gills
are adnate (at least when young) and never pink or chocolate-brown. Stropharia species
(e.g., Stropharia semiglobata and relatives) are also very similar, but have a darker (purple-
brown to black) spore print and stickier cap. Confusion with Pholiota and Rozites is also
possible, but the habitat and absence of scales or fibrils on the cap and stem should
distinguish it. In my experience, the veil forms a distinct ring on the stalk less than 50% of the
Left: This member of the Agrocybe praecox group is darker and more robust than the typical form
shown in the color plate. It usually grows in wood chips and may well be a distinct species. Right:
Agrocybe erebia (see comments below) has an even darker (brown) cap and slightly decurrent gills.

time, but usually leaves at least some vestiges on the cap margin or gills. The “typical”
form has a creamy cap (see color plate) and ranges from slender to fairly robust. Another
common form (probably a distinct species) favors wood chips, is usually gregarious or
clustered, has a darker (hazel-brown to olive-brown) cap when young, and is fairly robust
(see photo above). Also similar are: A. acericola, with a yellow-brown cap when young
and better-formed ring, growing on decayed wood; and A. dura (formerly Pholiota ver-
miflua), with a whitish to ochre-tinged, often cracked cap and slightly larger spores, found
in cultivated and disturbed ground. Other species with a membranous veil include: A.
erebia, a terrestrial woodland species with a dark brown, often viscid cap and/or slightly
decurrent gills (see above photo); and A. aegerita (-A. cylirtdracea), a medium-sized to
large edible southern species that usually grows in clusters on hardwoods (often living)
such as willow, poplar, and box elder. The latter species is prized in Europe and grown
there commercially. It has a yellowish to grayish-brown to fulvous-tinged, often wrinkled
cap and typically has a well-formed annulus (ring) on the stalk.

CONOCYBE (Cone Heads)

Mostly fragile, little brown mushrooms saprophytic on grass, dung, moss, and humus. CAP some¬
times convex but usually conical or bell-shaped. GILLS attached or free, usually rusty-brown to
cinnamon-brown at maturity. STALK central, usually thin andfragile, often hollow. VEIL present
or absent, sometimes forming an annulus (ring) on stalk. VOLVA absent. SPORE PRINT rusty-
brown to cinnamon-brown to ochraceous. Spores smooth or rough, typically with a germ pore.
Cap cuticle cellular.

CONOCYBES are often called “dunce caps” or “cone heads” because they usually have a
conical or bell-shaped cap. They are mostly fragile, often ephemeral, MycenaAike mush¬
rooms with a slender stem and rusty-brown to ochre-brown spores. They are some¬
times confused with Psilocybe, but have brighter brown spores. Among the brown-
spored mushrooms, they are apt to be confused with Bolbitius, which usually has a dis¬
tinctly viscid, striate cap, and Galerina, which has a filamentous rather than cellular cap
cuticle and an often viscid and/ or translucent-striate cap. One prominent exception to the
above is C. intrusa (keyed below), a rather fleshy mushroom with a thick stalk and convex
cap. It resembles a Hebeloma and was originally classified as a Cortinarius, but micro¬
scopic characteristics plus its rusty-colored spore print relate it to Conocybe.
The “cone heads” are partial to warm weather and fruit in great abundance on watered

470
CONOCYBE 471

lawns. Some, such as C. lactea, are so frail that they shrivel up or topple over a few hours
after appearing. The edibility of most Conocybes is unknown, but they are much too small
to be of food value and at least one species, C.filaris, is dangerously poisonous. Conocybe
is a fairly large genus with over 50 species in North America. As these are largely differ¬
entiated on microscopic characters, only three are described here.
Key to Conocybe
1. Veil present, usually forming a distinct ring (annulus) on stalk . 2
1. Veil absent or evanescent, not forming an annulus .4
2. Growing on dung or manure .C. stercoraria (see C. filaris group, below)
2. Growing in grass, woody chips, moss, etc... 3
3. Cap tawny-brown to orange-brown to brown; annulus often prominent and movable .
. C. filaris group, below
3. Cap reddish-brown to reddish-cinnamon when moist or sometimes yellower, fading as it dries;
annulus not movable, usually small or obscure .... (see Galerina, Tubaria, & Allies, p. 399)
4. Stalk 0.5-1.5 cm thick, usually with a bulbous or thickened base; cap convex to nearly plane,
2.5-9 cm broad, viscid when moist, whitish to tawny to ochre-brown or even reddish-brown;
growing in hothouses, mushroom farms, compost, etc.C. intrusa
4. Not as above . 5
5. Base of stalk aging or slowly bruising blue to bluish-green.
.C. smithii& C. cyanopus{see C. tenera group, p. 472)
5. Not as above . 6
6. Gills crisped, with prominent veins in between; found on lawns C. crispa{see C. lactea, p. 472)
6. Not as above . 7
7. Cap white to buff (or often darker at center), often wrinkled; fruiting body lasting only a few
hours before toppling over or withering ..-.C. lactea, p. 472
7. Cap brown to tan, yellow-brown, or cinnamon, but often fading to buff, more or less smooth 8
8. Cap 3-6 cm broad, broadly conical to bell-shaped or convex; growing in compost in a garden
. C. sp. (unidentified)
8. Cap typically smaller and/or habitat different. 9
9. Cap convex; growing in dung or manure . C. coprophila(see C. tenera group, p. 472)
9. Cap conical to bell-shaped; growing in many habitats . 10
10. Cap translucent-striate when moist, not growing in dung or manure .
.(see Galerina, Tubaria, & Allies, p. 399)
10. Not as above; growing in many habitats .C. tenera group, p. 472

Conocybe filaris group (Ringed Cone Head)

CAP 0.5-2.5 cm broad, conical or bell-shaped, expanding to convex or plane but usually
retaining an umbo; surface smooth, not viscid, tawny-brown to orange-brown or brown;
margin faintly striate when moist. Flesh thin, brown. GILLS pallid soon becoming brown¬
ish to rusty-brown, close, adnexed or notched, sometimes seceding. STALK 1-6 cm long,
1-3 mm thick, more or less equal, fragile, yellow-brown to brown; smooth. VEIL mem¬
branous, brown, forming a prominent but delicate, movable, median to superior ring on
stalk; ring often stained by spores and sometimes falling off. SPORE PRINT cinnamon-
brown to rusty-brown; spores 7.5-13 x 3.5-6.5 microns, elliptical, smooth, with germ pore.
HABITAT: Scattered to gregarious on lawns and other grassy places, also in moss, on
decayed wood and wood chips, etc.; widely distributed, but most common in the Pacific
Northwest. In our area I have found it only once, on a lawn in October, but a large number
of fruiting bodies were present.
EDIBILITY: POISONOUS—potentially fatal! It will not tempt most mushroom hunters
because of its small size, but is dangerous to toddlers in the so-called “grazing stage,”
particularly because it grows on lawns. Analysis has revealed the presence of amanita-
472 BOLBITIACEAE

toxins, and 20-30 caps are equivalent to about half of one cap ofAmanita phalloides—
a compelling reason not to eat “LBM’s!”
COMMENTS: Also known as Pholiotina filaris, this species and its close relatives are
distinguished from other Conocybes by the membranous ring on the stalk. In age, however,
the ring may fall off or be otherwise obliterated. Several poisonous Galerina species have
a ring, but they usually grow on wood and/or have a somewhat viscid, translucent-striate
cap (see G. autumnalis). There are also several Conocybes with an annulus (ring) that grow
on dung and manure (especially of horses), including C. stercoraria, fairly common,
with a yellow-brown to buff-colored cap.

Conocybe teneragroup (Brown Dunce Cap; Common Cone Head)

CAP 1 -2.5 cm broad and tall, conical to bell-shaped, sometimes expanding slightly in age;
surface smooth, not viscid, brown to cinnamon-brown or yellow-brown, usually fading
in age or as it dries to tan, yellowish, or buff; margin finely striate when moist. Flesh thin,
brownish. GILLS adnate to adnexed or free, close, pallid soon becoming pale brown, then
cinnamon-brown or rusty-brown. STALK 4-9 cm long, 1-4 mm thick, equal or with a
swollen base, smooth or minutely mealy, colored like cap or paler, fragile. VEIL absent.
SPORE PRINT rusty-brown or cinnamon-brown; spores 8-14 * 5-7 microns, elliptical,
smooth, with an apical germ pore.
HABITAT: Scattered to densely gregarious on lawns, in gardens, fields, rich soil, dung,
and humus; widely distributed. This species and its close relatives occur practically year-
round in our area in a variety of habitats (including the woods), but are not particularly
numerous. I’ve seen more than 100 fruiting bodies crowded into a small planter box filled
with lettuce seedlings.
EDIBILITY: Unknown, but do not experiment—the similar C. filaris is poisonous!
COMMENTS: The above description will fit a large number of brownish Conocybes with
a cute (but not necessarily acute) conical to bell-shaped cap, thin fragile stem, and no veil
or ring (annulus). They are reminiscent of Mycena and Panaeolus (with which they may
grow), but have rusty-brown gills at maturity and rusty-brown spores. The cap is darker
than that of C. lactea and not as wrinkled, and is not viscid as in Bolbitius. Other species:
C. coprophila has a slightly viscid convex cap and grows in dung or manure; C. smithiiand
C. cyanopus are two rather rare “pupil-dilating” species with a blue to blue-green stem
base, at least in age. The former grows in bogs, the latter in grass or sometimes moss. Neither
should be eaten by “magic mushroom” hunters because of their close resemblance to
ringless representatives of the poisonous C. filaris.

Conocybe lactea (White Dunce Cap; White Cone Head)

CAP 1-2.5 cm broad and/or tall, narrowly to bluntly conical (like a dunce cap), then
bell-shaped or thimble-shaped, the margin usually flaring; surface typically not viscid,
radially wrinkled or striate when moist, white to creamy or tinged buff to pale cinnamon,
especially at the center. Flesh very thin, whitish. GILLS very narrow, adnexed or free,
close, pallid soon becoming cinnamon-brown to tawny-ochre or brown. STALK 3-11 cm
long, 1 -2 mm thick, very thin and fragile, equal or with a small basal bulb, hollow, white or
whitish, often powdery or scurfy above. VEIL absent. SPORE PRINT reddish-brown or
cinnamon; spores 11-16* 6-10 microns, elliptical, smooth, with an apical germ pore.
HABITAT: Scattered or in troops on lawns, baseball fields, and other grassy places; widely
distributed and common in muggy weather. It usually appears in the early morning and
topples over or shrivels up by afternoon; it is one of our common summer lawn mushrooms.
Conocybe lactea is abundant on lawns in warm weather. These are rather mature specimens; younger
ones can have even narrower caps. The fruiting bodies collapse within a few hours.

EDIBILITY: Unknown, but worthless as food—to say it lacks substance is a gross under¬
statement. As Alexander Smith points out, toddlers would be the only ones tempted to
eat it, and if it were poisonous, we would probably know by now.
COMMENTS: This frail mushroom is easily told by its pallid “dunce cap” and thin, fragile
stem. The striate or wrinkled cap is not as viscid as in Bolbitius, and is paler than that of the
C. tenera group. The stem is so feeble that it is practically impossible to bring home speci¬
mens without breaking them. C. crispa is a somewhat similar widespread species with an
ochre-tinged cap and crisped, veined gills.

BOLBITIUS
Small, fragile, rapidly-decaying mushrooms found on dung, humus, grass, or wood. CAP viscid
when moist and usually conspicuously striate at maturity; usually yellow, white, or purple-gray.
GILLS typically adnexed or free at maturity, often dissolving somewhat in wet weather. STALK
slender, fragile, hollow. VEIL and VOLVA absent. SPORE PRINT rusty-brown to ochraceous.
Spores smooth, with a germ pore. Cap cuticle cellular.

THIS small genus of fragile, flaccid, putrescent fungi is reminiscent of Coprinus (particu¬
larly in the striate cap and tendency of the gills to liquefy somewhat), but has rusty-brown to
bright yellow-brown rather than black spores. The viscid striate cap separates Bolbitius
from Conocybe. The cap is rarely brown as in Galerina and the habitat is quite different.
Bolbitius species are worthless as food. The most common types fruit during wet weather
in dung, manure heaps, straw, tall grass, and other vegetable matter; a few grow on wood.
Two species are described here.

Key to Bolbitius
1. Cap bright yellow to pale yellow, at least when fresh .B. vitellinus, p. 474
1. Cap differently colored . 2
2. Usually growing on lawns or dung, rarely in woods . 3
2. Growing in woods or on rotten wood . 6

473
474 BOLBITIACEAE

3. Cap pinkish-grayish-cinnamon in age; on dung.B. coprophilus (see B. vitellinus, below)

3. Not as above; growing on lawns or in grass . 4
4. Cap milky white or with a tinge of buff, especially in age . B. lacteus (see B. vitellinus, below)
4. Not as above . 5
5. Cap conical or oval, very soft, some shade of brown; gills veined and often fused together; stalk
very fragile, often bending over; fruiting body ephemeral; found in hot, humid weather ....
. Gastrocybe lateritia
5. Not as above .(see Galerina, Tubaria, & Allies, p. 399)
6. Cap white; growing on wood or wood chips .B. sordidus
6. Cap not white . 7
7. Cap chocolate-brown to rusty-brown; stalk 3-10 mm thick and brown over lower portion . . .
. (seeAgrocybe, p. 467)
7. Not as above; stalk white or yellow, thin and fragile .B. aleuriatus & others, p. 475

Bolbitius vitellinus (Sunny Side Up)

CAP E5 -7 cm broad when expanded, oval to conical or bell-shaped when young, often
becoming plane in old age; surface smooth, viscid or slimy when moist, bright yellow to
pale yellow (the center sometimes yellow-orange), but often fading in age or as it dries to
whitish, brownish, grayish, etc.; margin striate, at times conspicuously grooved nearly to
the center. Flesh very thin, soft, yellowish. GILLS adnate to adnexed or free, close, soft,
dissolving somewhat in wet weather; pallid or pale yellow to pale brown, becoming rusty-
ochre to cinnamon-brown in age. STALK (3) 5-12 cm long, 2-8 (10) mm thick, equal or
thicker below, hollow, very fragile when thin (readily collapsing), whitish to pale yellow;
often delicately powdered or scurfy. SPORE PRINT rusty-orange to rusty-brown; spores
10-16 x 6-9 microns, elliptical, smooth, truncate from the large apical germ pore.
HABITAT: Solitary, scattered, or gregarious (or even tufted) on dung, manure, straw,
lawns, in tall grass, cultivated ground, etc.; widely distributed and ubiquitous, fruiting
throughout the mushroom season in our area. The most luxuriant fruiting I’ve seen was in
a horse corral where dozens of large, robust specimens were interspersed with massive

Bolbitius vitellinus varies considerably in size and shape, as evidenced by these photographs and
the one on p. 475. However, it is always viscid, yellow (when fresh), and very fragile. Left: Slender,
long-stemmed specimens such as these are common in lawns and other grassy areas. Right: Relatively
robust specimens found on horse dung. In old age the cap often becomes plane (flat).
BOLBITIUS 475

clusters of Peziza vesiculosa (a cup fungus). Greg Wright, who coined the common name,
says that in southern California it commonly occurs on decayed wood.
EDIBILITY: Harmless, but fleshless and flavorless.
COMMENTS: This fragile ephemeral mushroom varies greatly in size, shape, and habitat
but can generally be recognized by its viscid, yellow, striate cap, plus its rust-colored gills
(in age) and soft, fragile texture. The stems often collapse of their own accord, and in wet
weather the cap and gills tend to dissolve, much as in Coprinus (but not as the result of an
autodigesting enzyme). As a rule, those growing ingrass tend to be quite slender and fragile
(see photo), while those growing in a nitrogen-rich environment such as horse manure are
larger and more robust. Other species include: B. coprophilus, with a grayish-pinkish-
cinnamon cap in age, usually found on dung; and B. lacteus, with a small whitish or buff-
tinged cap, found in dung, grass, straw, etc., but not nearly as common as B. vitellinus.

Bolbitius aleuriatus
CAP 15 -4 cm broad, soon broadly convex to broadly umbonate or plane; surface smooth,
viscid when moist, gray to grayish-brown, often with a purplish or lilac tint (center some¬
times darker); sometimes fading in age; conspicuously striate, at least at margin. Flesh
very thin, whitish. GILLS adnexed to free, narrow, close, pallid soon becoming pinkish to
brown or cinnamon-brown. STALK 2.5-7 cm long, 2-3 mm thick, equal or thicker at base,
very fragile, smooth or minutely scurfy or powdery; entirely white or tinged yellow at base.
SPORE PRINT rusty-brown; spores 9-12 * 4-6 microns, elliptical, smooth, with an
apical germ pore.
HABITAT: Solitary or in small groups (rarely more than three) on rotting wood, sawdust,
and humus. It is fairly common in our area in the fall and winter, mainly under oak and
madrone, and probably has a much wider distribution.
EDIBILITY: Unknown.
COMMENTS: Also known as Pluteolus aleuriatus, this species is a fairly frequent but
oft-overlooked fungal feature of our oak wondlands. When the gills are free and pinkish,
confusion with Pluteus is likely. However, its extreme fragility and viscid, striate cap are
good fieldmarks. A form with a yellow stem—possibly distinct—also occurs in our area.
Other woodland species include: B. reticulatus, similarly colored but with a reticulate
(veined or netted) cap; and B. (-Pluteolus) callisteus, with a yellow stalk and a yellow-
olive to rusty-orange cap.

Left: Bolbitius vitellinus, young specimens with rounded yellow caps. (Nancy Burnett) Right:
Bolbitius aleuriatus is a fragile, slender-stemmed woodland species with a grayish to purple-gray cap
that is prominently striate (lined).
 PAXILLACEAE spores

Fleshy medium-sized mushrooms growing on wood or ground. CAP convex to plane or depressed,
margin sometimes strongly inrolled. GILLS typically decurrent, often forked or veined, yellow to
orange or dull-colored; often peeling easily from cap (Paxillus). ST ALK fleshy;, central or off-center
to lateral or even absent. VEIL and VOLV A absent. SPORE PRINT yellowish to brown(Paxillus &
Phylloporus), or white (Hygrophoropsis). Spores mostly elliptical, smooth, without a germ pore,
often dextrinoid.

THIS is a small family of medium-sized mushrooms with decurrent gills and yellowish to
brown (or in Hygrophoropsis, white) spores. The flesh is not dry, white, and brittle as in
Russula, and there is no latex as in Lactarius. The spore print is not rusty-orange as in
Gymnopilus, the stem is much fleshier than in Tubaria, and other brown-spored agarics
without a veil do not normally have decurrent gills.
There are three genera in the Paxillaceae. In the central genus, Paxillus, the margin of the
cap is usually inrolled when young and the gills are often veined or poroid near the stem and
peel rather easily from the cap. Like the dark-spored genera Chroogomphus and Gom-
phidius, Paxillus is thought to be closely allied to the boletes. The second genus, Phyllo¬
porus, includes several species which, despite the presence of gills, are considered true
boletes by many mycologists. The gills are bright yellow and often bruise slightly bluish
like the tubes in many species of Boletus, and the spores are “boletoid” (narrowly
elliptical to spindle-shaped). The third genus, Hygrophoropsis, resembles Paxillus but
has unpigmented (white) spores. It includes the fungus popularly known as the false
chanterelle, and has traditionally been placed in the Tricholomataceae (it is also keyed
out there).
The Paxillaceae are woodland fungi like the boletes, but are not nearly as numerous or
conspicuous. In South America the situation is reversed—Paxillus is a fairly prominent
group while the boletes are few and far between. In case you hadn’t noticed, this is not a
field guide to the fleshy fungi of South America, so only five members of the Paxillaceae
are described. They are not particularly good eating and at least one species, Paxillus
involutus, can be very poisonous.
Key to the Paxillaceae
1. Gills yellow and rather thick, broad, and widely spaced, often staining blue or greenish (some¬
times slowly) when bruised.Phylloporus rhodoxanthus, p. 480
1. Not as above; gills usually close or crowded. 2
2. Stalk velvety from a dense coating of rusty-brown to brown or blackish-brown hairs, typically
1-3 cm thick; growing on or around conifer stumps, logs, etc. Paxillus atrotomentosus, p. 478
2. Not as above; stalk not brown and velvety . 3
3. Odor fragrant (reminiscent of root beer); fruiting body pinkish to whitish .
.Hygrophoropsis olida (see H. aurantiaca, p. 479)
3. Odor mild or at least not fragrant . 4
4. Spore print white or creamy; stalk normally present; gills usually orange or orangish (but some¬
times differently colored) .Hygrophoropsis aurantiaca, p. 479
4. Spore print yellowish to brown; stalk present or absent; gills not orange . 5
5. Stalk lateral to absent; found on wood or lignin-rich humus .Paxilluspanuoides, below
5. Stalk present, central to off-center; usually terrestrial . . . Paxillus involutus & others, p. 477

Paxillus panuoides (Fan Pax)

CAP 1.5-7 (10) cm broad, petal-shaped to mussel- or fan-shaped, attached laterally to
substrate or with a stemlike base; surface minutely hairy or downy becoming smooth, not
viscid; buff to dingy yellowish, yellow-brown, olive-yellow, or dingy ochre; margin often
lobed and at first incurved. Flesh thin, soft, whitish to ochraceous. GILLS radiating from
Paxillus panuoides grows on wood and has little or no stalk. Note how gills are poroid near base of cap.

base of cap, close, pale or dingy yellowish to ochre or pinkish-buff, often crimped and
forked or connected by cross-veins, especially toward base. STALK absent or present
only as a small, narrowed, lateral base. SPORE PRINT yellowish-buff to brown or dingy
ochraceous; spores 4-6 x 3-4 microns, elliptical, smooth, many of them dextrinoid.
HABITAT: Solitary or ingroups or clumps on coniferous logs, stumps, debris, and humus
rich in lignin; widely distributed but not particularly common. In our area I have found it
several times on dead pine in the fall and winter. It is also said to occur on mine timbers,
causing a bright yellow discoloration in its host.
EDIBILITY: Unknown—do not experiment!
COMMENTS: The fan-shaped cap and absence or near absence of a stalk rescue this
listless little brown mushroom from the obscurity it so richly deserves. It might be mis¬
taken for a Crepidotus or Phyllotopsis, but the gills are usually forked or veined. The latter
feature can lead to confusion with the chanterelles, but the growth on wood, dingy color,
and poorly-developed stalk distinguish it. The false chanterelle (Hygrophoropsis
aurantiaca) is also somewhat similar, but has oranger gills, white spores, and a stalk.

Paxillus involutus (Poison Pax; Inrolled Pax)

CAP 4-15 (20) cm broad,atfirst broadly convex withastrongly inrolled margin, then plane
or centrally depressed with the margin eventually unfurled; surface viscid when moist but
otherwise dry; smooth or with soft matted hairs that wear away, sometimes cracked (areo-
late) in age; brown to dingy yellow-brown, olive-brown, or dingy reddish-brown, often
with darker brown stains; margin often slightly velvety or obscurely ribbed. Flesh thick,
firm, pale buff to yellowish, but usually stainingreddishtobrownwhencut. GILLS usually
decurrent, close or crowded, pallid to pale yellowish becoming dingy yellow to olive,
brownish, or yellow-brown, staining dark brown or reddish-brown when bruised or in old

Paxillus involutus is easily told by the strongly inrolled cap margin when young and the tendency
of the entire mushroom (especially the gills and flesh) to stain dark brown. Birch is one of its favorite
tree associates. See p. 41 for another photo, and see p. 478 for close-up of gills.
Left: The close relationship of Paxillus to the boletes is suggested by this close-up of the gills (and
“pores”) of Paxillus involutus. Right: Paxillus atrotomentosus is a wood-inhabiting species with
decurrent gills and a dark velvety stalk.

age; often forking and/or forming pores near the stalk. STALK 2-7 (10) cm long, 0.5-4
cm thick, usually shorter than width of mature cap, equal or tapered at either end, central
to somewhat off-center, solid, firm, dry, smooth; colored like cap or paler, often with dingy
reddish to dark brown stains. SPORE PRINT brown to yellow-brown; spores 7-10 x 4-6
microns, elliptical, smooth.
HABITAT: Usually scattered to densely gregarious on ground in woods, around the edges
of bogs, and in tree-studded parks or lawns; widely distributed, and by far the most
common member of the genus. Two possibly distinct forms occur in our area: the typical
one (shown in photo at bottom of p. 477) fruits mainly with planted birch trees in the fall,
while a larger form occurs with oak and pine in the late fall and winter.

EDIBILITY: Dangerous! It is often eaten in Europe and by transplanted Europeans in

America, but it can cause hemolysis (destruction of red blood cells) and kidney failure if
eaten raw, and sometimes even when thoroughly cooked! (Apparently the human body can
develop a devastating sensitivity to it.) It often has a sour taste anyway, so it is best avoided
and should never be eaten raw. The one recorded instance of a bonafide mycologist dying
of mushroom-poisoning was attributed to this fungus!
COMMENTS: The dingy brownish cap with inrolled margin when young plus the decur¬
rent gills that stain brown and are usually forked or veined near the stem are the
unappealing features of this unappealing fungus. Actually, its symmetry and compactness
are quite intriguing, but its choice of color is downright disastrous—an unbecoming blend
of dingy browns and murky yellows. The stature is reminiscent of a Lactarius or Russula,
but the gills do not exude a latex when broken and the flesh is not chalky or exceptionally
brittle. Other species: P. vernalis is similar but larger, with a paler cap when young and
reddish-brown to chocolate-brown spores. It grows mainly with aspen.

Paxillus atrotomentosus (Velvet Pax)

CAP 4-15 (20) cm broad, convex becoming plane or centrally depressed; surface dry,
unpolished, velvety or with matted hairs, yellow-brown to rusty-brown to dingy reddish-
brown becoming dull brown, dark brown, or blackish-brown in age; margin at first in-
rolled. Flesh thick, rather firm or tough, pallid to ochre or buff. GILLS close or crowded,
usually decurrent, tan to dull ochre, dingy yellowish, or paler; often forked or veined near

478
PAXILLUS 479

stalk. ST ALK 2-9 (12) cm long, 1 -3 (5) cm thick, often short and usually off-center or even
lateral; solid, tough, densely velvety from a coating of brown to dark brown or blackish-
brown, often matted hairs; apex often paler or yellowish. SPORE PRINT yellowish to
brownish; spores 5-6.5 * 3-4.5 microns, elliptical, smooth, many of them dextrinoid.
HABITAT: Solitary or in groups or tufts on oraround conifers(usually dead) or madrone;
northern in distribution. It is fairly common in the Pacific Northwest in the late summer
and fall, but rare in our area. It causes a carbonizing decay (brown rot) in its host.

EDIBILITY: Not recommended. Some people apparently collect it for the table, but the
same can be said of P. involutus, which has caused several deaths! (See comments on
edibility of that species.)
COMMENTS: The combination of brown cap, dark brown velvety stem, decurrent gills,
and growth on rotting conifers makes this one of the Pacific Northwest’s most distinctive
agarics. It does not discolor as much as P. involutus, and is quite attractive when fresh.

Hygrophoropsis aurantiaca (False Chanterelle) Color Plate 110

CAP 2-8 (14) cm broad, convex becoming plane or somewhat depressed, the margin at
first inrolled; surface dry, often somewhat felty or velvety, typically some shade of dark
orange, brownish-orange, brownish-yellow, yellowish-brown, olive-brown, or dark
brown (often darker or browner at center and orange to yellowish-orange at margin),
but in one form whitish and in another blackish. Flesh thin, pallid or tinged orange or cap
color; odor mild. GILLS decurrent, close, fairly thin and narrowat maturity (may be blunt
when young); usually forked dichotomously; typically deep orange to bright orange, but
sometimes pale orange or in one form yellowish. STALK 2-10 cm long, (0.2) 0.5-1 (2) cm
thick, central or off-center, equal or enlarged toward base, often curved; dry, yellowish
to orange or brownish-orange or colored more or less like cap. SPORE PRINT white to
creamy; spores 5-8 * 2.5-4.5 microns, elliptical, smooth, often dextrinoid.
HABITAT: Solitary, scattered, or in groups or tufts in humus and on rotting wood,
usually under conifers; widely distributed. In our area it fruits from the fall through early
spring, but seems most abundant in cool, dry weather when there are few other fleshy fungi
out and about. I usually find it under pine or redwood.
EDIBILITY: To be avoided. In my experience it is edible—but far from incredible. Some
sources, however, list it as mildly poisonous (whether or not this is a result of confusion
with Omphalotus, as has been suggested, is unclear).

Hygrophoropsis aurantiaca. The cap is usually brown at the center and lighter or brighter toward
the margin, but in some forms it is completely brown (or even black) and in others it is whitish. For
a close-up of the forked gills, see color plate.
480 PAXILLACEAE

COMMENTS: The typically bright orange, decurrent, dichotomously forked gills (see
color plate) and white spore print are the principal fieldmarks of this attractive but variable
fungus. In spite of its common name, it is difficult to confuse with the true chanterelle
(Cantharellus cibarius) if the following is kept in mind: the gills are thinner, crowded, and
bladelike at maturity (but often blunter when young), and are usually oranger than those
of the chanterelle. In other words, the false chanterelle has “true” gills while the true
chanterelle has “false” gills. Also, Hygrophoropsis has flimsier flesh, a browner cap, is less
robust, differently shaped (not as wavy or frilled) and sometimes grows on rotten wood.
The jack-o-lantern mushrooms (Omphalotus species) are also similar, but have brightly
colored flesh, typically grow in clusters on or under hardwoods, and do not have forked
gills. The false chanterelle was originally placed in Cantharellus, and is listed in many
mushroom books as Clitocybe aurantiaca, but the forked gills, frequently off-center stalk,
and dextrinoid spores connote a closer kinship to Paxillus. Other species: H. olida (-Cli¬
tocybe morganii) is a small (cap 1 -4 cm) species with a pinkish cap and stalk (that may fade
to buff), whitish to pinkish gills, and a flagrantly fragrant odor that is reminiscent of root
beer or cinnamon candy. At least some of its gills are forked and/or have cross-veins, its
stalk can be central or off-center, and its spores are white and dextrinoid. Like H. auran¬
tiaca, it favors conifers and is widely distributed, but seems to be rather rare. I have seen
it only once in California, near Mount Shasta, in June.

Phylloporus rhodoxanthus (Gilled Bolete) Color Plate 111

CAP 2-5 (12) cm broad, broadly convex to plane or with uplifted margin in age; surface
dry, minutely velvety to nearly smooth, dull brown to olive-brown or yellow-brown in one
form, red to reddish-brown in another; sometimes developing pallid to yellowish cracks or
fissures. Flesh thick, pallid to yellowish. GILLS widely spaced, broad, fairly thick, adnate
to decurrent but sometimes seceding; sometimes forked or with cross-veins; bright yellow
to ochre, typically bruising green or blue (but often slowly, sometimes not at all), some¬
times also staining brownish. STALK 3-10 cm long, 0.4-1 (1.5) cm thick, equal or tapered
below, dry, solid, yellow to dingy yellowish-buff or reddish, often stained dingy brown or
reddish-brown below; smooth. SPORE PRINT brown to yellowish- or olive-tinged;
spores 9-15 * 3-6 microns, narrowly elliptical to spindle-shaped, smooth.
HABITAT: Solitary to widely scattered or in small groups (usually twos or threes) on
ground in woods, widely distributed. The brown-capped form can be found nearly every
year in our area, but seldom in quantity. Like the related Boletus subtomentosus, it fruits
throughout the mushroom season, often along roads and trails.
EDIBILITY: Edible, but rarely found in sufficient numbers to bemorethanjustacurious-
ity. Some“bolete-your-meat’ers” rate it highly, but I’ve found it to be slimy and insipid.
COMMENTS: This evolutionary oddity is an adamant nonconformist. Thedry, minutely
velvety brownish to reddish cap is an almost exact replica of Boletus subtomentosus and
relatives, yet the underside of the cap has gills! These are thicker than the gills of many
agarics, and are bright yellow and often bruise blue like the tubes of many boletes (it may
take several minutes, however, for the color change to show). What’s more, the spores
are “boletoid”—long, narrow, and spindle-shaped—causing some taxonomists to place it
in the Boletaceae. Thus Phylloporus, like many other mushrooms, has been denied a stable
family and is at the complete mercy of the “authorities.” Like baseball’s itinerant tobacco-
chewing utility infielders of southern rural origin, its group affiliation is in a state of con¬
stant flux as it is systematically shuttled back and forth between one “team” and another.
Other species: P. arenicola is a very similar species with an olive-brown to olive-yellow cap
and adnexed (not decurrent) gills which don’t blue when bruised. It was originally de¬
scribed from coastal Oregon under pine, but I have not seen it.
 481

GOMPHIDIACEAE spores

THIS is a small but prominent family of fleshy-stemmed mushrooms with decurrent gills
and smoky-olive to black spores. No other group of black-spored mushrooms has con¬
sistently decurrent gills. Gomphos, meaning “peg” or “stake,” refers to the shape of the
young mushrooms, which look like tent stakes with their long stems and small, rounded to
conical caps. The spores are typically “boletoid”—i.e., long, narrow, and more or less
spindle-shaped, and the Gomphidiaceae are thought to be more closely related to the
boletes than to other gilled fungi. They are mycorrhizal exclusively with conifers and often
occur with the bolete genus Suillus. This may be purely coincidental, since Suillus is also
mycorrhizal with conifers, or it may be that the fungal mycelia are in some way associated
with each other as well as with their mutual host.
As far as is known the family is completely safe from an edibility standpoint. There are
two common genera, Gomphidius and Chroogomphus, but the latter is listed under
Gomphidius in many older books. A closely related gastroid genus, Brauniellula (see
p. 732) also occurs, but it often grows underground and does not have true gills.

Key to the Gomphidiaceae

1. Immature gills and flesh in the cap white to grayish . Gomphidius, below
1. Immature gills and flesh in the cap some shade of yellow, orange, buff, salmon, reddish, etc. .
.Chroogomphus, p. 484

GOMPHIDIUS
Fairly small to medium-large terrestrial mushrooms associated with conifers. CAP viscid or slimy
when moist, usually smooth. Flesh white to grayish. GILLS decurrent, soft or somewhat waxy,
fairly close to well-spaced, pallid or white when young but blackening in age. STALK fleshy, more
or less central; lower portion or base usually bright yellow( especially within). VEIL usually present,
but often disappearing or leaving only a slight ring (annulus) on stalk. VOLVA absent. SPORE
PRINT smoky-gray to black. Spores narrowly elliptical, smooth. Cap tissue not amyloid.

THIS genus is readily recognized by its soft, somewhat waxy decurrent gills, slimy-viscid
cap, white or pallid flesh, and smoky-black spores. In addition, the base or lower portion of
the stem is brilliant yellow in most species—a very striking and telltale feature. Gomphidius
is most likely to be confused with Chroogomphus, which has orange to yellowish or pinkish
flesh, and the waxy caps(Hygrophoraceae), which have white spores.
Gomphidius species are mycorrhizal exclusively with conifers—particularly fir,
spruce, Douglas-fir, hemlock, and larch—but in pine forests they are largely supplanted by
Chroogomphus. November rains can mean Gomphidius galore in our area, with large
numbers of the glutinous fruiting bodies littering our Douglas-fir forests, along with
maggoty Suillus species and assorted Russulas.
Gomphidius species are edible but not often collected—perhaps because they are soft,
sluglike, insipid, and putrescent. Also, they do not dry nicely like Chroogomphus, and
often blacken when cooked—but some people relish them nonetheless (maybe they can’t
find anything better!). About ten species occur in North America, all of them rather similar
in appearance. Two are described here and several others are keyed out.

Key to Gomphidius
1. Slimy veil present in young specimens; stalk usually viscid when moist (from veil remains);
associated with various conifers . 2
1. Veil absent; stalk not viscid; associated primarily (but not exclusively) with larch or tamarack;
found in northern North America . 5
482 GOMPHIDIACEAE

2. Cap pink to rosy-red or red, not large (up to 7 cm broad); stalk usually 1.5 cm thick or less . .
. G. subroseus, p. 483
2. Cap sometimes pinkish or salmon, but usually dingier or darker in color (whitish, grayish,
brownish, vinaceous, purplish, etc.); small to large . 3
3. Stalk with little or no yellow at base; fruiting body medium-sized to fairly small .
.G. smithii(see G. subroseus, p. 483)
3. Base or lower portion of stalk typically bright yellow; medium-sized to fairly large . 4
4. Often growing in small clumps, stalk often rooted deeply in soil; associated primarily with
Douglas-fir; spores typically less than 14 microns long. G. oregonensis, below
4. Not as above; found with various conifers (including Douglas-fir); spores longer than 14
microns; widely distributed .G. glutinosus & others (see G. oregonensis, below)
5. Cap whitish to yellowish when young; found in eastern North America; rare . . . G. nigricans
5. Cap pale cinnamon to brownish, etc., when young; widely distributed in northern latitudes
. G. maculatus {see G. oregonensis, below)

Gomphidius oregonensis (Insidious Gomphidius)

CAP 2-15 (18) cm broad, at first peglike, then broadly convex to plane or depressed; sur¬
face viscid or slimy when moist, smooth, color variable: whitish to salmon-buff to
ochraceous-salmon to dull pinkish when young, becoming dingier (brownish to purplish-
or vinaceous-gray to dark reddish-brown) in age, often spotted or stained smoky-gray to
black. Flesh rather soft, white or grayish (or tinged cap color under cuticle), but brilliant
yellow in lower part of stalk or at base. GILLS decurrent, soft and rather waxy, close to
fairly well-spaced, white or pallid, then gray and finally blackish as spores ripen. ST ALK 5-
15 cm long, 1 -5 cm thick, equal or tapered below (or occasionally swollen), solid, rather firm
or even tough, dry and white above the veil, whitish or dingy below and viscid when wet,
sometimes with darker streaks; lower portion bright yellow. VEIL whitish and fibrillose
beneath a layer of slime; disappearing or forming a slight hairy-slimy superior ring on stalk
which is subsequently blackened by falling spores. SPORE PRINT smoky-gray to black;
spores 10-14 (16) * 4.5-8 microns, spindle-shaped to narrowly elliptical, smooth.
HABITAT: Solitary, scattered, or gregarious on ground under Douglas-fir and other
conifers, often in small clumps which originate deep in the soil and may include one or more
aborted fruiting bodies; known only from western North America and very common,
(along with G. glutinosus—see comments) on the Pacific Coast. In our area it grows with
Douglas-fir, often mingling with Suillus species (S. caerulescens, S. lakei, S. ponderosus),
from the fall through early spring.
EDIBILITY: Edible, but its sliminess makes it undesirable—except as a possible escargot
substitute. To clean the cap, simply peel off the slimy pellicle (skin).

COMMENTS: The intensely yellow flesh in the lower stem plus the whitish flesh elsewhere,
viscid-slimy cap, soft decurrent gills, and smoky-black spores immediately identify this
mushroom as a Gomphidius. However, distinguishing it from its brethren (see below),
especially when it is not growing in clumps, can be difficult unless the spores—the shortest
in the genus—are measured. In age the fruiting body is often quite murky and insidious-
looking, hence its common name. The cap is quite variable in color but is usually dingier
and dirtier than that of G. subroseus, and the stalk is usually thicker—but not necessarily
slicker. Other species: The “Glutinous Gomphidius,” G. glutinosus, is probably the most
common and widespread member of the genus. It is mycorrhizal with a variety of
conifers, including spruce, fir, and in our area, Douglas-fir. It is quite similar to G. oregon¬
ensis, but has longer spores (over 14 microns), does not often grow in clumps, and usually
has a darker cap when young (purplish to purple-brown, brownish-gray, purplish-gray,
etc.). Another western species, G. largus, is essentially a giant version of G. glutinosus, and
also grows with spruce and fir, mainly at higher elevations. Finally, there is the “Hideous
Gomphidius,” G. maculatus, which has a pale cinnamon to reddish-brown to murky brown
Gomphidius oregonensis, a common associate of Douglas-fir. Gills are decurrent and darken in old
age, a veil is present when young (visible in specimen on right), and cap is slimy when moist. Specimen
at lower left has been sliced open to show the flesh, which is white in the cap and brightyellowinlower
part of stalk. Buttons (top left) of this species are often found in tight clumps.

or blackish-spotted cap and dark-fibered stalk. It lacks the slimy veil characteristic of
other Gomphidii and features both robust and very slender forms. It is a northern species
that grows mainly with larch, often in the company of Suillus cavipes and S. grevillei.

Gomphidius subroseus (Rosy Gomphidius) Color Plate 112

CAP 2.5 -7.5 cm broad, at first peglike, then broadly convex becoming plane or broadly
depressed; surface viscid or slimy when moist, smooth, dull to bright pink to rosy-red or
even red, often spotted grayish in old age. Flesh white (or tinged pink under cuticle), but
yellow in base of stalk and sometimes pinkish in extreme base. GILLS typically decurrent,
soft and rather waxy, fairly well-spaced, white becoming pale gray to smoky-gray or even
blackish as spores ripen. STALK 3-7.5 cm long, 0.5-1.5 cm thick, equal or narrowed
slightly at base, dry and white above the veil, usually viscid below and white to dingy-
colored with a pale yellow to bright yellow base; smooth or fibrillose, solid. VEIL white
and fibrillose beneath a layer of slime; disappearing or forming an obscure hairy-slimy
superior ring on stalk which is blackened by spores. SPORE PRINT smoky-gray to black;
spores (11) 15-21 * 4.5-7 microns, spindle-shaped to narrowly elliptical, smooth.
HABITAT: Widely scattered to gregarious or occasionally tufted on ground under
conifers (especially Douglas-fir), northern North America. It can be found throughout the
range of Douglas-fir and is fairly common in our area in the fall and winter. It is also said to
occur with spruce, fir, and hemlock.
EDIBILITY: Edible, but rather bland; peel off the slimy pellicle (skin) before cooking it.
COMMENTS: The beautiful rosy-red to pink cap and modest size separate this fungus
from other Gomphidius species, and make it the most attractive member of its clan. It
typically shows less yellow in the stem than either G. oregonensis or G. glutinosus, but more
than G. smithii{see below). It is often mistaken for a waxy cap(Hygrophoraceae) because
of its brightly colored cap and soft waxy gills. H owever, the smoky-gray to black spore print
distinguishes it. Other species: G. roseus has a redder cap and longer spores, but has not yet
been recorded in North America; G. smithiican also be rather small, but shows little or no
yellow in the base of the stalk and has a grayish to vinaceous-gray or brownish cap. The
latter occurs with Douglas-fir in coastal California but grows with other conifers as well.
484 GOMPHIDIACEAE

CHROOGOMPHUS (Pine Spikes)

Medium-sized terrestrial mushrooms associated with conifers. CAP convex to conical, umbonate,
plane, or depressed; smooth or woolly, viscid or dry. Flesh colored (pale orange to yellow-orange,
buff, salmon, pinkish, reddish, etc.). GILLS usually decurrent, fairly well-spaced, dull orange to
yellow-orange or ochraceous when young, blackening in age. STALK fleshy, more or less central,
dry. VEIL present but usually disappearing, fibrillose. VOLVA absent. SPORE PRINT olive to
smoky-gray to black. Spores long and narrow (narrowly elliptical to spindle-shaped). Gills with
prominent cystidia. Cap tissue amyloid.

A FLAGRANT fungal feature of our coastal pine forests, Chroogomphus (crow-oh-

gom-fus) is easily recognized by its decurrent gills, ochraceous to orange, salmon, buff-
colored, or reddish flesh, and smoky-olive to black spores. The fruiting bodies, in contrast
to Gomphidius, are very attractive: brightly colored with a lean, clean look and a ten¬
dency to become reddish or wine-colored in old age. The stem and gills are typically yellow-
orange to orange to orange-buff, but the latter blacken as the spores ripen.
Chroogomphus species grow exclusively with pine in our area and usually mingle with
slippery jacks (Suillus pungens, S. fuscotomentosus, etc.). They have an unusually long
growing season—from the onset of the fall rains through the spring—and may even turn
up in the summertime. They often seem quite abundant when other fleshy fungi are scarce.
They do not decay as rapidly as their cousins the Gomphidii and are usually free of
maggots. However, they are susceptible to attack by a greenish mold.
Chroogomphus is an excellent genus for beginners because all of its species are edible
and highly distinctive. True, fresh specimens are slimy and insipid when cooked (not to
mention purple), but they acquire a pleasant, chewy texture that is perfect for tomato
sauces if they are chopped up finely and then dried. At least that is the gospel according to
“Saint Ciro,” a wondrously demented and exceptionally resourceful toadstool-tester of
Sicilian extraction. With a large family to feed and an immense forest of unpicked Chroo¬
gomphus across the street, he isn’t the least bit deterred by their lack of renown. Every year
he gathers trunkloads and dries them for summer use! About ten species of Chroogomphus
occur in North America; most of these are keyed out here. Dr. Frank Dutra of Los Gatos,
California, reports that eating them can turn one’s urine red (just like beets!). This pheno¬
menon is harmless, but can cause anxiety if one mistakes the red pigment for blood.

Chroogomphus vinicolor is a common sight in pine forests and on lawns where pines have been
planted. Note the shape, which is very characteristic.
CHROOGOMPHUS 485

Key to Chroogomphus
1. Fruiting body robust (stalk at least 2 cm thick); cap and/ or stalk usually decorated with orange
to reddish to wine-colored fibrillose or felty patches or zones; cap convex (not umbonate),
not viscid or only very slightly so; spore print often olive-tinged . C. pseudovinicolor, p. 486
1. Fruiting body not so robust, or if so then not as above . 2
2. Cap not viscid or only slightly so, covered with flattened woolly or felty fibrils or fibrillose
scales; often associated with conifers other than pine (but also with pine) . 3
2. Cap usually smooth and viscid when moist, often lustrous when dry (but sometimes minutely
scaly or faintly fibrillose); associated principally with pine . 4
3. Cap pale to dull orange, yellow-orange, or ochraceous .C. tomentosus, p. 487
3. Cap usually grayish, at least at margin .C. leptocystis (see C. tomentosus, p. 487)
4. Cap small (1-4 cm broad) and usually pinkish- or vinaceous-tinged when young; restricted to
eastern North America; rare . C.flavipes
4. Not as above; widespread and common . 5
5. Cap often brightly colored (buff, yellow-orange, orange, ochre) until fairly mature; cystidia
thin-walled . C. ochraceus(see C. vinicolor, below)
5. Not as above (but cap may be orangish when very young); cystidia thin- or thick-walled .. 6
6. Cap often vinaceous (wine-colored) in old age; cystidia thick-walled .... C. vinicolor, below
6. Cap usually dull reddish-brown in age; cystidia thin-walled C. rutilus (see C. vinicolor, below)

Chroogomphus vinicolor (Pine Spike) Color Plate 113

CAP (1)2-10(13) cm broad, peglike to nearly conical or convex when young, becoming
somewhat top-shaped to plane or even shallowly depressed (but often with a slight umbo);
surface smooth to finely fibrillose (or sometimes scaly in age), viscid when wet, often
lustrous or silky when dry; color variable: dull orange to ochraceous to grayish, brownish,
yellow-brown, olive-brown, bister, or reddish-brown, often wine-red to dark vinaceous-
brown in age and tending to developing wine-red stains where rotten or when dried; mar¬
gin at first incurved. Flesh thick, usually pale orange but varying to buff, salmon, or
ochraceous-buff (often somewhat yellower in lower portion of stalk); often becoming wine-
red in old age or where injured. GILLS broad, fairly well-spaced, decurrent or occasionally
adnate, pale to dull orange or ochraceous when young, then clouded gray with spores and
finally blackish in old age. STALK typically long, slender, and sometimes sinuous, but at
other times short and rather thick; 2.5-15 (20) cm long, 0.5-2 (6) cm thick; equal or more
often tapered downward; dry, solid, firm; pale orange to yellow-orange to orange-buff,
ochraceous, or reddish-tinted when young, often more vinaceous-red in age and/ or with
reddish fibrils. VEIL dry, fibrillose, ochraceous to orange, soon disappearing or leaving a
slight hairy ring on stalk. SPORE PRINT smoky-gray to smoky-black, sometimes with an
olive tinge; spores (14) 17-23 * 4.5-7.5 microns, more or less spindle-shaped, smooth.
Cystidia on gills large, thick-walled.
HABIT AT: S olitary to scattered or gregarious on ground under conifers, particularly pine;
widely distributed, but especially common in northern and westernNorth America. Along
with C. rutilus (see comments), it is a prominent fungal facet of our pine forests and is also
frequent in yards and lawns where pines have been planted. The major crop is usually in
the late fall or winter and is often accompanied by masses of mushy slippery jacks (Suillus
species), but it can be found most any time. It is quite sporadic in its fruiting habits—
overwhelmingly abundant some years, very sparse during others.
EDIBILITY: Edible, but like all pine spikes, better dried than fresh (see comments on
p. 484). It is usually very clean, and unlike Suillus, is shunned by maggots.
COMMENTS: Anyone who hunts our coastal pine forests will come across this common
mushroom and its equally edible look-alike, C. rutilus (see next page). The variable color
or mixture of colors (orange, gray, brown, and/or wine-red) plus the pale orange flesh,
Chroogomphus rutilus is easily confused with C. vinicolor in the field. The critical difference is
microscopic (see coments below).

decurrent gills, and smoky-black spores make it as distinctive as it is attractive. The fruiting
body is often quite slender and long-stemmed, but robust forms also occur. C. rutilus
(formerly Gomphidius viscidus) is a very similar, widespread species with thin-walled
rather than thick-walled cystidia. It can often be told in the field by its slightly duller or less
vinaceous color and broader or flatter cap, but both species are so variable in shape and
color that a microscopic examination is often necessary to positively distinguish them.
Both differ from C. tomentosus and C. pseudovinicolor in their smoother cap that is viscid
or slimy when wet and often shiny when dry. Other smooth, viscid-capped species: C.
ochraceus, widely distributed, is closely related to C. rutilus (i.e., it has thin-walled cysti¬
dia), but is smaller, with a yellow-orange to buff, ochre, orange, or grayish cap, at least
when young; it also occurs in our area with pine. See also C.flavipes (in the key).

Chroogomphus pseudovinicolor (Robust Pine Spike)

CAP 5-15 cm broad, convex, sometimes becoming plane in age; surface dry to very slightly
viscid, orange-buff to dull orange to ochraceous-orange, sometimes flushed red in places or
often mottled with reddish or orangish patches of felty or woolly material, and often be¬
coming entirely dark dull red in old age; margin often fringed with veil remnants at first.
Flesh thick, quite firm, pale to dull orange or orange-buff (but often brighter or yellower
in base of stalk), reddening in old age or around maggot tunnels. GILLS fairly well-spaced,
decurrent or sometimes adnate, often forked; pale orange to dingy ochraceous, then
clouded olive to gray or black by spores. STALK 6-12 cm long, 2-5 cm thick at (or near)
apex; usually tapered below, solid, firm, dull orange to orange-buff or ochraceous, usually
with zones or patches of reddish to wine-colored fibrils or woolly-felty material; often
dingier, darker, or redder overall in age. VEIL scanty, fibrillose, soon disappearing or
leaving a slight zone of hairs at thickest part of stalk (just below apex). SPORE PRINT
greenish to smoky-olive to blackish (occasionally lacking olive tinge); spores 15-20 * 5-7.5
microns, spindle-shaped to narrowly elliptical, smooth. Cystidia on gills thick-walled.
HABITAT: Solitary, scattered, or in groups (often in small clumps) on ground under
conifers; known only from western North America, not common. In our area it seems to
occur only with ponderosa pine—usually in the fall and early winter.
EDIBILITY: Edible. It is firmer and meatier than other Chroogomphus species, but
unfortunately, not as numerous.

486
Left: Chroogomphus pseudovinicolor, a robust orange to reddish species with patches of fibrils on
the cap and stalk. Right: Chroogomphus tomentosus has a dry fibrillose, orangish to ochraceous
cap. It is very common under northern conifers.

COMMENTS: This robust but relatively rare Chroogomphus is a very beautiful mush¬
room when fresh—easily distinguished from other Chroogomphus species by its dry,
convex (never conical or umbonate) cap and thick, woolly-scaly stalk. Also, the spore
print is usually greener than that of its brethren, and it often grows in small clumps of
2-4 individuals rather than in the scattered fashion typical of other pine spikes.

Chroogomphus tomentosus (Woolly Pine Spike)

CAP 2-9 cm broad, peglike becoming broadly conical to convex, umbonate, or plane;
surface dry to very slightly viscid, covered with flattened woolly or felty fibrils or fibrillose
scales; pale buffy-orange to pale or bright ochraceous to ochraceous-orange, the fibrils
sometimes vinaceous-tinged; sometimes purple-stained in age. Flesh yellow-orange to dull
orange or pale orange-buff. GILLS well-spaced, usually decurrent but sometimes adnate,
yellow-orange to ochraceous or colored like cap, becoming smoky-gray to smoky-brown
as spores mature. STALK 4-18 cm long, (0.3) 0.7-1.5 (2) cm thick; equal or more often
narrowed below; solid, dry and somewhat fibrillose, colored more or less like cap. VEIL
dry, fibrillose, scanty, colored like cap; disappearing or leaving slight hairy remnants on
stalk near apex. SPORE PRINT smoky-gray to blackish; spores 15-25 * 6-9 microns,
narrowly elliptical to spindle-shaped, smooth. Cystidia on gills with fairly thick walls.
HABITAT: Solitary to widely scattered or gregarious onground underconifers(hemlock,
fir, Douglas-fir, pine); common in the mixed coniferous forests of the Pacific Northwest
and northern Rocky Mountains from late summer through early winter; also common at
times in northern California. It does not seem to occur in our area, but has been found on
the University of California campus in Los Angeles.
EDIBILITY: Edible, but better dried than fresh (see comments on p. 484).

COMMENTS: The dry, woolly-fibrillose cap, overall dull orange to ochraceous color,
decurrent gills, smoky-black spores, and growth with conifers form a distinctive set of
characteristics. The cap is drier and woollier than that of C. vinicolor and C. rutilus, and
not as variable in color, while the stalk is not as thick as that of C. pseudovinicolor. Also, it
seems to favor mixed coniferous forests, whereas the others grow almost exclusively with
pine. Other species: C. leptocystis of western North America tends to have a grayer cap
and thin-walled cystidia, but is otherwise quite similar.

487
488

Boletes
BOLETACEAE
BOLETES have a spongelike layer of tubes on the underside of the cap instead of gills.
Otherwise they resemble agarics: medium-sized to large, mostly terrestrial fungi with a
cap and central stalk. Polypores also possess a tube layer, but are tough or leathery and
usually grow on wood. The few polypores that are fleshy and terrestrial typically have an
off-center stalk and tubes which do not peel easily from the cap.
The tube layer in the boletes, on the other hand, usually peels away cleanly from the cap.
The spores are produced on basidia which line the inner surfaces of the tubes, which are
vertically arranged so that the spores, when discharged, drop into the air. The mouths
of the tubes are known as pores. They are sometimes stuffed with a pith when young,
making the pore surface appear smooth. In some species the pores are radially arranged
(arranged in rows which radiate from the stalk), giving a somewhat gill-like effect. This
is carried to an extreme in Phylloporus rhodoxanthus (p. 480), a “bolete” with true gills,
which serves to illustrate why boletes are thought to have more in common with gilled
mushrooms than with, say, the coral fungi, teeth fungi, or polypores.
Boletes used to be lumped together in one giant and unwieldy genus, Boletus. Several
genera are now recognized, the most prominent being Suillus, Leccinum, Tylopilus, and
Boletus. The common term “bolete,” however, is still applicable to any member of the
Boletaceae, not just those still retained in Boletus.
The salient characters for identification are much the same as for agarics. It is particu¬
larly important, however, to note color changes on the pore surface, cap, stalk, and flesh.
Many boletes will stain blue or greenish-blue when bruised (see Color Plate 135); a few will
stain brown or some other color, while others will not stain at all. In a number of boletes
the cap is typically areolate, i.e., it develops an extensive system of shallow cracks or
fissures as it matures, exposing the flesh beneath.
Another important feature to note is the type of stalk ornamentation, if any. In Boletus
and Tylopilus, the upper portion or sometimes the entire stalk is frequently reticulate
or “netted”: covered with a network of veins. Some species are coarsely reticulate (see
Color Plate 139), others are very finely so (see photo on p. 489). In Leccinum, the stalk is

Close-up of the pores (tube mouths) in Boletus subtomentosus. Boletes are one of two groups of
fungi that produce their spores in tubes. The other group is the polypores (p. 549).
Close-up of the finely reticulate (netted) stalk of a young Boletus edulis.

always scabrous: decorated with tufted hairs or rough, scurfy scales (scabers) which
typically darken at maturity (see Color Plates 145-147). In Suillus, on the other hand, the
stalk is often speckled or smeared with glandular dots (see Color Plate 120). These pinkish
to brown or blackish spots exude a resinous substance that will stain your fingers brown.
Spore color ranges from yellow to olive, brown, reddish-brown, chocolate-brown, or
black. A spore print is easily obtained, but pigmentation in the tubes may stain the paper,
making the spore color look brighter (usually yellower or greener) than it actually is. The
spore shape is characteristically “boletoid”: long and narrowly elliptical or spindle-shaped
in face view, inequilateral in profile. Viewed through the microscope, a bolete spore is an
unusually large, handsome, and healthy-looking specimen, somewhat reminiscent of a
surfer with a good tan.
The boletes are one of the safest groups for the table, as wellas one of the most substantial
and rewarding. A few members of the genus Boletus are poisonous, but the majority are
edible (though some are distinctly better than others). Since they are large and fleshy,
boletes are a popular picnicground for maggots. These may enter in the usual manner, via
the stem, or surreptitiously, through the tubes. Always check for maggots in the field, and
cut away soggy or infested areas (assuming you already know the species) before popping
them in your basket. That way you are less likely to have a maggot-infested mush two days
later. Some people peel off the tubes—when old and spongy they are mostly wateranyway.
Also, boletes are vulnerable to attack by a variety of moldy-looking parasites. Discard
all such individuals when picking for the table. For more on hunting boletes, see p. 546.
The boletes attain their greatest diversity in the deciduous forests of eastern North
America, but are also a conspicuous and colorful component of the West’s woodland
fungi. The cap can be very hefty and the pores are often brightly colored. The vast maj ority
are mycorrhizal. Several hundred species occur in North America, more than 80 on the
west coast. Though not complete (what book is?) and more than a trifle expensive (what
book isn’t?), Harry Thiers’ California Mushrooms: A Field Guide to the Boletes is a
definitive must for the serious and self-respecting bolete buff. (Thiers hasalso co-authored,
with Alexander Smith, the bolete bible for easterners, entitled The Boletes of Michigan.)

Key to the Boletaceae

1. Fruiting body often appearing aborted or misshapen and often buried or partially buried in the
humus; tubes at least somewhat disoriented (i.e., not necessarily arranged vertically); spore
print unobtainable; not common.Gastroboletus, p. 544
1. Not as above; tubes vertically arranged; spore print usually obtainable; usually found above
the ground when mature. 2

489
490 BOLETACEAE

2. Cap and stalk shaggy or coarsely scaly, the scales gray to brown or black; pores whitish becoming
gray or black in age; veil present when young; spore print dark brown to black; spores spiny,
warty, or reticulate; found in eastern North America and Southwest . Strobilomyces, p. 543
2. Not as above . 3
3. Stalk roughened by small tufted hairs or scales (scabers) which are usually gray to dark brown
or black by maturity (but differently colored or remaining pallid in a few species); stalk not
glandular-dotted and typically not reticulate; pores rarely yellow .Leccinum, p. 536
3. Not as above; stalk typically without scabers ..4
4. Bright yellow, dry, cottony-powdery or cobwebby veil present when young, usually leaving
cottony remnants on cap and/or stalk; pores and flesh usually blueing (sometimes slowly)
when bruised .Pulveroboletus, p. 509
4. Veil absent, or if present then not as above . 5
5. Veil present when young, sometimes forming a ring (annulus) on stalk, at other times clinging
to the margin of cap, and in still other cases disappearing in age .9
5. Veil absent (check young specimens if unsure), but cap sometimes fringed with sterile tissue 6
6. Stalk with glandular dots or smears at least when mature (the dots often slightly resinous to the
touch); stalk usually more or less equal or at least not markedly bulbous; cap often (but not
always) viscid; spore print olive to brown or dull cinnamon .Suillus, p. 491
6. Stalk not glandular-dotted (or very rarely so, but then with much darker spores).7
7. Pores often radially elongated, sinuous, arranged in rows, and/or veined; tubes often (but not
always) decurrent; taste of cap not peppery .8
7. Pores not as above, or if so then taste peppery .10
8. Veil absent when young; tube layer often difficult to separate from cap; stalk often short,
curved, and/or off-center; spores broadly elliptical to nearly round; associated with hard¬
woods (mainly ash and alder); common in eastern North America, rare in the West .... 14
8. Not with above features; veil present or absent; spores elliptical to spindle-shaped; associated
with conifers or less commonly hardwoods; widespread and common .9
9. Spore print dark grayish-brown to dark reddish-brown, chocolate-brown, or vinaceous-brown;
veil typically present (at least when young) and glandular dots typically absent(rarely present);
associated with larch and other northern conifers .Fuscoboletinus, p. 505
9. Spore print olive-brown to brown, yellowish, or dull cinnamon; veil present or absent; stalk
glandular-dotted or not; found with many kinds of trees (including larch) .. Suillus, p. 491
10. Cap viscid to very slimy when moist; stalk glandular-dotted or if not then stalk typically white
to yellow and not reticulate; associated with conifers .Suillus, p. 491
10. Not with above features . 11
11. Spore print flesh-colored to pinkish-brown, reddish-brown, vinaceous, or chocolate-brown;
pores typically white to pinkish, vinaceous, gray, or brown but not red and only rarely yellow
(pores if pallid often but not always staining brown when bruised); common in eastern North
America but rather infrequent in the West .Tylopilus, p. 532
11. Not as above; spore print yellow to yellow-brown, olive-brown, olive, brown, or more rarely
cinnamon-brown; pores white, yellow, olive, red, orange, or sometimes gray or brown (but if
pallid then not typically staining brown when bruised) . 12
12. Spore print pale yellow to yellow; stalk often hollow or partially hollow at maturity (especially
near the base); common in eastern North America, rare in the West.Gyroporus, p. 510
12. Spore print olive-brown to olive or brown, or occasionally yellow-brown or cinnamon-brown;
stalk not normally hollow; common and widespread . 13
13. Spores ornamented with ridges, grooves, pits, etc.; stalk usually relatively long(7 cm or more)
and slender and coarsely reticulate or conspicuously lined with jagged ridges for most of its
length; pores yellow to greenish-yellow, not usually blueing when bruised; typically terrestrial;
found in eastern North America and the Southwest .... Boletellus& Austroboletus, p. 508
13. Not with above combination of features; spores typically smooth; widespread Boletus, p. 511
14. Common under hardwoods (especially ash) in eastern North America; cap olive to dingy yellow-
brown or brown; flesh not blueing or blueing only slightly; stalk often quite dark in old age,
off-center or lateral .Boletinellus(-Gyrodon) merulioides
14. Associated with alder; known from southern California (and Europe), apparerltly rare; cap
yellowish to reddish-brown; flesh usually blueing when bruised . Gyrodon lividus
 491

SUILLUS (Slippery Jacks)

Medium-sized to fairly large, fleshy, terrestrial bo\eits associated almost exclusively with conifers.
CAP usually viscid or slimy, but sometimes dry and scaly. PORES and tubes typically white to
yellow, sometimes radially arranged, rarely bruising blue. STALK typically more or less equal
(rarely bulbous), rarely reticulate; often with resinous brown to pinkish glandular dots and smears.
VEIL frequently present and forming an annulus (ring) on stalk, but often absent. SPORE PRINT
brown (usually olive-brown becoming dull cinnamon-brown as moisture escapes). Spores typically
elliptical to spindle-shaped, smooth. Bundles of cystidia on inner surfaces of tubes staining dark
brown to black in potassium hydroxide (KOH).

MOST slippery jacks have a veil or a glandular-dotted stem (see Color Plate 120) or both.
In addition, the cap is often “slippery” (viscid or slimy), the pores are radially elongated
in some species (that is, in rows radiating from the stalk toward the cap margin), the
stalk is neither conspicuously bulbous nor reticulate as is commonly the case in Boletus,
and they are mycorrhizal almost exclusively with conifers. Suillus is a genus of exceptions,
however: several species are not slippery, others lack a veil, still others are not glandular-
dotted, many do not have radially-arranged pores, and two or three species in eastern
North America occur with hardwoods. However, by using the above features in combi¬
nation, you should have little trouble distinguishing Suillus from other bolete genera with
the exception of Fuscoboletinus, which has darker spores and doesn’t occur in California.
Like other boletes, Suillus can be found in older mushroom books under Boletus. It is
perhaps the most primitive genus in the Boletaceae, leading to Boletus on the one hand and
to the Gomphidiaceae (via Fuscoboletinus) on the other. Suillus splits neatly into two
groups. In the first (the genus Boletinus of some authors), a veil is always present and the
cap is frequently dry and fibrillose, thus belying the moniker“slippery jack.” Also, the stalk
is not glandular-dotted and the pores are often elongated and/ or radially arranged. In our
area this group occurs only with Douglas-fir, where it is often accompanied by species of
Gomphidius. Elsewhere it also occurs with larch and various pines. The second group
includes the “true” slippery jacks—or “slippery jills” as they are sometimes called. They
have a slimy cap or glandular-dotted stem or both, and a veil may or may not be present.
They occur principally with pines, often mingling with species of Chroogomphus.
Slippery jacks are remarkable for their fecundity. Under favorable conditions they
fruit in quantities that must be seen to be believed. Bushels can be harvested from a single
row of pine trees and truckloads from a pine plantation. They are indisputably among
the most prominent fungal facets of temperate coniferous forests, and in fact, wherever
there are conifers of the pine family (Pinaceae)—be it in a forest or park, on a lawn or by a
freeway—there are bound to be one or more Suillus species also. They are partial to cool
weather and generally fruit later than Boletus and Tylopilus species, which like it warmer.
In our area they can be found most any time but peak in the late fall and winter.
None of the slippery jacks is known to be poisonous, but a few have caused “allergic”
reactions. In my experience even the so-called “choice” species are insipid and slimy when
cooked, but one novel solution is to take advantage of their inherent wetness by using
them as an escargot substitute. (You can boil them, flavor them with herbs, stuff them into
empty snail shells, and call it “Parsley, Sage, Rosemary, and Slime.”) Slippery jacks also
tend to be wormy and do not dry well in the tasty tradition of Boletus edulis. They are
indisputably available, however, so I heartily recommend that you try them. If you find a
species that pleases you, you’ll never have to worry about a mushroom shortage again!
Nearly half of the 70+ North American species of Suillus have been found in California.
As they pose a threat to those who don’t watch where they step (they are at least as
dangerous as banana peels!) and inevitably arouse the curiosity of fungophiles and other
weird elements (for their prodigious numbers as well as their slipperiness). I’m offering a
generous selection here. Fourteen species are described and many others are keyed out.
492 BOLETACEAE

Key to Suillus
l. Veil present (check young specimens if possible!), either completely covering the pores when
young (and often forming an annulus or ring on stalk after rupturing) or present only as a roll
of cottony tissue on cap margin or patches on cap (and not forming an annulus) .2
1. Veil and annulus absent; margin of cap naked, even when young .28
2. Glandular dots or smears present on stalk and conspicuous at least in age; associated mainly
with pine, sometimes with spruce . 11
2. Stalk lacking glandular dots (or occasionally with a few visible in old age); found with Douglas-
fir, larch, hemlock, or sometimes pine in the West and with larch, pine, and oak in East . . 3
3. Stalk usually hollow or partially hollow, at least at base; cap not viscid; associated only with
larch .S. cavipes, p. 494
3. Not as above; stalk not normally hollow .4
4. Cap with tawny to orangish, red, reddish-brown, pink, or purple-red scales or fibrils, dry or with
a viscid layer beneath the scales (i.e., can be viscid in wet weather if fibrils are obliterated) 5
4. Cap more or less smooth (but sometimes streaked) and viscid to slimy when moist . 7
5. Found in eastern North America, mainly with pine .43
5. Found in western North America, principally with Douglas-fir ..6
6. Cap fibrillose or fibrillose-scaly, the fibrils reddish-brown to brick-red or pinkish, or sometimes
orange-buff or tawny; surface dry, or viscid only in wet weather or old age . . S. lakei, p. 495
6. Cap with only scattered fibrils, but often streaked; surface usually viscid when moist, not reddish
but often cinnamon-brown, orangish, greenish, etc.S. caerulescens & others, p. 496
7. Associated with hardwoods (e.g., oak) in Great Fakes region; veil thick, tough, often gelatinous;
spore print olive-yellow to yellow-brown; spores more or less round .N. sphaerosporus
7. Not as above . 8
8. Base or lower portion of stalk often (but not always) staining blue or green when cut (often
staining slowly or weakly); associated primarily with Douglas-fir. 9
8. Not as above; stalk not blueing and/or associated with larch or pine . 10
9. Veil viscid and yellow to bright orange while still covering the pores; cap more or less smooth
.S. ponderosus & others (see V. caerulescens, p. 496)
9. Veil dry and whitish while covering the pores; cap smooth or fibrillose S. caerulescens, p. 496
10. Cap yellow to golden-yellow to dark red or bay-red; flesh usually yellow when young; associated
with larch .S. grevillei, p. 497
10. Cap duller (honey-colored to dingy yellow-brown, brown, etc.); flesh white becoming pale
yellow in age; associated primarily with pine . 11
11. Cap at first covered with a whitish veil that usually leaves small patches or scales on the surface
of cap, especially near margin; found mostly with white (5-needle) pines in northern Rocky
Mountains (particularly common in northern Idaho).S. albivelatus
11. Not as above . 12
12. Pores large (many of them typically 1 mm or more broad at maturity), often (but not always!)
elongated and at least somewhat radially arranged (in radiating rows). 13
12. Pores mostly less than 1 mm broad and not radially arranged . 15
13. Veil typically forming a gelatinous annulus(ring) on stalk; cap often umbonate, dingy-colored,
and rather small; found mainly with lodgepole and beach pines .S. umbonatus, p. 498
13. Not as above; veil not forming a gelatinous annulus or forming only a very slight one .... 14
14. Stalk often short, poorly developed, and/or off-center; pores often elongated and irregular;
associated with pines and other conifers in the Sierra Nevada and other mountains.
. S. riparius& S. megaporinus(see S. sibiricus, p. 498)
14. Not as above; stalk typically well-developed; found with 5-needle (white) pines; widespread 45
15. Veil usually forming a distinct annulus (ring) on stalk . 16
15. Veil typically attached only to margin of cap or if attached to stalk when young then typically
forming only a slight or very obscure annulus . 20
16. Annulus usually thin and/or fibrillose; glandular dots on stalk often inconspicuous until old
age .S. pseudobrevipes, p. 500
16. Annulus usually well-formed; stalk with conspicuous glandular dots (unless very young) . 17
SUILLUS 493

17. Stalk and/ or underside of veil (annulus) often with purplish or dull lilac shades; cap reddish-
brown to dark reddish-brown or sometimes yellow-brown .S. luteus, p. 500
17. Not as above; purplish shades absent; cap white to dingy yellowish, olive-brown, etc.18
18. Found in eastern North America (including the Gulf Coast) . 19
18. Found in the West (especially common in Pacific Northwest) .S. subolivaceus, p. 499
19. Annulus tall (i.e., broad) and baggy or bandlike (both lower and upper edges often flaring
out) .S', subluteus & others (see S. subolivaceus, p. 499)
19. N ot as above; annulus not baggy; stalk slender; cap whitish to pale yellow, pale cinnamon, etc.,
the slime strongly acidic-tasting.S. acidus
20. Veil purplish to lilac-brown on its underside; cap soon brown to dark brown; associated mainly
with western white pine .S. borealis (see S. pseudobrevipes, p. 500)
20. Not as above .21
21. Cap white becoming olive-gray or olive, then spotted, streaked, or becoming entirely yellow,
cinnamon, orange, etc.; common with various pines in central and southern California, and in
regions where Monterey pine has been planted .S. pungens, p. 503
21. Not as above; cap not olive or grayish-olive when young .22
22. Veil rudimentary (not well-developed); cap yellow to dingy yellowish, mustard-yellow, or
orange-buff, often with brown to reddish spots, streaks, or scales or with well-developed brown
to grayish fibrils; known only from eastern North America and the Southwest.34
22. N ot as above; if similarly colored then veil well-developed at least when young(asa roll of cottony
tissue extending from cap margin toward stalk, or as a true partial veil covering the pores) 23
23. Cap yellow to dingy olive-yellow or mustard-yellow (even in age), often with reddish to brown
scales, spots, or streaks; stalk markedly glandular-dotted; found with 5-needle pines ... 45
23. Not with above features . 24
24. Young specimens with yellowish to pale tan caps and very dense, conspicuous glandular dots on
stalk .S. glandulosipes (see S. pseudobrevipes, p. 500)
24. Not as above; young specimens differently colored or with inconspicuous glandular dots . 25
25. Slime on cap becoming chocolate-brown in age; associated mainly with sugar pine on the west
coast, lodgepole pine in Rocky Mountains . S', brunnescens(see S. pseudobrevipes, p. 500)
25. Not as above .26
26. Growing in volcanic soil, often buried; cap yellow to tawny, often fibrillose or streaked and
often developing pinkish to reddish-brown areas S. volcanalis(see S. pseudobrevipes, p. 500)
26. Not as above; habitat different . 27
27. Veil material present only on margin of cap, never attached to stalk; cap whitish to buff‘or at
times brown to tawny or cinnamon .S. albidipes(see S. pseudobrevipes, p. 500)
27. Veil attached to the stalk when young; cap dingy yellowish to honey-colored or brown, but
not whitish .S. pseudobrevipes, p. 500
28. Pores large (many of them 1-3 mm in widest dimension), often elongated radially or arranged
in radiating rows and often decurrent . 29
28. Not as above . 30
29. Cap hairy or velvety and brown to vinaceous-brown; found under hardwoods in southeastern
United States .S. castanellus
29. Not as above .46
30. Cap dry to viscid but not thickly slimy, with hairs, fibrils, or scattered fibrillose scales when
young (but these often absent in age or rainy weather); pores typically dark brown to yellow-
brown, pale cinnamon, or orangish when young (but often dingy yellowish in age) .31
30. Cap viscid to very slimy when moist, smooth (bald) or streaked; fibrils typically absent on cap
(occasionally present, but if so then pores typically white to pale yellow when young) ... 35
31. Flesh and/ or pores staining blue when bruised or cut (but sometimes staining slowly or only
slightly—check several specimens if unsure!) .S. tomentosus, p. 504
31. Not as above . 32
32. Found in eastern North America and the Southwest; cap sometimes yellow or orangish . . 33
32. Found in California and the Pacific Northwest; cap not yellow . S. fuscotomentosus, p. 504
33. Pores dark brown when young; cap soon more or less bald; associated with conifers, often in
boggy areas; odor often fragrant when several specimens are put together . ... S. punctipes
33. Not as above; cap usually with scales or colored fibrils; found with hardwoods and conifers 34
494 BOLETACEAE

34. Associated with conifers (mainly pine); cap scales or fibrils usually conspicuous and glandular
dots on stalk prominent at maturity .S. hirtellus
34. Favoring hardwoods; scales on cap usually small and glandular dots obscure . S’, subaureus
35. Glandular dots or smears present on stalk and usually conspicuous by maturity . 36
35. Glandular dots absent or obscure (in old age sometimes visible at apex) .40
36. Cap white when young, whitish to pale yellowish in age (or with darker slime); associated with
white pines in eastern North America; common .S. placidus(see S. granulatus, p. 502)
36. Not as above; cap darker, at least in age, and/or found in the West .37
37. Cap smooth, at first white but soon becoming olive to olive-gray and then orange, cinnamon,
yellow, brown, etc. (or splashed with these colors); found with various pines in coastal central
and southern California, or in regions where Monterey pine is planted . . S. pungens, p. 503
37. Not with above features; cap variously colored but lacking an olive or olive-gray phase . . 38
38. Cap with grayish to dark brown streaks, fibrils, or scales; found with pine (especially Monterey
pine) in California; taste harsh, unpleasant . ... S. acerbus(see S. fuscotomentosus, p. 504)
38. Not as above . 39
39. Cap buff to yellowish or pale cinnamon at maturity; common with ponderosa pine in the
Southwest .S. kaibabensis(see S', granulatus, p. 502)
39. Not as above; either found elsewhere or cap darker at maturity S’, granulatus & others, p. 502
40. Taste peppery (chew on a small piece of cap) and base of stalk usually bright yellow or pores
brilliant yellow to intense greenish-yellow and associated with hardwoods (see Boletus, p.511)
40. Not as above .41
41. Stalk typically white to yellow (not brown or red) and not viscid or slimy; associated with
conifers (mainly pine); pores not blueing when bruised .42
41. Stalk viscid or slimy and cap sometimes yellow, or if not then not as above (see Boletus, p. 511)
42. Cap dark brown to reddish-brown to dull cinnamon when young .S. brevipes, p. 501
42. Cap whitish or at least paler than above when young .
.S. pallidiceps & S. occidentals (see 5. brevipes, p. 501)
43. Pores and flesh typically blueing when bruised; spores ridged or otherwise ornamented .
.(see Boletellus& Austroboletus, p. 508)
43. Pores and flesh not blueing; spores smooth .44
44. Cap fibrils pink to reddish to purple-red; growing with white pine S.pictus(see S', lakei, p. 495)
44. Not as above; cap dull yellowish to orangish or pinkish-orange; especially common along the
Gulf Coast .S. decipiens (see S. lakei, p. 495)
45. Veil sometimes forming a slight annulus (ring) on stalk; base of stalk often vinaceous-stained;
common in western North America, including the Southwest .S. sibiricus, p. 498
45. Not as above; veil typically not forming an annulus; common in eastern North America and
also found in the Southwest .S. americanus (see S. sibiricus, p. 498)
46. Cap small (up to 5 cm broad) and stalk slender .S. helenae (see S. umbonatus, p.498)
46. Cap medium-sized to large; stalk usually quite thick S. punctatipes(see S. granulatus, p. 502)

Suillus cavipes (Hollow-Foot) Color Plate 122

CAP 3-12 cm broad, convex or broadly umbonate becoming plane or slightly uplifted;
surface dry (not viscid), densely hairy-fibrillose, often with a suede-like or felty texture;
typically dark brown to reddish-brown to orange-brown or tawny (rarely paler), the
margin and/ or tips of the fibrils often paler. Flesh firm, white or pale yellow, not blueing
when bruised. PORES arranged more or less radially, large, angular (elongated); pale
yellow becoming dark yellow or greenish-yellow, not blueing; tubes same color, usually
decurrent. STALK4-9 cm long,0.5-2 cm thick, equal or swollen below,dry, lemon-yellow
above the ring, colored like cap or paler below; glandular dots absent; lower portion hollow
inside, at least in age; not blueing when cut. VEIL fibrillose-cottony, white, forminga thin,
fragile, fibrillose ring on stalk and/ or leaving remnants on cap margin. SPORE PRINT
dark olive to brown; spores 7-10 * 3.5-4 microns, elliptical to spindle-shaped, smooth.
SUILLUS 495

HABITAT: Scattered to gregarious on ground in late summer and fall, associated with
larch trees; common throughout the northern hemisphere wherever larch occurs. I have
seen it in Oregon and Idaho, but not in California.
EDIBILITY: Edible and “choice” according to some sources, but I find it bland. Its one
saving grace is that it isn’t as slimy as many slippery jacks.
COMMENTS: Also known as Boletinus cavipes, this is the only Suillus with a consistently
hollow stem and dry, densely hairy, brown to tawny cap. Its growth with larch is also diag¬
nostic. Fresh specimens have an unusually clean, pristine appearance that is readily notice¬
able but difficult to describe (see color plate); the pale pores contrast strikingly with the
darker cap, making it one of our most beautiful boletes. Since larch does not occur in
California, S. cavipes probably doesn’t either. However, it is a prominent feature of the
coniferous forests of Idaho, Montana, and the Cascades, as well as the northeastern
United States and Canada. For other larch-loving boletes, see S. grevillei and the genus
Fuscoboletinus (particularly F. ochraceoroseus and F. aeruginascens).

Suillus lakei (Western Painted Suillus) Color Plate 124

CAP 5-15 cm broad, convex becoming plane or shallowly depressed; surface covered
with reddish-brown to brick-red to pinkish or tawny fibrils or fibrillose scales on a yellow
to dingy orange or tan background; dry, but in one form viscid in wet weather beneath the
fibrils; margin sometimes hung with veil remnants. Flesh thick, yellow, often discoloring
pinkish when bruised. PORES large (1-3 mm in length), sometimes radially arranged,
yellow when young becoming dingy yellow or ochre and usually discoloring reddish to
dingy reddish-brown where bruised or in age; tubes same color as pores, adnate to de¬
current. STALK 3-8 (12) cm long, 1-3 (4) cm thick, more or less equal, firm, dry, solid;
yellow above the ring, usually with reddish to brown streaks below; glandular dots absent;
interior usually staining weakly blue or green when cut, at least near base. VEIL white or
becoming cap color, dry, usually forming a fibrillose ring or ragged zone on stalk, but
sometimes disappearing. SPORE PRINT brown to dull cinnamon; spores 8-11 * 3^4
microns, spindle-shaped to elliptical, smooth.
HABITAT: Scattered to gregarious on ground in woods; associated with Douglas-fir
and common throughout the West where its host occurs. In the Rocky Mountains it fruits
in the summer and early fall and is the most common bolete associated with Douglas-fir.
In our area it is frequent in the fall and winter, but is not as numerous as S. caerulescens
and S. ponderosus, which also grow with Douglas-fir. It prefers poor, exposed soil and
often fruits on roadbanks or in campgrounds.
EDIBILITY: Edible, but not choice.
COMMENTS: This is one “slippery jack” that is not slippery except in very wet weather.
The dark red to reddish-brown fibrils or scales on the cap plus the presence of a veil and
association with Douglas-fir set it apart. When young it is quite attractive, the scales often
being more pink then red. Heavy rain will wash them off, however, and gelatinize the cap
surface. Then the yellow to dingy orange-brown background color predominates, making
it resemble S. caerulescens and S. ponderosus. In the Pacific Northwest it can also be
confused with two larch-lovers—S. cavipes, which has a hollow stalk, and Fuscoboletinus
ochraceoroseus, which is more brightly colored and has darker spores. The “Painted
Suillus” of eastern North America, S. pictus, is a similar but more brightly colored species
associated with white pine. It has a dry cap with pink to red fibrillose scales, a somewhat
longer stalk with similarly colored fibrils, and a soft, white, cottony annulus (ring). It is
edible a-nd highly touted by some, mediocre according to others. Another dry-capped
eastern species, S. decipiens, has an orangish to dingy ochraceous to pinkish-orange,
fibrillose or fibrillose-scaly cap.
Suillus caerulescens commonly occurs with Douglas-fir. Note the presence of a veil which may or
may not form a distinct annulus (ring) on the stalk. A closely related and equally common species,
S. ponderosus, is shown in color plate.

Suillus caerulescens (Douglas-Fir Suillus)

CAP 5-18 cm broad, convex becoming plane or shallowly depressed; surface viscid when
moist, with scattered fibrils, scales, or streaks, color variable: dull cinnamon to yellowish,
but most often dull reddish-brown, orange-brown, or yellow-brown toward the center and
yellowish to buff near the margin, with dingy greenish stains sometimes developing in cold
weather; margin sometimes hung with veil remnants. Flesh thick, pale yellow or dis¬
coloring pinkish to vinaceous. PORES yellow when young, dingier in age, fairly large
(1 mm or more in diameter at maturity) and sometimes radially arranged, not blueing
when bruised but often discoloring dingy reddish-brown or brown; tubes same color,
adnate to decurrent. STALK 2.5-10 cm long, 1.5-3 (4) cm thick, equal or tapered at either
end, firm, solid, dry, yellow above the ring, fibrillose and dingier below and often mottled
with reddish or brown stains and usually staining brownish after handling; sometimes
spotted but not glandular-dotted; flesh near or at base usually staining blue or green when
cut (sometimes slowly). VEIL white or pallid when young, becoming colored like the cap
in age, not slimy, usually forming a slight to distinct ring on stalk; ring fibrillose or bandlike,
median to superior. SPORE PRINT brown to dull cinnamon; spores 8-11 * 3-5 microns,
elliptical to spindle-shaped, smooth.
HABIT AT: Scattered to densely gregarious on ground under or near Douglas-fir; known
only from the Pacific Northwest and California, very common. In our area it is sporadi¬
cally abundant, along with S. ponderosus (see comments), from the fall through early
spring. The largest fruiting is usually around Thanksgiving.
EDIBILITY: Edible. It is generally listed as mediocre, but one collection I sampled had
a rather pleasing lemony flavor.
COMMENTS: If you hunt Douglas-fir forests regularly, you will soon tire of finding this
mundane mushroom. The dull cinnamon to orange-brown or yellowish cap, presence of a
veil, tendency of the stalk base to stain blue when cut, and association with Douglas-firare
key characteristics. A very similar species, S. ponderosus (COLOR PLATE 117), is just
as common in our area and just as mediocre an addition to a meal. It differs in having a
viscid, yellow to orange veil (before breaking) and a smoother, viscid to slimy cap which
ranges from reddish-brown to dull yellowish. Its name refers to its sometimes ponderous
SUILLUS 497

size, and not to the fact that it isn’t associated with ponderosa pine. In fact, it is associated
with Douglas-fir just like S. caerulescens, and the two sometimes mix company. A third
very similar species, S. imitatus, has a smooth viscid cap and shorter spores; it occurs
under conifers in the Pacific Northwest. A dark or dingy greenish variety (S. imitatus var.
viridescens) also occurs, and both S. caerulescens and S. ponderosus can develop greenish
stains on the cap and stalk in cold weather. This in no way affects their edibility, however.

Suillus grevillei (Tamarack Jack)

CAP 3-15 cm broad, convex to nearly plane; surface smooth, viscid or slimy when moist,
often shiny when dry; color variable: deep red to reddish-brown with a yellow margin,
varying to golden-yellow throughout or yellow with a cinnamon- or rusty-tinged center;
margin sometimes with veil remnants. Flesh thick, yellow but often aging or staining pink¬
ish to reddish; not blueing. PORES creamy when young becoming yellowand finally dingy
ochre or olive-yellow, not blueing but usually turning brownish or rusty where bruised;
often somewhat radially arranged; tubes same color, adnate to decurrent. STALK 4-10
cm long, 1 -3 cm thick, equal or slightly thicker below, solid, firm, pale yellow when young
but soon mottled reddish to deep brown or cinnamon below the veil and often staining
brown when handled; glandular dots absent. VEIL yellowish before breaking, the under¬
side often viscid in wet weather; usually forming a median to superior, cottony, whitish ring
on stalk. SPORE PRINT olive-brown to dull cinnamon; spores 8-10 * 3-3.5 microns,
elliptical to spindle-shaped, smooth.
HABITAT: Scattered to gregarious or in small clusters in woods or bogs, associated
exclusively with larch (tamarack); common from late summer through early winter
throughout the northern half of the northern hemisphere wherever larch occurs. It is
very common in Idaho and Montana and also grows on the eastern slope of the Cascades
(frequently in the company of S. cavipes), but is not known from California.
EDIBILITY: Edible, but like most slippery jacks, far from incredible.
COMMENTS: Also known as S. elegans, this attractive slippery jack is easily told by its
smooth golden-yellow to deep red cap, presence of a veil, absence of glandular dots on the
stem, and monogamy with larch. In the West the cap tends to be dark red to cinnamon,
while in Europe and eastern North America it is yellower. S. ponder osus and S. caeru¬
lescens are somewhat similar but prefer Douglas-fir. Other species: S. proximus of
eastern North America is closely related, but its spore print is chestnut-brown when moist
and its stalk often stains greenish when cut open (at least below); it also favors larch.

Suillus grevillei, a slippery jack that grows only with larch. Note slimy cap and prominent annulus
(ring). This is the eastern form with a golden cap; in the West the cap is usually reddish-brown.
m

Suillus umbonatus is a small slippery jack that favors lodgepole pine. Note the glutinous (slimy)
ring on the stalk and the rather large pores.

Suillus umbonatus (Umbonate Slippery Jack)

CAP 3 -9 cm broad, obtusely convex becoming convex or plane, often with a low umbo, the
margin sometimes uplifted in age; surface smooth, viscid or slimy when moist, often
streaked, olive-buff to dingy tan or dull yellowish-brown, often more olive in age, especially
toward margin. Flesh soft, pale yellow to buff, bruising dingy pinkish-brown. PORES
large, more or less radially arranged, pale yellow when young, dingy greenish-yellow in old
age; sometimes staining dingy pinkish-brown when bruised; tubes same color. STALK 3-8
cm long, 4-9 mm thick, more or less equal, solid, usually slender; pale yellow or tan above
the ring, same color or paler below, with pallid to yellow-brown, often obscure glandular
dots; often brownish-stained where handled. VEIL viscid-gelatinous, pale brownish,
forming a median to superior gelatinous ring on stalk. SPORE PRINT olive-brown to dull
cinnamon-brown; spores 7-11 * 3.5-4.5 microns, elliptical to spindle-shaped, smooth.
HABITAT: Scattered to gregarious or in small clumps under conifers, particularly lodge-
pole and beach pines (Pinus contorta), often in large numbers; fruiting in the late summer
and fall or early winter, western North America. It is common in the Sierra Nevada and
on the California coast north of San Francisco; it has also been taken in eastern Canada.

EDIBILITY: Too gooey and gluey to be of value.

COMMENTS: This species is one of our most distinctive slippery jacks, easily recog¬
nized by its dingy yellow-brown to olive-buff color, frequently umbonate cap, viscid
annulus (ring), and rather small, slender stature. The glandular dots are quite obscure
when young but typically darken as the fungus matures. It is closely related to S. sibiricus
and S. americanus, which, however, are more brightly colored and have reddish scales on
the cap. Other species: S. flavidus is slightly yellower but otherwise similar if not the
same; S. helenae, found under lodgepole (beach) pine in coastal Oregon, is also very
similar but lacks a veil.

Suillus sibiricus (Siberian Slippery Jack) Color Plate 116

CAP 3-10 cm broad, convex to broadly convex or broadly umbonate; surface viscid or
slimy when moist, bright yellow to dull yellowish, olive-yellow, ochre-yellow, or dark yel¬
low, with scattered reddish to cinnamon-brown or dark brown spots, scales, streaks, or
patches of fibrils, especially toward the margin (but these sometimes washed off by rain);
margin incurved and usually hung with cottony veil remnants. Flesh soft, thin, yellow, not
blueing but often staining vinaceous or pinkish when bruised. PORES large(l -2 mm wide)

498
SUILLUS 499

when mature, often somewhat radially arranged, mustard-yellow becoming duller or

darker (ochre to yellow-brown) in age, not blueing when bruised (but may stain vinaceous
or pinkish); tubes same color or dingier. STALK 3-11 cm long, 0.5-1.5 cm thick, more or
less equal, often curved, solid or sometimes hollow in age, yellow to ochre-yellow, often
staining vinaceous or brown when handled and often vinaceous-stained at or in the base;
glandular dots and smears present, often inconspicuous when young but typically brown to
cinnamon and quite conspicuous in age. VEIL cottony, whitish to pale yellow or in age
sometimes brownish, usually remaining attached to margin of cap, but sometimes forming
a slight ring on stalk. SPORE PRINT brown to dull cinnamon-brown; spores8-l 2 * 3.5-4.5
microns, elliptical to spindle-shaped, smooth.
HABITAT: Scattered to gregarious or clustered under conifers, associated with white
(5-needle) pines; widely distributed, but especially common in the summer and fall with
western white pine in the Pacific Northwest and northern California. It is also found in
the Southwest with southwestern white pine and limber pine.The very similar A. ameri-
canus{see comments) replaces it in eastern North America.
EDIBILITY: Edible but thin-fleshed, insipid, and slimy. Alexander Smith relates the case
of one person who cannot even touch it without suffering an acute allergic reaction.

COMMENTS: The combination of viscid yellow cap with reddish to brown spots or
plaques, cottony veil, and glandular-dotted stalk make this one of the most distinctive of
all the slippery jacks. The cap is usually bright yellow in dry weather and duller or dingier
when moist. The closely related S. americanus is often abundant with eastern white pine
and also occurs in the Southwest. Its cap is often more streaked than spotted and its stalk
is usually thinner (3-10 mm), plus it lacks an annulus because its veil normally doesn’t come
into contact with the stalk (instead it extends in from the cap margin as a roll of cottony tis¬
sue). In the S outhwest the two species seem to intergrade; the color plate may represent one
such “hybrid.” In the Sierra Nevada two somewhat similar species, S. riparius and S.
megaporinus, are often common. Both have very large, irregular or even gill-like pores
(up to 5 mm long and 3 mm broad) and yellow to brownish caps with brown to rusty-brown
scales, streaks, and/or fibrils. S. riparius usually has a well-developed stalk while in S.
megaporinus the stalk is often poorly developed and off-center and the cap is usually small.

Suillus subolivaceus (Slippery Jill) Color Plate 121

CAP 4-15 cm broad, convex or obtusely umbonate becoming plane or broadly umbonate;
surface smooth but often streaked or with flattened darker fibrils, viscid or slimy when
moist; dull olive to olive-brown, grayish-olive-brown, dingy yellow-brown, or dingy tan.
Flesh pallid to yellowish or tinged olive-gray, not bruising blue. PORES fairly large at
maturity, grayish-olive to grayish-buff or olive-buff when young and often beaded with
clear droplets in wet weather, becoming dingy yellowish to brownish-yellow in age (and
droplets often drying blackish); not bruising blue; tubes same color or yellower. STALK
6-12 (17) cm long, 0.8-2 cm thick, equal or thicker below, solid, white or yellowish above
the ring, whitish to brownish below; densely covered throughout with prominent pinkish-
brown to blackish glandular dots and smears. VEIL membranous, whitish, with a gelatin¬
ous olive to brownish underside; forming a large bandlike, median to superior ring on stalk
which may shrink considerably in age. SPORE PRINT brown to dingy cinnamon; spores
8-11 x 3-4 microns, elliptical to spindle-shaped, smooth.
HABITAT: Scattered to gregarious under conifers, particularly western white pine;
known only from western North America. In the Pacific Northwest it is often common in
the late summer and fall. It probably occurs in California, but I have not seen it south of
the Cascades. Similar species occur in eastern North America (see comments).
500 BOLETACEAE

EDIBILITY: Edible, but of poor quality.

COMMENTS: The large bandlike ring, overall dingy olive-brown to yellow-browncolor,
slimy cap, and very conspicuous glandular dots on the stem which become blackish in age
make this slippery jack easier to identify than it is to grip! The glandular dots are often quite
sticky and resinous and will stain your fingers brown more quickly than those of other
slippery jacks. S. umbonatus is somewhat similar but grows with lodgepole pine and has a
gelatinous annulus (ring) and pale yellowish pores when young. S. luteus is easily distin¬
guished by its reddish-brown cap and sheathing, purplish-tinged veil. In eastern North
America there is a group of “slippery jills” with a yellowish to olive-brown cap and
strikingly broad (1 -2 cm high), thick annulus that is baggy (flares at both the bottom and
top edges) when young and bandlike in age. The most common northern member of this
group is S. subluteus. Several species are common in the South, including^, cothurnatus,
S. salmonicolor (with salmon-tinged pores), and S’, pinorigidus. None are worth eating.

Suillus luteus (Slippery Jack) Color Plate 118

CAP 5-12 cm broad, obtuse or convex becoming plane; surface smooth, viscid or slimy
when moist, often shiny when dry; typically reddish-brown to dark reddish-brown or
chestnut-brown, but ranging sometimes toward yellow-brown or rusty-brown, especially
in age, and sometimes streaked; margin often hung with veil remnants. Flesh thick, white to
pale yellow, not bruising. PORES minute, white at first, becoming yellow and finally dark
yellowish, often brownish-spotted inage; not bruising blue; tubes same color. STALK 3-10
cm long, 1 -3 cm thick, more or less equal, solid, white to pale yellow above, often purplish
below or with a purplish zone; prominent pinkish to brown glandular dots and smears soon
developing both above and below. VEIL sheathing the stalk, membranous, persistent,
white above, purplish to dull lilac below and viscid when wet; usually forming a large, often
flaring or sleevelike ring that turns brown from falling spores. SPORE PRINT brown to
dull cinnamon; spores 6-10 * 2.5-3.5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Scattered to gregarious in cool wet weather under conifers, particularly pine
but also spruce; widely distributed. It is abundant in pine plantations in northeastern North
America, and hasalso beenreportedfromthePacificNorthwestandSouthwest. Afavored
associate is Scots pine (Pinus sylvestris), and it may very well turn up in California where
Scots pine has been planted.

EDIBILITY: Edible and widely collected, though some people are apparently “allergic” to
it. According to one source it is “the best of the slippery jacks”—a classic case of damning
with faint praise, if you ask me. As in most slippery jacks, the slimy skin peels off easily.
COMMENTS: This species is the “original” slippery jack, i.e., the one from which all the
others take their name. It can be told by its reddish-brown cap, glandular-dotted stem, and
sheathing veil which typically forms a prominent annulus (ring) on the stalk. The dull
purplish color of the lower stalk or underside of the ring is also distinctive. S. borealis
(see comments under S. pseudobrevipes) is a closely related westerner with no annulus.

Suillus pseudobrevipes (Veiled Short-Stemmed Slippery Jack)

CAP 5-15 cm broad, convex becoming broadly convex or plane; surface slimy or viscid
when moist, smooth or with a few patches of veil tissue near margin, but often appearing
fibrillose or streaked; dingy yellowish to yellow-brown, dark yellow-brown, or honey-
colored, often becoming darker or more cinnamon in age; margin often hung with whitish
veil remnants. Flesh thick, white to pale yellow, not blueing. PORES and tubes pale yellow
to yellow becoming dingy yellowish in age, not blueing. STALK 2-8 cm long, 1-3 cm
Suillus pseudobrevipes, a widespread slippery jack with a thin veil and inconspicuous glandular dots.
The veil may form a slight ring on the stalk and/ or leave remnants on the margin of the cap, or even
disappear entirely. These are rather small specimens.

thick, often rather short and thick, equal or tapered downward, solid, firm, white when
young, yellowish in age; glandular dots absent or obscure when young but tending to
become brown and more visible in age. VEIL white to dingy lavender or lavender-brown,
usually forming a median fibrillose ring on stalk, but the ring often slight or collapsed;
sometimes forming a sheath over the base of stalk or merely leaving remnants on cap
margin. SPORE PRINT brown to pale brownish; spores 7-9 * 2.54 microns, spindle-
shaped to elliptical, smooth.
HABITAT: Scattered to gregarious under pines (mainly ponderosa and lodgepole);
known only from western North America. In our area it fruits only in scattered localities
with ponderosa pine, in the fall and winter. In the Southwest and Rocky Mountains it is
quite abundant, however, and I’ve also seen it in the Sierra Nevada.
EDIBILITY: Edible.
COMMENTS: This slippery jack tends to have a short, stocky stem like its namesake,
V. brevipes, but the presence of a veil and paler cap color distinguish it. Sometimes the veil
persists as a roll of cottony tissue on the cap margin instead of forming a distinct annulus
(ring), leading to confusion with several species that typically boast veil remnants on the
margin of the cap but not the stalk. These species include: S. volcanalis, glandular dots
obscure, cap yellowish or tawny (or often with pinkish to reddish-brown areas), growing
in volcanic soil (often buried) with Jeffrey pine in Mt. Lassen National Park, California;
S. brunnescens, cap white with the slime becoming chocolate-brown, glandular dots small,
associated with sugar and lodgepole pines; S. borealis, cap dark brown, veil dull lavender
or purplish (as in S. luteus), stalk glandular^dotted in age, associated with western white
pine; S. glandulosipes, cap fibrillose and pale ochre to cinnamon, glandular dots very
conspicuous, locally common in northern coastal California; and S. albidipes, whose cap
is whitish to buff, brownish, or dull cinnnamon and has a roll of cottony veil tissue on the
margin when young. I have seen enormous fruitings of the latter species under lodgepole
pine in Oregon and Washington.

Suillus brevipes (Short-Stemmed Slippery Jack) Color Plate 115

CAP 5-13 cm broad, convex becoming broadly convex to plane; surface smooth, viscid
or very slimy when moist, often shiny when dry; dark vinaceous-brown to dark brown
when young, often fading in age to reddish-brown, dull cinnamon, or even tan (or in one
form yellow-brown) and sometimes appearing streaked; margin naked. Flesh thick, white
or often becoming yellow in age, soft, not blueing when bruised. PORES and tubes pale
when young, becoming darker or dingier yellow in age and finally olive-yellow; not blueing.
STALK2 -7 cm long, 1 -2 (3) cm thick, equal or slightly thicker belowand often quite short;
firm, solid, white becoming pale yellow in age; glandular dots absent or sometimes barely
visible in age (never,prominent). VEIL absent. SPORE PRINT brown to dull cinnamon;
spores 7-10 * 34 microns, elliptical to spindle-shaped, smooth.
HABITAT: Scattered to densely gregarious under conifers (particularly 2- and 3-needle

501
502 BOLETACEAE

pines, also spruce); very widespread and common. In the West it is especially abundant
with lodgepole and bishop pines. In our area it fruits in the fall and winter but is en¬
countered infrequently, perhaps due to the superabundance of S. pungens.
EDIBILITY: Edible and perhaps the best of our local slippery jacks—which is hardly a
compliment. Peel the slimy pellicle (skin) before cooking it.
COMMENTS: This widespread and often abundant slippery jack is best recognized by
its smooth, slimy, dark brown to reddish-brown cap (which may fade in age), absence of a
veil, and absence of obvious glandular dots on the stem. True to its name, the stalk is often
rather short, but it is a variable species and longer-stemmed individuals are not unusual.
The viscid cap and non-reticulate, more or less equal stalk distinguish it from Boletus; the
cap is never olive-gray as in S. pungens, and there is no veil as in S. pseudobrevipes, while
5. granulatus and its relatives have a distinctly glandular-dotted stem. Other species witha
smooth viscid cap, no veil, and absent or inconspicuous glandular dots include: S. pallidi-
ceps of the Rocky Mountains, with a white cap when young that becomes pale cinnamon
or yellowish in age; and S. occidentalis, common under ponderosa pine in the Southwest,
with a paler (buff to light brown or pinkish-brown) cap that is not white when young.

Suillus granulatus (Granulated Slippery Jack)

CAP 3-15 cm broad, convex to plane or slightly wavy; surface smooth, viscid or very slimy
when moist, often shiny when dry; cinnamon-brown to brown, orange-brown, rusty-
cinnamon, or yellow-brown (but in some forms whitish when young), often streaked or
mottled and paler or duller in old age; margin naked. Flesh thick, soft, white when young
but soon pale yellow. PORES and tubes whitish when very young and often beaded with
milky droplets, soon becoming buff or yellow and eventually dingy yellowish or brownish-
spotted when mature. STALK 3-8 cm long,0.7-2.5 cm thick, more or less equal, firm, solid,
whitish when young, becoming yellow above and dingy cinnamon- or reddish-stained
toward base; covered with pinkish to reddish-tan or brown glandular dots and smears in
age. VEIL absent. SPORE PRINT brown to dull cinnamon or ochre; spores 7-10 * 2.54
microns, spindle-shaped to elliptical, smooth.
HABITAT: Scattered to densely gregarious on ground under pines in the summer, fall,
and early winter; very widespread and common, but replaced in our area by S. pungens.
EDIBILITY: Edible, but bland (see S. pungens).
COMMENTS: This widespread slippery jack can be found almost anywhere where there
are pines. It is closely related to S. pungens, but lacks the olive-gray tones so characteristic
of that species. The combination of glandular-dotted stem, slimy or shiny cap, and absence
of a veil are quite distinctive, but there are a number of closely related species which are
rather difficult to distinguish (fortunately, allare apparently edible). These variants, which
are especially numerous and confusing in the pine forests of the Rocky Mountains and
Southwest, include: S.flavogranulatus of the Rockies, witha yellowish to dingy ochre cap
at maturity and larger pores; 5. kaibabensis, common in the Southwest, with a buff to pale
cinnamon or yellowish cap; and S. wasatchicus, with reddish pores when young that
become yellowish in age. In the Pacific Northwest another species, S. punctatipes, is quite
common under various conifers. It also has glandular dots or smears and lacks a veil, but
is larger and thicker-stemmed with larger yellow pores that are often somewhat radially
elongated and/ or decurrent. Its flesh is white and its cap color variable: orange-brown to
dark brown, dingy tan, pinkish-brown, gray, vinaceous-gray, or purple-tinged. 5. mon-
ticolus of the Sierra Nevada resembles S', granulatus but has a swollen or bulbous stem.
S. placidus of eastern North America is very similar, but its cap remains whitish for a long
time before darkening or turning yellowish. Also see S. albidipes (under S. pseudobrevi¬
pes), which differs in having veil tissue on the cap margin when young.
Suillus pungens. Note slimy cap and presence of milky droplets on pore surface of young specimens,
plus the glandular-dotted stem. Cap color is variable (see color plate and description). (Joel Leivick)

Suillus pungens (Pungent Slippery Jack) Color Plate 114

CAP 4-18 cm broad, convex becoming broadly convex or plane; surface smooth, slimy or
viscid when moist, often shiny when dry; color extremely variable: usually white when very
young but soon olive to grayish-olive and then becoming yellow, tawny-cinnamon, rusty-
brown, orange, reddish-brown, or often mottled and streaked with various combinations
of the above colors; margin typically naked or with a slight roll of cottony tissue when
young. Flesh thick, white when young, soon becoming lemon-yellowand soft, not blueing;
odor pungent or pleasant (somewhat banana-like). PORES white when young and often
beaded with white to pinkish droplets in wet weather, becoming yellow and finally dark
yellow-brown or dingy ochre in age, not blueing when bruised; tubes same color. STALK
2-10 cm long, 1 -2 (3) cm thick, equal or with a slightly tapered or swollen base, firm, solid,
white when young becoming yellow in age, with conspicuous reddish to brown glandular
dots and smears. VEIL absent. SPORE PRINT olive-brown to dull cinnamon-brown;
spores 9-10 * 3-3.5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Scattered to densely gregarious or clustered on ground near or under Mon¬
terey pine; apparently endemic to our area, but also occurring where M onterey pines have
been planted (it is a common “lawn mushroom” in our area, often growing with Chroo-
gomphus vinicolor and Helvetia lacunosa). It is most abundant in the fall and winter but
can be found whenever it’s damp enough. It is also quite common under knobcone and
ponderosa pines—but apparently only within the geographical range of Monterey pine.
EDIBILITY: Edible; disdained by some due to its“harsh, unpleasant, subnauseous” taste,
but prized by others. In my opinion its soft flesh and slimy texture are much bigger draw¬
backs than its flavor. Since it occurs in such large quantities and is easy to identify, it is
definitely worth experimenting with. To remove the slime, simply peel off the skin.
COMMENTS: Locally this is our most prolific bolete, appearing in colossal numbers
wherever there are pines. The remarkable series of color changes it undergoes is apt to
baffle the color-conscious beginner. On the other hand, when several individuals are
present it affords an instant means of recognition! It is closely related to S. granulatus, but
that species is never grayish-olive when young. The glandular-dotted stem and milky
droplets on the pore surface of young specimens are also distinctive.
S. pungens vies with Hygrophorus eburneus and the “Insidious Gomphidius” (Gom-
phidius oregonensis) for the title of “Slipperiest and Slimiest Fungus Among Us.” An
organic version of hockey or caroms can be played ona wet surface, using the caps of young
S. pungens for pucks. Overmature individuals, on the other hand, are more aptly called
S. “spongens” than S. pungens—they literally seethe with fat, agitated maggots and sag
with so much excess moisture that they practically demand to be wrung out like a sponge!

503
504 BOLETACEAE

Suillus fuscotomentosus Color Plates 119,120

(Poor Man’s Slippery Jack)
CAP 4-15 cm broad, convex to plane; surface dry, or viscid in age or wet weather, at first
covered with olive-brown to dark brown or fuscous (deep grayish-brown) fibrils or small
fibrillose scales (in this stage usually dry), the scales often sparser in age, revealing the paler
(dull ochre to buff) background, sometimes more cinnamon-colored overall at maturity
or appearing streaked; margin naked. Flesh thick, yellow to pale orange or orange-buff,
not blueing when bruised; taste mild to slightly unpleasant. PORES typically orange-buff
when young and sometimes beaded with droplets, becoming yellowish-buff to dark dingy
yellow or olive-yellow in old age, not blueing when bruised; tubes same color. STALK
4-12 cm long, 1 -3 cm thick, equal or somewhat thicker below, solid, firm, pallid to yellow
or brownish-buff or often pinkish-orange to pale orange, especially toward base; glandular
dots and smears present and usually conspicuous at least in age, often elongated, same
color as stalk or darker (brown to cinnamon). VEIL absent. SPORE PRINT olive-brown
to dull cinnamon-brown; spores 9-12 * 3-4 microns, spindle-shaped to elliptical, smooth.
HABITAT: Scattered to densely gregarious on ground under pines, often in large num-
bersr; originally described from Santa Cruz County, California, but occurring in the Sierra
Nevada and Cascades as well. In our area it is often abundant in the late fall and winter
under ponderosa, knobcone, and digger pine, while the very similar S. acerbus (see
comments) favors Monterey pine.
EDIBILITY: Edible, but slimy and insipid. In blind tastings of local edible boletes, it has
consistently placed last. Asked to rate each bolete on a scale of 1-10, several tasters gave
it a zero and one, a minus five!
COMMENTS: This common bolete resembles S. tomentosus but does not stain blue. The
pores are usually orange-buff when young, rather than white as in S. pungens and S.
granulatus, and the brownish to grayish-brown cap fibrils and absence of a veil are also
distinctive. The “Starving Man’s Slippery Jack,” S. acerbus, is a very similar species with
a more viscid, less fibrillose-scaly cap. It is supposedly associated exclusively with Mon¬
terey pine, but I wonder whether it is really distinct from S. fuscotomentosus. The degree
of viscidity seems to depend on age and weather conditions, and I’ve found specimens
under Monterey, knobcone, and digger pines which could be referred to either species.

Suillus tomentosus (Blue-Staining Slippery Jack)

CAP 5-15 (20) cm broad, convex to plane; surface dry, but in wet weather or age often
viscid, at first covered with a dense coating of minute hairs (fibrils) which break up to form
scattered scales and are often quite sparse (or even absent) in age; fibrils or scales usually
grayish-brown to brown (reddish in one form), sometimes pallid or buff when young; back¬
ground usually pale yellow to yellow-orange, orange-buff, or dark dull orange; margin
naked. Flesh thick, pallid to yellow, usually blueing(but sometimes slightly or slowly) when
bruised; taste mild or acidic. PORES dingy brown to dark cinnamon when young,
becoming dingy yellow to buff or olive-yellow in age; usually blueing at least slightly when
bruised (at least in age); tubes yellowish to olive-yellow. STALK 3-11 (15) cm long, 1-3 cm
thick, equal or thicker below, firm, solid, colored like background of cap or oranger, with
numerous small glandular dots which may be same color or browner and are often obvious
in age; base sometimes reddish-stained. VEIL absent. SPORE PRINT dark olive-brown
to dull cinnamon; spores 7-11 * 3.5-4.5 microns, elliptical to spindle-shaped, smooth.
HABITAT: Scattered to gregarious or in troops onground under pines and other conifers,
late summer through early winter; widely distributed, but especially common in the West,
where it is one of the most common of all the slipperyjacks. I haveseenenormousfruitings
Suillus tomentosus is a common and widespread species easily told by the tendency of its pores and/
or flesh to stain bluish when bruised. Note presence of fibrillose scales on cap. Older or rain-battered
specimens, however, may be practically bald (without scales or hairs).

in Idaho, Washington, northern California, and the Sierra Nevada, especially with lodge-
pole pine and in mixed forests of aspen and pine. In our area, however, it is largely re¬
placed by the similar S.fuscotomentosus.
EDIBILITY: Edible, and every bit the equal of S. fuseotomentosus (see comments on
the edibility of that species).
COMMENTS: The fibrillose to scaly cap, brownish pores when young, absence of a veil,
glandular-dotted stem, and tendency of the poresand/ orthe flesh tostainblue orgreenish-
blue when bruised form a distinctive combination of characters. It is one of the commonest
and most variable slippery jacks, with several slightly different color forms. Other species:
S. variegatus of Europe is very similar if not the same; S', reticulatus is a rare, reticulate-
stalked species that stains blue.

FUSCOBOLETINUS (Larch Boletes)

Medium-sized, fleshy, terrestrial boletes associated mainly with larch. CAP viscid, or if dry then
fibrillose. PORES pallid to yellowish, gray, or grayish-brown, often large and radially arranged.
STALK not usually reticulate or glandular-dotted; not scabrous. VEIL present, frequently forming
an annulus (ring) on stalk. SPORE PRINT dark grayish-brown to dark reddish-brown to
chocolate-brown or chocolate-gray. Spores elliptical to spindle-shaped, smooth. Cystidia on inner
surfaces of tubes staining dark brown in KOH (potassium hydroxide).

THIS small genus is an off-shoot of Suillus. It differs principally in its darker spore color,
and is included in Suillus by some boletologists. A veil is present in all species but the stalk
usually lacks glandular dots. The cap varies from smooth and slimy to dry and fibrillose.
Fuscoboletinus species are mycorrhizal mainly with larch or tamarack (Larix). This is a
helpful feature in identification, though several Suillus species (e.g., S. grevillei and S.
cavipes) also grow with larch. Some Fuscoboletinus species grow with other northern
conifers, but their geographical range still corresponds to that of larch. As larch does not
occur in California, Fuscoboletinus probably doesn’t either. However, it could con¬
ceivably appear where larches have been planted. Only two species are known from the
western United States, and they are described here; most of the other species occur in the
Northeast and Canada. Like slippery jacks, they are edible but of mediocre quality.

505
506 BOLETACEAE

Key to Fuscoboletinus
1. Stalk distinctly glandular-dotted (the dots dark); known from Minnesota; rare . F. weaverae
1. Stalk not glandular-dotted (but may have reddish spots when young); common .2
2. Tubes and pores yellow to yellow-brown or yellow-olive when fresh . 3
2. Tubes and pores whitish to grayish or grayish-brown, not yellowish . 10
3. Cap 4-10 cm broad, at first covered by cottony grayish to yellow or red veil material which breaks
up to form coarse scales, streaks, or plaques; cap viscid beneath the veil material and usually
dark red to brownish or red-and-yellow in age; partial veil with both a cottony and a viscid or
gelatinous layer; stalk with veil remnants (like cap), usually a mixture of red, yellow, and/ or
gray; pores and tubes yellow; associated with larch (tamarack), usually growing in bogs;
found in the northeastern United States and Canada (a very striking mushroom—see Color
Plate 126) .F. spectabilis
3. Not as above .4
4. Cap viscid or slimy when moist and more or less smooth . 5
4. Cap dry (not viscid) and fibrillose or scaly . 8
5. Flesh in lower stalk staining greenish when cut (see Suillusproximus under S', grevillei, p.497)
5. Not as above .6
6. Very common with larch in both eastern and western North America; spore print olive-brown
to dull cinnamon .(see Suillus grevillei, p. 497)
6. Found in northeastern U .S., Canada, and Alaska, usually with conifers other than larch; spore
print darker than above . 7
7. Veil(and stalk below veil) viscid; cap red-brown to mahogany, the stalk often with reddish spots
when young; found under conifers in Alaska, Canada, and northeastern U.S. F. glandulosus
7. Not as above; stalk dry (not viscid) .F. sinuspaulianus
8. Stalk typically becoming hollow in lower portion; cap color variable: dark brown to reddish-
brown, tawny, etc.; spore print dark olive-brown to brown .... (see Suillus cavipes, p. 494)
8. Stalk not normally hollow; cap rosy to dark red (but sometimes overlaid with white) when
fresh; spore print dark reddish-brown to dark vinaceous-brown . 9
9. Stalk typically 1-3 cm thick and cap 6-20 cm broad; known only from western North America
.F. ochraceoroseus, below
9. Not as above; typically smaller and eastern .F. paluster (see F. ochraceoroseus, below)
10. Cap only slightly viscid; flesh not blueing when bruised; pores usually elongated near the stalk
or forming gill-like rows; found in the eastern United States and Canada, usually in bogs
.F. grisellus(see F. aeruginascens, p. 507)
10. Not as above; cap viscid to slimy when moist; flesh often (but not always) staining bluish or
bluish-green when exposed; found in both eastern and western North America . 11
11. Cap more or less chocolate-brown when fresh; known from the eastern United States and
Canada .F. serotinus (see F. aeruginascens, p. 507)
11. Not as above; widespread .F. aeruginascens, p. 507

Fuscoboletinus ochraceoroseus Color Plate 123

(Rosy Larch Bolete)
CAP 7-20 (25) cm broad, convex to plane or slightly umbonate, or in age the margin some¬
times uplifted; surface dry, densely fibrillose or fibrillose-scaly and often uneven or pitted;
rosy-red to bright pink, but often overlaid with whitish hairs(fibrils),and oftendarkerand
duller (brick-red) in age; margin sometimes yellowish and hung with veil remnants. Flesh
thick, yellow or tinged pink under cap cuticle, bruising slightly greenish-blue or not at all;
taste often slightly bitter or acrid. PORES large, elongated (2-5 mm long) and usually
arranged radially; yellow becoming dark yellow to olive-ochre and finally dingy brown in
old age, not bruising blue; tubes shallow, same color, adnate to decurrent. STALK 3-7 (10)
cm long, 1-3 cm thick, equal or with an enlarged base (and often a flaring apex); solid,
firm, yellow(colored like pores) or with reddish to brownish stains, base often whitishand
FUSCOBOLETINUS 507

fibrillose; apex often slightly reticulate from tubes; glandular dots absent. VEIL
membranous, white or yellowish, thin, sometimes forming a slight ring on stalk but more
often clinging to margin of cap. SPORE PRINT dark reddish-brown to dark vinaceous-
brown; spores 7.5-9.5 * 2.5-3 microns, elliptical to spindle-shaped, smooth.
HABITAT: Solitary to scattered or in groups on ground under conifers, associated with
larch; known only from the northern Rocky Mountains and Pacific Northwest. I have seen
large fruitings in Idaho in September, but apparently it isn’t common every year.
EDIBILITY: Edible, but of very poor quality due to the slightly bitter taste.
COMMENTS: The beautiful rosy-red to pink cap, radially arranged pores, presence of a
veil, and association with larch are the most distinctive characters of this most distinctive
bolete. Its color is somewhat reminiscent of Suilluspictus, which grows with eastern white
pine, and S. lakei, which grows with Douglas-fir, but the spore print is darkerand the stalk
usually thicker. Other species: F. paluster of northeastern North America is somewhat
similar (cap deep red) but much smaller and more slender, with even larger pores; it grows
in swamps and bogs. Another very striking bog-lover, F. spectabilis (COLOR PLATE
126), is also restricted to the Northeast and Canada. One glance is usually sufficient to
identify it (as evidenced by the color plate), but see the key to Fuscoboletinus for details.

Fuscoboletinus aeruginascens (Grayish Larch Bolete)

CAP 3-12 cm broad, convex becoming plane or slightly umbonate; surface viscid or slimy
when moist, smooth or streaked with flattened fibrils, or at times with darker fibrillose
scales, sometimes cracking in dry weather; grayish-white to gray, grayish-brown, olive-
gray, or dingy yellowish, sometimes with darker scales or developing grayish to dingy olive
spots; margin usually hung with veil remnants. Flesh soft in age, white to yellowish,
typically turning bluish-green when bruised (but often slowly or only very slightly), at
least in the stalk. PORES small and round when young but often rather large and somewhat
radially arranged in age; white at first becoming gray to grayish-brown at maturity, usually
staining blue to dingy greenish when bruised (but often only slightly or slowly); tubes same
color. STALK 4-6 (9) cm long, 0.7-1.5 cm thick, equal or tapering upward or sometimes
pinched at base; solid, pallid or tinged olive above the ring, gray to olive-grayish to
brownish below and usually somewhat viscid; glandular dots absent, apex often reticulate
from tubes. VEIL membranous, white to gray or yellowish, usually forming a median to
superior ring on stalk. SPORE PRINT dark reddish-brown or vinaceous-brown; spores
8-12 x 3.5-5 microns, spindle-shaped to elliptical, smooth.

HABITAT: Scattered to densely gregarious in woods and around the edges of bogs, asso¬
ciated with larch (or tamarack) and common wherever larch occurs—the most widespread
member of the genus. In the Pacific Northwest it is often common in the spring and again in
the late summer and fall; it probably does not occur in California.
EDIBILITY: Edible, but unappealing because of its sliminess, thin flesh, and dingy color.
COMMENTS: The whitish to grayish to yellowish cap, white (never yellow!) pores that
become grayish in age, presence of a veil, and tendency to bruise weakly bluish-green are the
distinguishing fieldmarks of this undistinguished larch-lover. It is likely to be mistakenfor
a Suillus (and is even placed in that genus by some mycologists), but the spore print is
vinaceous-brown. In eastern North America it often grows with F. spectabilis, a very
striking species described in the key to Fuscoboletinus and shown in Color Plate 126.
F. serotinus is a similar species with chocolate-brown slime on the cap; F. grisel/us is
colored like F. aeruginascens but does not normally bruise bluish-green, has a more conical
and less viscid cap, and more elongated pores. Both species are associated with tamarack
(eastern larch) and are fairly common in the northeastern U.S. and Canada.
508 BOLETACEAE

BOLETELLUS& AUSTROBOLETUS
Medium-sized woodland boletes. CAP fleshy, dry or viscid, sometimes areolate or scaly. PORES
and tubes usually (but not always) yellow, sometimes blueing when bruised. STALK fleshy but
usually long, slender, and more or less equal; lacking glandular dots or dark scabers but often
longitudinally ridged or jaggedly reticulate. VEIL absent (except in B. ananas). SPORE PRINT
olive to brown or cinnamon-brown. Spores elliptical to spindle-shaped, longitudinally wrinkled,
ridged, grooved, or “winged" (Boletellus) or minutely pitted (AustroboletusJ.

THESE are gracile (slender-stemmed) boletes with ornamented spores (ridged or wrin¬
kled in Boletellus, pitted in Austroboletus). The stalk is usually long in relation to the cap
and not bulbous as in many species of Boletus. In addition, it is often raggedly or jaggedly
reticulate or ridged. Both genera are centered in the tropics. In the U nited States they are
most prevalent in the southeastern sector and along the Gulf of Mexico. A few species
range north to Canada and west to southern Arizona, but none have been found in Cali¬
fornia. One Boletellus is depicted here; several other species are keyed out.

Key to Boletellus & Austroboletus

1. Stalk smooth to fibrillose or scurfy but not reticulate, jagged, or markedly ridged; pores and
flesh usually blueing when bruised; often growing at or near the bases of trees . 2
1. Stalk markedly or jaggedly reticulate or ridged; flesh and pores not blueing; terrestrial .... 3
2. Veil present when young, usually leaving white flaps on cap margin; cap coarsely scaly or fibril-
lose-scaly; usually growing on or near pine trees .B. ananas (see B. russellii, below)
2. Veil absent; cap often areolate but not fibrillose-scaly or shaggy.B. chrysenteroides
3. Cap and stalk whitish to pale yellow or buff; stalk prominently and coarsely reticulate; taste
bitter; usually found in sandy soil in southern U.S. (subtropical) .A. subflavidus
3. Not as above; cap and stalk darker or brighter . 4
4. Cap smooth and viscid when fresh and moist .A. betula (see Boletellus russellii, below)
4. Cap not viscid, often areolate (cracked) in age.B. russellii, below

Boletellus russellii (Jagged-Stemmed Bolete) Color Plate 125

CAP 3-9 (13) cm broad, convex to broadly convex or rarely plane; surface dry, yellow-
brown to buffy-brown or olive-gray, varying to brownish, reddish, or cinnamon-brown;
minutely velvety to obscurely scaly, often becoming areolate (breaking up into small
scales) in age, revealing the flesh beneath; margin at first incurved. Flesh yellow, not
blueing when bruised. PORES rather large (1 mm broad or more), yellow when young,
greenish-yellow in age, not blueing; tubes same color. STALK 10-20 cm long, 0.8-2 cm
thick, equal or slightly thickened downward, typically long and slender, often curved at
base; coarsely reticulate-lace rate (ragged and deeply ridged) more or less throughout; dull
reddish to reddish-brown or cinnamon; solid, dry or with a viscid base when fresh. VEIL
absent. SPORE PRINT dark olive to olive-brown; spores 15-20 ><7-11 microns, elliptical
to spindle-shaped, deeply ridged or wrinkled longitudinally, with a cleft at apex.
HABITAT: Solitary, scattered, or in small groups on ground under hardwoods (espe¬
cially oak) or occasionally conifers; fairly common in the summer and early fall in eastern
North America, but rarely fruiting in large numbers. It also occurs in southern Arizona,
like many other “eastern” species.
EDIBILITY: Edible, but rather soft and bland.
COMMENTS: This distinctive bolete is easily recognized by its long, slender, lacerated
(ragged or jagged) stalk (see the color plate), dry cap that is frequently areolate, and
yellow to greenish-yellow pores that do not stain blue. Another species with a long, slim,
lacerated stalk, Austroboletus (-Boletus, Boletellus) betula, differs in having a smooth,
viscid, yellow-orange to reddish-brown cap, a more southern distribution, and pitted
BOLETELLUS 509

spores. Another southerner, Boletellus ananas, is quite different. It has a fibrillose to

coarsely scaly or shaggy, purplish to reddish (or sometimes yellowish) cap plus a whitish
veil when young. The veil usually leaves remnants on the cap margin rather than forming
a ring. The stalk is whitish to tan and smooth to slightly fibrillose, and the flesh and pores
turn blue when bruised or cut. Neither of the above species is worth eating.

PULVEROBOLETUS
AS DEFINED here, Pulveroboletus contains an awesome total of one common species.
The prefix pulvero means pulverulent, which in turns means “powdery.” It pertains to
the powdery to somewhat cottony or cobwebby consistency of the bright yellow veil that
covers the cap and stalk of the young fruiting body—a unique feature among the boletes.
Some mycologists, however, broaden the genus concept to include veil-less species such
as P. auriporus and P. hemichrysus (treated in this book under Boletus).

Pulveroboletus ravenelii (Veiled Sulfur Bolete)

CAP 2-8 (12) cm broad, convex or nearly round when young, becoming plane in age; sur¬
face dry to slightly viscid if wet, bright yellow with orange, pinkish, or reddish-brown tones
developing at the center, especially in age; covered with yellow cottony or powdery veil
material, at least when young; margin often covered with veil remnants also. Flesh thick,
white to yellow, changing slowly to blue when exposed, then eventually yellowish to dingy
brown. PORES and tubes bright yellow when fresh, becoming dingier or olive-yellow in
age; usually bruising bluish-green, and then sometimes turning dark brownish. STALK
6-15 cm long, 0.5-1.5 cm thick, equal to slightly irregular, often rather long; dry, bright
yellow, powdery or cottony at least when young, except at apex. VEIL bright yellow,
cottony-powdery to cobwebby, leaving remnants on cap and cap margin and stalk (and
sometimes a slight ring on stalk). SPORE PRINT dark olive-brown to brown; spores
8-12 x 4-6.5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary or in small groups in woods; widely distributed, but rather rare on
the west coast. In our area I have found it in the fall and early winter in mixed woods of oak
and pine and under manzanita.
EDIBILITY: Like myself, not firmly established.
COMMENTS: I was dumbfounded by the sight of my first Pulveroboletus. As boletes go
it is an odd creature—a gaudy misfit, an audacious anomaly. The brilliant yellow cottony
veil which covers the cap and stalk when young sets it apart from all other boletes. In old
age or rainy weather the veil remnants may be obliterated, but the color and relatively
long, slender stalk help identify it. Though rare in California, it is included here because
it is so bizarre. If you should be (un)fortunate enough to fall on top of one, as I did, I would
like you to know what it was.

An unusually gracile (long and slender) example of Pulveroboletus ravenelii. The distinctive
yellow veil is not clearly visible in this picture.
510 BOLETACEAE

GYROPORUS
Small to medium-sized, terrestrial boletes. CAP typically dry. PORES pallid or whitish when
voung, pale yellow to yellow in age. STALK lacking scabers and glandular dots and typically not
reticulate; usually hollow at maturity, at least toward the base. VEIL absent or rudimentary.
SPORE PRINT pale yellow to yellow. Spores elliptical to spindle-shaped, smooth.

THE pale yellow spore print and hollow or partially hollow stem are the distinctive
features of this small genus. Four species—all choice edibles—are included in the key, but
only one, G. castaneus, has been found in California. In addition to the generic charac¬
teristics, G. castaneus can be told by its chestnut-brown to orange-brown cap, white to pale
yellow pores, and often slender, uneven, similarly-colored, non-reticulate stalk plus its
“failure” to stain blue when bruised.

Key to Gyroporus
1. Flesh and pores quickly turning deep indigo-blue or violet-black when bruised or cut; cap
yellowish to buff, dry, often somewhat fibrillose; pores whitish becoming yellowish; stalk
colored like cap, often thicker toward base, smooth or fibrillose; common in sandy soil and
woods in eastern North America (also said to occur rarely in Pacific Northwest) G. cyanescens
1. Flesh and pores not blueing when bruised . 2
2. Cap and stalk white to yellowish or tinged pink or apricot, sometimes spotted with cinnamon or
brown in age; common under oak and pine in southeastern North America . G. subalbellus
2. Cap and stalk darker (tawny to orange-brown, chestnut-brown, wine-red, etc.) . 3
3. Cap more or less wine-red to burgundy; found under hardwoods in eastern North America
. G. purpurinus
3. Cap brown to chestnut-brown to orange-brown or tawny; widespread . G. castaneus, below

Gyroporus castaneus (Chestnut Bolete) Color Plate 148

CAP 2.5-7 (10) cm broad, convex to plane or shallowly depressed; surface dry, minutely
hairy to smooth, sometimes with a delicate whitish bloom when young; color variable:
chestnut-brown to brown, cinnamon-brown, orange-brown, or rusty-tawny. Flesh thick,
firm, white, not blueing when bruised. PORES and tubes white, becoming pale yellowish
in old age, not blueing. STALK 3-9 cm long, 0.5-1 (3) cm thick, often rather slender and
more or less equal (but a robust form also occurs), sometimes thicker or swollen below;
surface dry, uneven, brown to tawny (colored more or less like cap or slightly paler); hollow
or partially hollow at least at maturity (especially toward base). SPORE PRINT pale
yellow to yellow; spores 8-12 * 4.5-6 microns, elliptical-oblong, smooth.
HABITAT: Solitary or in groups under oaks and other hardwoods; fairly common in
Europe and eastern North America, but extremely rare in the West. I have found it only
twice in our area, under a tanoak in July (while truffle-hunting) and under a large live oak
in October. The slender form is the common one in North America.
EDIBILITY: Edible and highly esteemed in Europe, but alas, hard to come by in our area.
One day I was picking fairy ring mushrooms (Marasmius oreades) on a lawn. A round
Polish woman came out and said the mushrooms I was picking were no good. Then she
invited me inside, where she proceeded to stuff me with sour cream cookies (she thought I
was too skinny). She showed me a picture of her son, who was in the navy. Then she gave me
a necklace of dried Gyroporus castaneus. They were her last ones, she said, and they came
from “the old country, where everything tastes better.” They made a fabulous soup. S o did
the Marasmius.

COMMENTS: In the West it is unlikely to be confused with anything, as it is unlikely to

be found.
 511

BOLETUS
Medium-sized to very large, mostly terrestrial and mycorrhizal, woodland boletes. CAP fleshy,
usually dry (but sometimes viscid), sometimes areolate. PORES and tubes typically white, yellow,
orange, red, brown, or gray, often blueing when bruised. STALK fleshy, sometimes bulbous, thick
or slender, often reticulate but not glandular-dotted and without dark scabers. VEIL absent.
SPORE PRINT usually olive to olive-brown or brown, but sometimes yellow-brown or cinna¬
mon-brown. Spores spindle-shaped to elliptical, smooth.

IF YOUR bolete is not a Suillus, Fuscoboletinus, Leccinum, Tylopilus, Pulveroboletus,

Austroboletus, Boletellus, Gyroporus, Strobilomyces, or Gastroboletus, then it’s pro¬
bably a Boletus. In other words, it is much easier to characterize what a Boletus isn’t than
what it is. There is neither a veil nor glandular dots as in Suillus, the stem does not have the
dark scabers characteristic of Leccinum, the spores are neither yellow as in Gyroporus nor
reddish as in Tylopilus, nor are they ornamented as in Boletellus and Austroboletus.
As such Boletus is a large and varied genus that splits nicely into two natural groups. The
first group includes the hefty familiar forms like B. edulis, and can be subdivided into
those with white pores when young(e.g., B. edulis), those with red pores(e.g., B. satanas),
and those with yellow pores (e.g., B. appendiculatus). The stem is usually thick, some¬
times bulbous, and often reticulate, the pores small (up to 1 mm broad), and the different
species are relatively easy to distinguish in the field. They are fairly finicky as to their
mycorrhizal mates and have exceedingly fickle fruiting habits. For instance, in our area
heavy early rains may elicit a bumper crop of nearly every species, whereas if the rains
are delayed until December, they may not fruit at all!
Species belonging to the second group (the genus Xerocomus of some books) tend to be
smaller, with a dry, minutely velvety (subtomentose) cap. The pores are often large (1-2
mm or more in diameter), and the stem is relatively slender (or at least not bulbous) and
not reticulate or only very coarsely so from decurrent tube walls. M embers of this group are
more cosmopolitan than those in the first group—that is, they occur in a wider range of
habitats and may even grow on rotting wood. They tend to have a much longer growing
season and more dependable fruiting habits, but seldom appear in large numbers.
Identifying them is often difficult without resorting to chemical and microscopic
characters, but few species in the group are good edibles.
Boletus boasts some of the finest and most flavorful of all the fleshy fungi (see photo on
p. 512). First and foremost, of course, is that fabulous fungus cherished by the Europeans
above all others—the king bolete or cepe, B. edulis. Its sister species, B. barrowsii and B.
aereus, are also excellent, and the butter boletes (T?. appendiculatus and B. regius) are little-
known local treasures. On the other hand, some species are bitter-tasting(see comments on
the edibility of B. “marshii”\) and others, such as B. satanas, are so poisonous that their
consumption can result in hospitalization. A general rule of thumb is to avoid all those
species whose pores bruise blue. But this rule eliminates roughly half of all Boletus species
(including some very good ones such as B. appendiculatus), and at least on the west coast it
can be amended to: avoid all those blue-staining species with pink, red, or orange pores.
However, this does not mean that you should munch on those without red pores
indiscriminately. The edibility of some species has yet to be established and many (even B.
edulis) can cause stomach upsets if not thoroughly cooked.
Even in its modern “restricted” sense, Boletus is the major genus of boletes. It is espe¬
cially numerous and diverse (over 100 species) during the humid summer months in the
hardwood forests of eastern North America. On the west coast, however, the rains come
during the cooler months of the year and the Boletus flora is much smaller, with only
about 40 known species. Our oak-madrone woodlands boast a modest but unique
bolete bounty. Prominent species include B. satanas, B. erythropus, B. amygdalinus, B.
appendiculatus, B. regius, B. aereus, B. barrowsii, and B.flaviporus (along with Leccinum
Five delectable species of Boletus (gathered the same day!): B. regius, three at top left; B. barrowsii,
three at top right; B. aereus, three at bottom right; B. appendiculatus, small button at bottom and
the one just above it; and B. edulis, two at bottom left. The smallest bolete in this picture is about
10 cm (4 inches) high! All of these species favor hardwoods (at least in our area) except B. edulis.

manzanitae). Pine forests, on the other hand, are favored by B. edulis, which can be har¬
vested by the bushel under favorable conditions. Boletus species are easily preserved by
slicing and drying, but remember: if you should stumble into a“Boletus bonanza,” take
only as many as you can use. This will allow others (namely me) to share your luck, and
aside from the good social practice it builds, it will test your will power to its limits!
More than 30 species of Boletus occur in California. Nineteen are described here, plus
three extralimital species. Don’t be intimidated by the apparent length of the key—it is
actually composed of two separate keys that diverge at couplet #7: one key to western
species and another to eastern ones. (Since Boletus is such a prominent group in eastern
North America, I have keyed out many of the more distinctive and common species that
occur there.) If you happen to be in southern Arizona, southern New Mexico, Mexico,
or other regions where the eastern and western fungal floras overlap, try the key to western
species first; if that doesn’t work, then try the key to eastern ones (couplet #42).

Key to Boletus
1. Growing on earthballs (Scleroderma species); fruiting body small to medium-sized, usually
rather soft; especially common in southern latitudes .B. parasiticus
1. Growing on ground or wood, not on earthballs . 2
2. Taste distinctly acrid (peppery) when a small piece of the cap is chewed; fruiting body typically
rather small, with bright yellow mycelium at base of stalk .... B. piperatus & others, p. 517
2. Not as above (but taste may be bitter or sour) . 3
3. Growing on or near wood or in sawdust (especially coniferous) .4
3. Growing on ground . 7
4. Cap yellow to ochre (or developing rusty-orange tones toward center in age); pores and flesh
blueing when bruised; often in groups or clusters; spores short-elliptical . 5
4. Not as above . 6
5. Pores reddish to reddish-brown when young; cap at first powdery .B. hemichrysus
5. Not as above; pores typically yellow .B. sphaerocephalus
6. Cap and stalk dark reddish-brown to maroon-brown or chocolate-brown; cap plushlike or
fibrillose-scaly; stalk usually long and often streaked or lined; pores yellow to olive-yellow,
not blueing when bruised; found on or near northern conifers .B. mirabilis, p. 521
6. Not as above . 7

512
BOLETUS 513

7. Found in western North America (Rocky Mountains, Southwest, and westward)* .8

7. Found in eastern North America (east of the Rockies, also southern Arizona)* .42
8. Pore surface red, pink, vinaceous, or orange when fresh(but may be yellowish or dingier in age,
or in one case yellow when young becoming red in age); pore surface and flesh usually turning
blue or blue-black when bruised or cut .9
8. Pore surface yellow to olive, brown, white, etc., but not red or orange; pore surface may or
may not turn blue when bruised . 15
9. Stalk distinctly reticulate (netted), at least over upper portion . 10
9. Stalk not reticulate, or only very slightly so from decurrent tube walls. 13
10. Stalk equal to bulbous, usually at least 2.5 cm thick at apex; reticulation on stalk usually red
or pink or vinaceous (but may be stained olive or brown by spores); widespread in West 11
10. Stalk equal or slightly thicker below, typically 1 -3 cm thick at apex; reticulation red or not red;
found only in parts of the Southwest (e.g., southern Arizona) .46
11. Stalk with an exaggerated basal bulb; cap whitish to grayish to olive-buff, or sometimes suf¬
fused with pink in age; associated with oak . B. satanas, p. 527
11. Stalk equal or swollen below, but without an exaggerated bulb; cap some shade of brown or red
(or in one variety, pallid); associated mainly if not exclusively with conifers . 12
12. Cap brown to olive-brown or yellow-brown (not often reddish); pores often yellow when very
young; found in mountains . B. haematinus (see B. pulcherrimus, p. 528)
12. Cap brown to reddish-brown; pores never yellow .B. pulcherrimus & others, p. 528
13. Cap intensely yellow, quite large; rare .B. orovillus(see B. erythropus, p. 526)
13. Not as above; common . 14
14. Flesh changing to blue slowly or not at all when mushroom is cut open; cap usually areolate in
age; not common .B. mendocinensis(see B. chrysenteron, p. 519)
14. Flesh blueing quickly when cut; cap not normally areolate .. . B. erythropus & others, p. 526
15. Stalk typically long, relatively slender, and scurfy or minutely scaly (as in Leccinum); cap yellow
to ochre, reddish-orange, or cinnamon; pores yellow, not normally blueing when bruised;
rare (reported from northern Idaho and the Southwest) .67
15. Not with above features; common. 16
16. Cap viscid or slimy when moist, cinnamon-brown to chestnut- or reddish-brown; pores an
intense, brilliant yellow or greenish-yellow, not blueing when bruised; stalk not reticulate or
only slightly so (but sometimes ridged) ...B. flaviporus, p. 522
16. Not as above; cap not viscid, or if viscid then not as above . 17
17. Stalk entirely or partially pink to red to dark red (or with red flakes or granules) . 18
17. Stalk white to buff, yellow, or brown (i.e., lacking obvious red tones, but may have reddish
to cinnamon-brown stains, especially near or at base) . 27
18. Cap red to purple-red, pink, or at times reddish-brown (but sometimes overlaid at first with
olive-yellow to olive-gray to olive-brown hairs or fibrils) . 19
18. Cap not markedly reddish (but may be dark brown or black with a slight reddish cast) ... 22
19. Cap small(2-5 cm broad), red todark red to purple-red; stalk less than 1 cm thick; poresand flesh
typically not blueing when bruised; rare .B. coccyginus (see B. bicolor, p. 521)
19. Not as above .20
20. Cap and stalk red or reddish; known only from the Southwest . . . B. bicolor & others, p. 521
20. Not as above; common in the Pacific Northwest and California, but not in the Southwest; stalk
usually only partially red .21
21. Associated with live oak in central and southern California .B. dryophilus, p. 520
21. Associated with northern conifers .B. smithii(see B. dryophilus, p. 520)
22. Taste distinctly bitter (chew on a small piece of the cap), even when cooked; stalk at least 2 cm
thick at apex . 23
22. Not as above (but taste may be acidic or sour) .24
23. Upper part of stalk typically reticulate (often finely so) .B. calopus, p. 523
23. Stalk not reticulate or only obscurely so at the very apex .B. rubripes, p. 524
24. Cap black to blackish-brown to deep olive-gray when fresh, often developing a slight reddish
cast toward the margin (especially when wet) and sometimes fading in age; surface of cap not
typically areolate, but occasionally becoming so in age .B. zelleri, p. 518
24. Cap dark olive-brown to olive-gray to brownish, tan, etc.; surface often areolate .25

*In regions where the eastern and western fungal floras overlap (e.g., southern Arizona), try both choices!
514 BOLETACEAE

25. Fruiting body medium-sized to large (cap 6-18 cm broad); flesh in base of stalk not normally
reddish; spores not truncate; found under mountain conifers .B.fragrans
25. Not as above; fruiting body small to medium-sized, or if larger than more common along the
coast and flesh in base of stalk often reddish; spores truncate or not truncate; common . 26
26. Cap when areolate usually showing pink or reddish tints in the cracks; stalk typically rather
slender (less than 2 cm thick); cap medium-sized (usually less than 9 cm broad); spores
not truncate .B. chrysenteron, p. 519
26. Not as above; cap when areolate may or may not show pink or reddish tints; size and shape
of fruiting body variable, but sometimes more robust than above species (stalk 1 -3 cm thick,
cap 5-15 cm broad); spores often amyloid or truncate (appearing chopped off at one end) when
viewed under the microscope .B. truncatus & others (see B. chrysenteron, p. 519)
27. Fruiting body medium-sized to large; stalk 2 cm thick or more, often bulbous or thicker below
(but often not); pores small (mostly less than 1 mm broad) . 28
27. Fruiting body small to medium-sized; stalk usually 2 cm thick or less at apex, or if thicker
then pores at least 1 mm broad; stalk usually equal or tapered below (not bulbous) .38
28. Pore surface whitish when young, becoming yellowish, greenish, or brown inage and not blueing
when bruised; stalk distinctly reticulate (netted) at least over upper portion; flesh white or
tinged reddish, not blueing (or sometimes blueing slightly when tubes are peeled off) ... 29
28. Not as above; if stalk reticulate then pores not white when young and/ or pores blueing when
bruised . 31
29. Cap white to pale grayish or dingy buff, usually dry; associated mainly with oak in central
California, with pine and other conifers in the Southwest .B. barrowsii, p. 529
29. Not as above; cap typically darker (but may be whitish while still under the duff) .30
30. Cap dark brown to blackish-brown beneath a thin whitish bloom when young(but often fading
to cinnamon-brown or paler in age); associated with hardwoods . B. aereus, p. 531
30. Not as above; cap biscuit-brown to yellow-brown, cinnamon-brown, brown, or dark red;
associated mainly with conifers but also with oak and other trees . B. edulis & others, p. 530
31. Taste distinctly bitter (chew on a small piece of the cap), even when cooked; stalk typically at
least 2 cm thick at apex . 32
31. Not as above (but may taste slightly sour or acidic) . 33
32. Found under northern conifers; stalk usually reticulate B. coniferarum(see B. calopus, p.523)
32. Found mostly with oak in California; stalk not reticulate .B. “marshii, ”p. 524
33. Stalk distinctly reticulate, at least over upper portion. 34
33. Stalk typically not reticulate; found under mountain conifers .B.fragrans
34. Stalk yellow to buff (at least when fresh), sometimes with reddish stains below .35
34. Stalk brown, or white with brown fibrils (apex sometimes yellow) .B. fibrillosus, p. 523
35. Cap dark brown when fresh; pores and flesh not blueing when bruised .
.B. sp. (unidentified) (see B. fibrillosus, p. 523)
35. Not as above; pores normally turning blue when bruised (but may not blue in button stage);
flesh in very base of stalk often tinged pinkish or vinaceous; cap pink, red, brown, tan, or
yellowish, but not normally dark brown. 36
36. Cap with distinct fibrillose scales, especially in age; associated with mountain conifers (parti¬
cularly fir) . B. abieticola(see B. appendiculatus, p. 525)
36. Associated with hardwoods, or if with conifers then cap lacking obvious scales .37
37. Cap usually brown to reddish-brown, sometimes yellowish .B. appendiculatus, p. 525
37. Cap reddish to pink or rose-colored, sometimes with yellow .B. regius, p. 526
38. Flesh, stalk, and pores staining dark blue almost instantly when exposed; cap dull brown to
dark brown; stalk usually yellow at apex and brown or reddish-brown at base; pores yellow to
olive; rare in California, more common in the Pacific Northwest .B. pulverulentus
38. Not as above . 39
39. Spore print amber-brown to bright yellow-brown; cap vinaceous-brown to dark brown, yellow-
brown, tan, etc., often with paler spots; pores whitish to buff or pale tan; reported from the
Southwest (but mainly eastern in distribution) .B. affinis
39. Not as above; spore print typically brown to olive or olive-brown .40
BOLETUS 515

40. Cap brown to dark brown or blackish; stalk pallid to buff (or yellowish at apex); pores brilliant
yellow, not blueing when bruised .B. citriniporus (see B. fibrillosus, p. 523)
40. Not as above; pores typically yellow but not as brilliantly colored as above .41
41. Cap usually extensively fissured (areolate) by maturity . . . . B. chrysenteron & others, p. 519
41. Cap not usually areolate, but sometimes showing a few cracks in age, especially near margin
.B. subtomentosus & others, p. 517
42. Pore surface dark red to bright red, orange, orange-brown, red-brown, or dark yellow-brown
when fresh (may fade to yellow in age) .43
42. Pore surface white to yellow, olive, grayish, etc. when fresh (but sometimes brown in age) 49
43. Pore surface reddish-brown to dark yellow-brown or orange-brown when young, or if not then
fruiting body rather small and not blueing when bruised . 73
43. Pore surface dark red to red, brick-red, or orange; medium-sized to large, usually blueing 44
44. Cap whitish to buff, grayish-olive, or sometimes tinged pink; stalk neither bulbous nor mar¬
kedly reticulate; found under southern hardwoods .B. piedmontensis
44. Not as above; cap typically darker or brighter than above .45
45. Stalk typically reticulate, at least over upper portion .46
45. Stalk not reticulate or sometimes slightly so at apex, but often scurfy from numerous minute
granules or dots .48
46. Cap yellow to olive-brown, brown, or reddish-brown B. luridus{ see B. pulcherrimus, p. 528)
46. Cap deep red to bright red to rosy-red, at least when fresh .47
47. Cap viscid when moist; pores dark red, sometimes with yellow droplets; stalk very coarsely
reticulate nearly to the base .B.frostii, p. 528
47. Not as above; cap not viscid or only slightly so; stalk only finely reticulate .
.B. flammans & B. rubroflammeus (see B.frostii, p. 528)
48. Cap red or rosy-red . B. bicolor var. borealis (see B. bicolor, p. 521)
48. Cap yellowish to brown or reddish-brown .
.B. erythropus, B. subvelutipes, & others (see B. erythropus, p. 526)
49. Cap white or pallid, at least until old age . 50
49. Cap more highly or deeply colored, at least when young . 51
50. Stalk white or whitish, usually not reticulate (but sometimes slightly so); very common under
hardwoods; taste mild to bitter .B. pallidus
50. Stalk at least partially red or with bright red to pink reticulation; taste bitter .... B. inedulis
51. Pore surface and/ or flesh typically staining blue, blue-green, or blue-black when bruised or
cut (but sometimes slowly or weakly) . 52
51. Neither the pore surface nor the flesh blueing when bruised .60
52. Fresh fruiting body bright yellow to lemon-yellow (but cap may be yellow-brown at center or
streaked with red), quickly staining blue or blue-black when bruised or cut; stalk sometimes
reticulate but usually not . B. pseudo sulphur eus
52. Not as above . 53
53. Cap bright red to rose, pink, or rusty-red when fresh (may fade in age) .54
53. Not as above (but cap may show slight red or pinkish tints) . 56
54. Stalk slender (usually less than 1 cm thick) .B. rubellus& others (see B. bicolor, p. 521)
54. Stalk usually 0.8-3 cm or more thick . 55
55. Stalk distinctly reticulate, at least over upper portion.B. speciosus(see B. regius, p. 526)
55. Stalk not reticulate or only slightly so from decurrent tube walls . B. bicolor Si others, p. 521
56. Taste decidedly bitter (chew on a small piece of the cap), even when cooked; stalk typically
at least 2 cm thick; not common . B. calopus, p. 523
56. Not as above (but taste may be slightly acidic or sour) . 57
57. Flesh, stalk, and pores staining dark blue almost instantly when exposed; cap dull brown to
dark brown; stalk usually yellow at apex and brown or reddish-brown toward base; pores
fairly small, yellow to olive-yellow; stalk typically not reticulate .B. pulverulentus
57. Not as above (but may stain blue to blue-green) . 58
58. Cap typically areolate (extensively fissured or cracked) at maturity; stalk entirely or partially
red to dark red (or with red fibrils or granules) . 59
58. Not as above; cap not areolate or only occasionally so andjor stalk lacking red .66
516 BOLETACEAE

59. Stalk scurfy from small rough scales; spores striate under the microscope; often growing at
the bases of trees .(see BoletellusSc A ustroboletus, p. 508)
59. Not as above; spores not striate; usually found on ground . . B. chrysenteron & others, p. 519

60. Cap rosy or pink when fresh (but often fading to tan in age); stalk not reticulate; stalk base bright
yellow inside and out; pores whitish when young .(see Tylopilus, p. 532)
60. Not as above .*.61
61. Pore surface white to grayish when young(but usually yellowish, brownish, grayish, or greenish
in age); stalk distinctly reticulate (netted), at least over upper portion; flesh white or tinged
reddish; spore print olive to brown but not yellow-brown .62
61. Not as above; pores differently colored and/ or stalk not reticulate or only faintly so at apex 64
62. Cap grayish (or with darker fibrils) when fresh; pores white becoming gray to grayish-brown
in age; common under hardwoods .B. griseus(see B. ornatipes, p. 522)
62. Not with above features; pores not typically grayish in age .63
63. Cap tan to gray-brown, dark brown, or yellow-brown, dry, often areolate in age; cap and stalk
without purplish tones; associated with hardwoods.B. variipes (see B. aereus, p. 531)
63. Not with above combination of features (but may have some of them); found with hardwoods or
conifers .B. edulis & others, p. 530
64. Cap and stalk bright orange-brown to golden-orange; fairly common in North Carolina and
adjacent areas, but exact distribution uncertain . .B. auriflammeus (see B. ornatipes, p. 522)
64. Not as above .65
65. Cap and stalk viscid or slimy and yellow, or if not then pores brilliant yellow to bright greenish-
yellow; not large.75
65. Not as above (pores often yellow but not intensely so).66
66. Stalk typically long, relatively slender, and scurfy or minutely scaly as in Leccinum; cap yellow
to ochre, reddish-orange, or cinnamon; pores yellow, not normally blueing when bruised 67
66. Not with above features . 68
67. Dots or minute scales on stalk reddish; cap usually viscid (sometimes thinly so) when moist
.B. longicurvipes & B. rubropunctus
67. Not as above; stalk yellow .B. subglabripes
68. Stalk largely yellow to buff when fresh (but may develop brownish to cinnamon stains from
the base upward, especially in age) . 69
68. Stalk typically brownish to reddish, tan, etc., or white with brown tones, even when fresh 71
69. Stalk prominently reticulate throughout or nearly throughout; pores small (about 2 per mm);
taste mild or bitter; found under hardwoods .B. ornatipes & others, p. 522
69. Not as above; stalk not reticulate and/ or pores larger; taste usually mild .70
70. Cap often with brownish to reddish hairs or small scales on a yellowish to orange-buff back¬
ground, usually viscid beneath hairs when wet (hairs may be absent in age) (seeSuillus, p. 491)
70. Not as above .B. subtomentosus & others, p. 517
71. Spore print yellow-brown or amber-brown; cap yellow-brown to reddish-brown or dark brown,
often with paler spots; pores white to buff or pale tan or yellow .B. affinis
71. Not as above; spore print olive to brown or olive-brown; pores yellow to olive . 72
72. Stalk with prominent raised ridges and/ or coarsely reticulate, often long and slender in relation
to cap; cap grayish or olive-tinged when young but usually reddish-brown to dark reddish-
brown or bay-red in age; pores not blueing .B. projectellus (see B. mirabilis, p. 521)
72. Not as above; cap slightly viscid when young or wet, usually reddish-brown to bay-red to
chestnut-brown, but at times yellow-brown or olive-tinged; pores may or may not stain blue
when bruised; stalk not coarsely reticulate or strongly ridged B. badius(see B. zelleri, p. 518)
73. Pores and/ or flesh not blueing when bruised; under hardwoods or conifers . . . B. rubinellus
73. Pores and/ or flesh staining blue to blue-black when cut; usually under hardwoods .74
74. Pore surface dark reddish-brown to orange-brown when young; stalk typically not reticulate
.B. vermiculosus & others
74. Pore surface dark yellow-brown when fresh; stalk often reticulate .B.fagicola
75. Fruiting body mostly bright yellow to lemon-yellow (including cap); cap and stalk viscid or slimy;
stalk usually with a white cottony base; common in South .B. curtisii
75. Not as above; pores brilliant yellow when fresh; cap and stalk viscid or dry.
.B. viridiflavus & B. auriporus (see B. flaviporus, p. 522)
Boletuspiperatus is our smallest species of Boletus. It has a peppery taste and bright yellow mycelium
at the base of the stalk.

Boletus piperatus (Peppery Bolete)

CAP 2-8 cm broad (but usually less than 5 cm), convex to plane; surface slightly viscid to
dry, usually smooth, yellow-brown to buff, rusty-cinnamon, orange-brown, or reddish-
brown. Flesh thin, yellowish-buff to pinkish in cap, bright yellow in base of stalk; taste
distinctly acrid (peppery). PORES rather large (1-2 mm in diameter), yellow-brown to
cinnamon, reddish-brown, coppery or brick-red, darkening slightly when bruised but not
blueing; tubes tawny to reddish-yellow. STALK 2-8 (12) cm long, 0.4-1 (2) cm thick,
equal or tapered downward, often slender; smooth, solid, colored more or less like the
cap; base coated with bright yellow mycelium. SPORE PRINT brown to dull cinnamon;
spores 8-12 * 3-5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary to scattered or gregarious on ground in woods, associated mainly
with conifers; widely distributed. In our area it fruits in the fall and winter with pine and
Douglas-fir, but is not common (I find it only once or twice each year). Farther north,
however, it is sometimes abundant.

EDIBILITY: Questionable. It’s not as bitter as some of the peppery R ussula and Lactarius
species, and therefore could be useful as a spice. However, according to one source the
peppery taste disappears when thoroughly cooked, and according to another it is mildly
poisonous unless thoroughly cooked.
COMMENTS: Our smallest Boletus, this species is suggestive of a Suillus (a.nd was once
placed in that genus), but has neither glandular dots nor a veil. Some view it as a possible
“missing link” between Suillus and Boletus. If you have any doubt as to its identity, just
chew a small piece of the cap for a couple minutes! The bright yellow mycelium and yellow
flesh at the base of the stalk are also characteristic. B. piperatoides is a similar widespread
peppery species with blue-staining pores.

Boletus subtomentosus (Boring Brown Bolete)

CAP 5-15 (20) cm broad, convex to plane; surface dry, minutely velvety (subtomentose),
but may appear smooth in age; yellow-brown to drab olive-brown to dull brown, or
occasionally yellowish, but in age or especially in wet weather often redder (cinnamon-

517
Boletus subtomentosus is a boring brown and yellow bolete with large yellow pores (see p. 488 for a
close-up of the pores).

brown); sometimes with pallid, yellow, or reddish-tinged cracks, especially near margin.
Flesh pallid to pale yellow, sometimes blueing slightly when exposed. PORES large (1-3
mm in diameter), dull yellow to bright yellow, blueing weakly or not at all when bruised.
STALK 4-14 cm long, 1-2 (3) cm thick, equal or tapered either way, smooth or scurfy, the
apex often coarsely reticulate from downward-extending tube walls; firm, yellow to buff,
or often stained brown to dull cinnamon (especially below), but never red. SPORE
PRINT olive-brown; spores 10-16 * 3.5-5 microns, elliptical to spindle-shaped, smooth.
HABITAT: Solitary to widely scattered or in small groups on ground in woods; widely
distributed and very common, but not often occurring in large numbers. In our area
it fruits throughout the mushroom season with a variety of tree hosts (oak, conifers, etc.).
EDIBILITY: Edible, but definitely not choice. In blind tastings of local boletes it has
consistently placed near the bottom.
COMMENTS: This boring but ubiquitous bolete with the boring brown cap does not
show red on the stem, in contrast to B. zelleri and B. chrysenteron. In addition, the cap
is not often as conspicuously areolate as that of the latter species, and the pores are larger.
Xerocomus subtomentosus is a synonym. B. spadiceus is an equally boring but ubiqitous
bolete. It has the same general appearance and grows in the same places at about the same
time. True, its cap may be more reddish-brown and its pores may blue more readily, but
the only way to separate it with certainty is by applying a drop of ammonium hydroxide
to the cap surface. A fleeting blue-green to blue-black reaction means B. spadiceus;
otherwise, it’s B subtomentosus. Since neither is wortheating, thedistinctionisacademic.
Inundating boring brown boletes with ammonia may be your idea of fun, but it’s not
mine!

Boletus zelleri (Zeller’s Bolete) Color Plate 128

CAP 3-16 cm broad, convex to plane; surface dry, often wrinkled or uneven and with a
frosted or finely powdered appearance when young; black to dark gray or dark olive-brown
when fresh, often reddening somewhat in age or wet weather, especially toward the
margin; surface cracking only slightly or not at all. Flesh thick, white to pale yellow, some¬
times blueing erratically when exposed. PORES and tubes yellow to dark yellow or olive-

518
BOLETUS 519

yellow, often blueing when bruised (but often not blueing). STALK 4-12 cm long,0.5-3 (5)
cm thick, equal or slightly thicker at either end; firm, yellow to tan with delicate red
granules when young, usually dark red above or throughout in age. SPORE PRINT
olive-brown; spores 12-16 * 4-5.5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary, scattered, or in groups on ground or rotten wood; very common
along the Pacific Coast in woods of all kinds. In our area it fruits in the fall, winter, and
early spring and is one of the few boletes with a tolerance for redwood (I have even found
it growing on a redwood stump!). It is also abundant under oak and farther north, under
alder.
EDIBILITY: Edible and highly rated by some sources. In my experience, however, it
cooks up slimy and tasteless.
COMMENTS: The sensational combination of black, yellow, red, and often blue makes
this our most colorful bolete. The color plate shows specimens whose caps have begun to
develop a reddish tinge, but perfectly fresh caps can be coal-black or deep gray. B. chry-
senteron and B. truncatus often have a more copiously cracked (areolate) cap that is
usually paler or duller in color. B. citriniporus has a dark cap, but its stem is not red.
In eastern North America, B. zelleri is supplanted by B. badius, a common edible species
with a dark yellow-brown to bay-red cap. Like B. zelleri, it grows on rotten wood (as well
as on the ground), usually under conifers.

Boletus chrysenteron (Cracked-Cap Bolete)

CAP 3-11 cm broad, convex to plane; surface dry, minutely velvety when young, in age
usually conspicuously cracked or fissured (areolate), especially toward the margin;
color variable: dark olive-brown to dark grayish-olive, grayish-brown, or brown; often
paler (tan to olive-buff) in age, with pink to reddish tints usually visible in at least some of
the cracks (especially those toward the margin). Flesh fairly thick, whitish to yellow,
usually blueing slowly when exposed. PORES rather large (about 1 mm in diameter),
yellow to greenish-yellow or sometimes dingy brownish, usually (but not always) bruising
blue or greenish (sometimes slowly!); tubes also yellow. STALK 4-13 cm long, 0.5-1.5 cm
thick, equal or tapering downward, smooth or minutely scurfy and often longitudinally
ridged or striate, firm; color variable, but typically a mixture of yellow and red (yellow

Boletus chrysenteron is one of several species that has a conspicuously cracked (areolate) cap at
maturity. Others include B. truncatus, B. porosporus, and Tylopilus amylosporus (not illustrated).
520 BOLETACEAE

with reddish fibrils or yellow above and red to dark rhubarb-red below). SPORE PRINT
olive-brown; spores 10-15 * 3.5-6 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary or in small groups under trees and in wooded areas, often near trails
or on roadbanks; widely distributed and ubiquitous, but rarely fruiting in large numbers.
In our area it can be found throughout the mushroom season.
EDIBILITY: Edible, but not choice. It is rather mushy and insipid when cooked.
COMMENTS: The conspicuously fissured (areolate) cap with pinkish-tinted cracks
plus the yellow pores that usually bruise blue are the hallmarks of this cosmopolitan
bolete. Also known asXerocomus chrysenteron, it is sometimes confused with B. zelleri,
which has a darker cap that is not usually areolate. There are several very similar“cracked-
cap” boletes that are difficult to distinguish in the field, including: Tylopilus amylosporus,
with dark reddish-brown, erratically amyloid spores and pallid (not pink or red) flesh in
the cracks; B. mendocinensis, with pinkish to reddish pores; B. truncatus, with truncate
(“chopped off’) spores; and B. porosporus, with truncate spores and no pinkish tints
in the cracks. The latter two species are particularly widespread and common, and seem to
intergrade. A robust form (cap 5-15 cm broad, stalk 1-3.5 cm thick) of B. truncatus bears
special mention because it occurs commonly in our coastal forests. Its cap is often only
slightly areolate and shows little or no pink in the cracks. Since none of the above species
are worth eating, it hardly matters whether or not you identify them correctly, unless
you aspire to be a professional boletologist (or plan on entertaining a professional bole-
tologist). All of them, along with B. zelleri and B. dryophilus, are prone to attack by a
powdery white to bright yellow parasitic fungus (Hypomyces chrysospermum) which
eventually engulfs the entire fruiting body, making it look like a very sick puffball.

Boletus dryophilus (Oak-Loving Bolete) Color Plate 132

CAP 3-10 cm broad, convex to plane or somewhat irregular; surface dry, minutely velvety
when young, often cracked or fissured in age, usually reddish to reddish-brown or pinkish,
but sometimes overlaid with minute olive-brown to olive-gray hairs. Flesh thick, yellow,
usually blueing when exposed. PORES often somewhat irregularly shaped, yellow to
olive-yellow and usually blueing when bruised; tubes same color. ST ALK 3-8 (12) cm long,
1 -2 (2.5) cm thick, often rather short and stout, usually pinched or narrowed at the base and
often slightly swollen above it; sometimes nearly equal; firm, solid, usually distinctly
yellow at apex and red to dark reddish below. SPORE PRINT brown to olive-brown;
spores 12-16 * 5.5-8 microns, spindle-shaped to elliptical, smooth.
HABIT AT: S olitary or in groups in humus under live oak, fall through early spring, known
only from California. It is rather rare in our area, but sometimes abundant in southern
California. In fact, it is one of the most common boletes in Los Angeles and San Diego
counties.
EDIBILITY: Edible and fairly popular in the above-mentioned region. Collections I have
sampled were mediocre, and in our area they are more readily parasitized by Hypomyces
chrysospermum than any other bolete!
COMMENTS: This interesting endemic species is closely related to B. chrysenteron, but
is somewhat stouter and apparently grows only with oak. The reddish tones usually present
in the cap plus the distinct yellow and red zones on the stalk help identify it. The abruptly
tapered or “pinched” stem base is also distinctive when present. Other species: B. smithii
has a similarly colored cap, but is larger and stouter. Its stalk may be narrowed at the base
but is usually pallid or yellow with a bright red band near or at the apex. It seems to grow
only with conifers and is quite common in northern California and the Pacific N orthwest.
BOLETUS 521

Boletus bicolor (Red and Yellow Bolete)

CAP 5-15 cm broad, convex to plane or somewhat irregular; surface dry, smooth or
minutely velvety, sometimes cracking in age; deep red to pinkish-red or rose, the margin
sometimes yellowish in age. Flesh thick, pale yellow, blueing erratically or not at all when
exposed. PORES yellow to bright yellow, sometimes with reddish areas in old age;
typically staining blue or blue-green slowly when bruised; tubes also yellow. STALK 5-10
cm long, 1-3 cm thick, equal or thicker below or with a tapered base; smooth, firm, solid,
not reticulate or only slightly so at apex; deep red to rosy except for yellow apex. SPORE
PRINT olive-brown; spores 8-12 * 3.5-5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Scattered to gregarious on ground in woods, associated mainly with oak and
perhaps aspen; common in the summer and early fall in eastern North America and also
occurring in the Southwest, but apparently not found on the west coast. A related species,
B. coccyginus (see comments), occurs in California and the Pacific Northwest, but is rare.
EDIBILITY: Edible, and according to some, excellent. However, gastrointestinal upsets
have been attributed to closely related species, so exercise caution.
COMMENTS: The rosy to red cap and stalk contrast vividly with the yellow pores, making
this a most beautiful bolete. There are a number of similar red-capped, yellow-pored,
blue-staining species that form a close-knit group within Boletus. They are difficult to
distinguish in the field and are largely confined to the humid forests of easternand southern
North America and the tropics. They include: B. sensibilis, somewhat larger, with a brick-
red cap and yellow stalk; B. miniato-pallescens, cap rosy-red fading to yellow and stalk
yellow to reddish-brown, common; and several smaller, slimmer (stalk less than 1 cm
thick) species such as B. rubellus, widespread, B. campestris, especially common on
shaded lawns, and B.fraternus, with an areolate cap at maturity. All of these species bruise
blue (often quickly) and do not have the robust, reticulate stalk of B. regius. Other species:
B. coccyginus is a rare, small, slender western species with a red to rose or purple-red cap
and yellow pores and flesh that do not stain blue; B. bicolor var. borealis resembles the
typical variety but has red to orange pores; it occurs mainly in the Southeast.

Boletus mirabilis (Admirable Bolete) Color Plates 127,129

CAP 5-15 (20) cm broad, convex to plane; surface moist to dry, granulose to plushlike
(roughened by numerous small, often erect, fibrillose scales); dark reddish-brown to
maroon-brown, bay-brown, or chocolate-brown; margin often hung with fragments of
tissue. Flesh thick, white to dingy pinkish or yellow, rarely blueing when bruised. PORES
fairly large (1-2 mm in diameter), pale yellow becoming mustard-yellow and finally
greenish-yellow, not blueing when bruised but sometimes staining darker yellow; tubes
also yellow. STALK 7-20 cm long, 1-3.5 cm thick, usually club-shaped (thicker toward
base); dark brown to maroon-brown or reddish-brown, sometimes with yellow, buff, or
beige streaks; firm and often roughened, pitted, or longitudinally ridged, the apex often
coarsely reticulate; base frequently with yellow mycelium. SPORE PRINT olive-brown;
spores 18-24 * 7-9 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary or in small groups on or near rotting conifers (especially hemlock),
but sometimes appearing terrestrial; common in the fall in the Pacific Northwest and
northern California, also reported (rarely) from Michigan. It does not seem to venture
south of San Francisco. (Neither does hemlock, and neither would I if I could help it.)
EDIBILITY: Edible and delicious—some collections have a distinct lemony flavor.
Don’t use specimens attacked by a whitish mold (Hypomyces chyrospermum).
COMMENTS: This beautiful northern bolete is practically unmistakable. The plush¬
like maroon-brown cap and long roughened, similarly-colored stalk plus the yellow
522 BOLETACEAE

pores that do not stain blue and growth on or near rotting conifers form a unique set of
characters. Other species: B. projectellus of eastern North America is a closely related,
terrestrial species that favors sandy situations under pine.

Boletusflaviporus (Viscid Boletus) Color Plate 131

CAP 4-15 cm broad, convex to plane; surface viscid or slimy when wet, smooth or fibrillose,
reddish-brown to cinnamon-brown or chestnut-brown, not normally cracking into scales.
Flesh thick, white to pale pinkish, not blueing when bruised. PORES and tubes brilliantly,
intensely yellow, becoming slightly greenish-yellow in age; not bruising blue. ST ALK 6-15
cm long, 0.6-2 (3) cm thick, equal to slightly thicker below or often with a short, tapered,
rooting base; smooth, viscid in wet weather but otherwise dry; color variable, but usually
yeilow at apex and pallid to reddish-brown or dark brown below; typically not reticulate
or only slightly so at apex. SPORE PRINT dark olive-brown; spores 11-17 x 4-6 microns,
spindle-shaped to elliptical, smooth.
HABITAT: Solitary, scattered, or in small groups in mixed woods and under hardwoods;
endemic to the west coast. In our area it is fairly common in the fall and winter under oak,
madrone, and manzanita, but seldom fruits in large numbers.
EDIBILITY: Edible, but mediocre.
COMMENTS: This is our only Boletus with a distinctly viscid cap and intensely yellow
pores that do not stain blue. It might be mistaken for a Suillus, but it isn’t associated with
conifers and has neither glandular dots nor a veil. The stem is not truly reticulate like that of
B. edulis and B.fibrillosus, but the tubes may form a slight raised network at the stalk apex.
Its eastern counterparts with brilliant yellow, non-blueing pores are B. viridiflavus,
with a reddish- or pinkish-tinged stalk and sometimes viscid cap; and B. auriporus
{-B. caespitosus?), with a yellower stalk and somewhat more velvety cap.

Boletus ornatipes (Ornate-Stalked Bolete)

CAP 4-20 cm broad, convex to nearly plane; surface dull, often velvety when young, dry
or slightly viscid when wet, color variable: gray, purple-gray, olive, olive-brown, yellow,
or mixtures thereof. Flesh firm, thick, yellow, not blueing; taste mild to somewhat
bitter. PORES and tubes lemon-yellow to bright yellow, usually staining orange-
yellow to rusty-brown when bruised, but not blueing; pores small. STALK 7-15 cm long,
1-3.5 cm thick, more or less equal but often curved, solid, firm, bright yellow to yellow-
orange throughout and prominently reticulate nearly to base (reticulation often coarse);
dingier in age. SPORE PRINT brown to olive-brown; spores 9-13 * 3-4 microns,
spindle-shaped to elliptical, smooth.
HABITAT: Solitary, scattered, gregarious or in small clusters on ground under oaks
and other hardwoods in eastern North America (particularly the northern half); it is
common in the summer and early fall.
EDIBILITY: Non-bitter specimens are said to be edible; I haven’t tried it.
COMMENTS: This beautiful eastern bolete is readily told by its yellow pores and flesh
and its beautifully patterned (reticulate) golden-yellow stalk. The cap color varies consi¬
derably but is usually some shade of gray, olive, or yellow-olive. B. retipes is a bitter¬
tasting southern version (probably a form of the same species); B. auriflammeus, also
southern, is similar but has a brownish-orange to brilliant gold cap and creamy or pink-
tinged flesh. All of these stain your fingers yellow when handled. Another closely related
species, B. griseus, is grayish but often shows some yellow (at least at the base of the stalk)
as it ages. It has softer, mild-tasting, pallid, edible (but often wormy!) flesh, white to gray
or grayish-brown (never yellow!) pores that may stain brown, and a reticulate stalk. It is
common under hardwoods in eastern North America and also occurs in southern Arizona.
BOLETUS 523

Boletus fibrillosus
CAP 6-17 cm broad, convex to more or less plane; surface dry, minutely velvety becoming
fibrillose or sometimes fibrillose-scaly at the center; brown to dark brown to cinnamon-
brown, sometimes with paler blotches or paler at the margin (but not yellow). Flesh thick,
white or buff, not blueing when bruised; taste mild. PORES and tubes pale yellow to
yellow or dingy olive-yellow (occasionally pallid when very young), not bruising blue.
ST ALK 8-16 cm long, 2 A cm thick at apex, equal or thicker below(but sometimes pinched
at base); solid, firm; light brown to brown(often paler than cap); apexoftenyellowand/ or
base whitish; usually lined or fibrillose, reticulate at least at the top. SPORE PRINT
dark olive-brown; spores 13-17.5 x 3.5-5.5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary to gregarious on ground in mixed woods and under conifers;
known only from the west coast, sometimes common in the fall in coastal northern
California and the Cascades. A similar species (see below) occurs in our area.
EDIBILITY: Edible and fairly good, but certainly not the equal of B. edulis or B. aereus.
COMMENTS: The dark brown frequently fibrillose cap, yellow pores, brown reticulate
stalk, and “failure” of all parts to turn blue when bruised are the telltale traits of this
northern bolete (see photo on p. 529). It resembles B. edulis and B. aereus, but its cap is
more fibrillose and its pores are often yellow even when young. Tylopilus pseudoscaber
is also somewhat similar, but does not have yellow pores. Both T. pseudoscaber and B.
fibrillosus have passed under the defunct name B. olivaceobrunneus. A similar, uniden¬
tified Boletus occurs rarely in our area under live oak, but differs in having a yellow stalk.
B. citriniporus is a smaller Californian species with a brown to blackish cap, brilliant
yellow pores that do not bruise blue, a whitish to buff (or yellow at apex) stalk that is not
reticulate or only slightly so, and mild taste. I have found it in mixed woods and under oak.

Boletus calopus (Bitter Bolete)

CAP 10-30 cm broad, convex to plane or somewhat irregular; surface dry, dull, smooth to
minutely hairy or fibrillose, often cracking (areolate) in age; dull brown to olive-brown,
grayish-brown, or olive-buff, sometimes yellow-brown or in age darker brown. Flesh very
thick, pale yellow or whitish, quickly blueing when exposed; taste distinctly and
persistently bitter. PORES and tubespaleyellowbecomingdarkerordingieryellowinage;
typically bruising blue or blue-green quickly. STALK 6-15 (20) cm long, 3-7 cm thick at
apex, equal or bulbous; solid, firm; yellow, but usually with pink to red zones or
discolorations also present, or sometimes entirely reddish; blueing when bruised; apex or
upper half finely reticulate. SPORE PRINT dark olive-brown; spores 13-19 x4-6 microns,
spindle-shaped to elliptical, smooth.
HABITAT: Solitary, scattered, or in groups onground in mixed woodsand underconifers
in the late summer and fall (or occasionally spring); common in western North America,
rare in the East. It is particularly prominent at higher elevations (e.g., in the Sierra
Nevada), but 1 have also seen large fruitings on the Olympic Peninsula in Washington.
In our area it is apparently supplanted by another bitter bolete, B. “marshii.”
EDIBILITY: Bitter-tasting (see comments on the edibility of B. “marshii”).
COMMENTS: This sometimes massive bolete is easily recognized by its bitter taste,
blueing of all parts when injured, yellow pores, and reticulate stalk. The bitter taste always
comes as a disappointment to those seduced by its large size. Our other large “bitter
boletes,” B. “marshii” and B. rubripes, have non-reticulate stalks. Still another bitter
bolete, B. coniferarum, has a yellow reticulate stalk (without any red or pink), a darker
(olive-gray to deep brown) cap, and a northern distribution. I have seen it under conifers
in northern Idaho and British Columbia.
524 BOLETACEAE

Boletus rubripes (Red-Stemmed Bitter Bolete) Color Plate 130

CAP 6-15 (25) cm broad, convex to broadly convex or sometimes plane; surface dry, dull,
smooth or velvety, often becoming cracked or furrowed in age; pale buff to olive-buff to
tan. Flesh thick, firm, buff to pale yellow or even whitish, but blueing when exposed; taste
usually bitter. PORES and tubes pale yellow becoming darker or duller yellow in age;
blueing when bruised. STALK 7-13 (20) cm long, 1-4 cm thick, sometimes swollen below
(especially when young), but often more or less equal in age; firm, solid, not reticulate but
often longitudinally striate in age; color variable, but usually yellow at apex with bright
pink to reddish areas below, becoming dark red throughout in age. SPORE PRINT olive-
brown; spores 12.5-17.5 * 4-5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary to scattered or in groups, associated primarily with conifers; known
oniy from western N orth America and Mexico. I have not seen it in our area, but it is fairly
common in northern California under Sitka spruce and also occurs in the Sierra Nevada.
It fruits in the summer and fall. The color photograph was taken in New Mexico.
EDIBILITY: Bitter-tasting (see comments on the edibility of B. “marshii”).
COMMENTS: This beautiful but bitter bolete can be told from B. calopus by its non-
reticulate stalk (use a hand lens if unsure!), and from B. “marshii” by the red to pink stalk
which beomes dark red or dull purple-red in age. Several Tylopilus species are also bitter¬
tasting, but do not have yellow pores.

Boletus “marshii” (Ben’s Bitter Bolete)

CAP 5-25 cm broad, convex to nearly plane or somewhat irregular; surface dry, smooth
or cracking in age, whitish to gray, pale grayish-brown or buff when young, becoming
dull brownish and developing darker brown stains in age or upon handling. Flesh thick,
dense, white or grayish, turning blue to bluish-gray erratically when exposed; taste bitter,
at least latently. PORES and tubes pale yellow becoming dingy yellow or olive-yellow,
bruising blue quickly, then eventually turning dingy brown. STALK 4-15 cm long, 2-5
cm thick at apex, usually enlarged below but sometimes tapered at the base, smooth, not
reticulate; pallid to buff or yellow above (sometimes with a very slight reddish zone), dingy
brown below, darker brown throughout in old age; turning blue when bruised. SPORE
PRINT olive-brown; spores 11-14 * 4.5-6 microns, spindle-shaped to elliptical, smooth.

HABITAT: Solitary to gregarious on ground near or under live oak in the summer and
early fall (before the onset of the fall rainy season); common in the vicinity of Santa Cruz,
California, and also reported by several collectors from the foothills of the Sierra Nevada.

This bitter-tasting bolete is known locally as Boletus “marshii’' (or the “Shucks Bolete,” because
that’s what you say when you taste it!). Note pale cap, blue-staining pores, and non-reticulate stalk.
BOLETUS 525

EDIBILITY: Sauteed delicately in butter with a pinch of pepper and a clove of garlic,
served steaming hot on toast with cream cheese and celery, broiled belligerently on a
skewer with spiced lamb and bell peppers, or layered lovingly in a casserole with parmesan
cheese, egg noodles, and onions, Boletus “marshii” is still inedible.

COMMENTS: This bulky bolete with the bitter taste appears to be unnamed. However,
it is known to local yokels as B. “marshii” because it bears an uncanny resemblance to
its discoverer, Ben Marsh, who is also dense, bulky, bitter, and bulbous, and who spent
many fruitless hours (and ruined many otherwise marvelous meals) in a highly commen¬
dable if ill-conceived attempt to make it palatable. It is easily distinguished from B. calopus
by the non-reticulate stalk, from B. appendiculatus by the paler cap, non-reticulate stalk,
and bitter taste, from B. rubripes by the paler cap and absence of red on the stalk, and
from B. barrowsii by the bitter taste, blue-staining pores that are yellow when young, and
non-reticulate stalk. Its habit of fruiting before the rains arrive is odd, but quite reliable.
A European species,!?, albidus(-B. radicans), is similar in color and taste, but is usually
described as having a reticulate stalk with a rooting base. Although B. “marshii” never
seems to have a reticulate stalk, it often has a rooting base, and a critical comparison may
reveal that the two species are one and the same.

Boletus appendiculatus (Butter Bolete) Color Plate 134

CAP 6-20 (30) cm broad, convex becoming broadly convex or even plane; surface dry or
slightly viscid, smooth or with a fine bloom when young, sometimes cracked inage; brown
to cinnamon-brown or yellow-brown, or sometimes buff to yellowish, often with reddish
stains or blushes. Flesh thick, very firm and dense, pale yellow to yellow, changing slowly
and erratically to blue when cut, or not at all; taste mild. PORES and tubes lemon-yellow to
bright butter-yellow or in old age olive-yellow; usually blueing when bruised, but
sometimes not (especially in button stage). STALK 5-15 cm long, 2-6 cm thick at apex,
usually bulbous or thicker below, but sometimes equal or tapered at base; solid, very firm;
uniformly yellow or butter-yellow, sometimes with brownish or reddish stains; upper
portion finely reticulate; flesh in base usually pinkish or vinaceous. SPORE PRINT dark
olive-brown; spores 12-15 * 3.5-5 microns, spindle-shaped to elliptical, smooth.
HABIT AT: S olitary to scattered or gregarious on ground under hardwoods, especially live
oak and tanoak; fruiting shortly after the first fall rains and apparently more common in
our area than anywhere else in North America. It is abundant some years, practically
absent others; it often grows with B. satanas, B. regius, and/ or B. aereus.
EDIBILITY: Edible and popular (although some people are “allergic” to it). It lacks the
nuttiness of B. edulis but is remarkable for its firmness—a joy to find as well as to eat,
something you can really sink your teeth into. When cooked it will often turn blue, then
gray, then back to yellow,
COMMENTS: The butter bolete is easily recognized by its heaviness (it is the densest of
our boletes—the young buttons are sometimes as hard as rocks), mild (never bitter!) taste,
yellow pores that usually bruise blue quickly, and finely reticulate, butter-yellow stalk.
Don’t be misled by the variation in cap color—it is usually brown, but ranges from yel¬
lowish to somewhat reddish or rusty-brown. Sometimes it seems to intergrade with B.
regius, which is also edible. The reticulation is usually the same color as the stalk (unless
stained brown by spores), and thus can be difficult to see(a hand lens will help). Specimens
of the poisonous B. satanas which have lost their red pore color are easily distinguished
because they do not have a yellow stalk. B. “marshii” differs in its paler cap, white flesh,
non-reticulate stalk, and bitter taste. Other species: B. abieticola, sometimes common
under fir in the Siskiyou Mountains of northern California and southern Oregon, is quite
similar, but has a more fibrillose, tan to rose-colored cap.
526 BOLETACEAE

Boletus regius (Red-Capped Butter Bolete) Color Plate 133

CAP 6-20 cm broad or more, convex to plane or somewhat irregular; surface smooth to
uneven or pitted, sometimes minutely hairy when young, dry or slightly viscid; pink to
rosy-red or dark red, sometimes also with brownish tones or yellow areas. Flesh very thick,
firm, yellow, slowly and erratically or irregularly blueing when exposed; taste mild.
PORES and tubes bright yellow becoming darker yellow or olive-yellow in old age; usually
blueing when bruised. STALK 4-14 cm long, 2.5-6 cm thick at apex, usually bulbous or
thicker below, often rather short in relation to cap; solid, firm, pale to bright yellow but
usually bruising blue and often reddish-stained below; finely reticulate over at least the
upper half; flesh in base often pinkish or vinaceous. SPORE PRINT olive-brown; spores
12.5-16.5 x 3.5-5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary to gregarious on ground in woods; known from California, Oregon,
and Washington. In our area it fruits in the fall under live oak, often with B. barrowsii, B.
satanas, and/ or B. appendiculatus. Like those species it does not seem to fruit every year.
In the Sierra Nevada and Cascades it grows under conifers in the spring, summer, and fall.
EDIBILITY: Edible and very similar to B. appendiculatus in flavor, but not as firm and
therefore not as good; it is often riddled with maggots.
COMMENTS: This sometimes massive bolete can be told by its blue-staining yellow pores,
yellow flesh, pink to reddish cap, and large, thick, reticulate yellow stem. It closely mimics
B. appendiculatus except for cap color, but since both are edible it hardly matters if you
confuse them. Small specimens of B. regius are shown on p. 512. B. speciosus is a similar
eastern species with a longer, slimmer stalk and a tendency to stain blue more readily.

Boletus erythropus Color Plate 135

CAP 5-15 (20) cm broad, convex becoming nearly plane to slightly depressed or pitted in
age; surface dry, minutely velvety when young, yellow-brown to cinnamon-brown, dark
brown, or reddish-brown, quickly staining bluish-black when bruised. Flesh thick, firm,
yellow or yellowish but blueing very quickly when exposed; often reddish-brown or reddish
in base of stalk. PORES red to brick-red to orange-red, orange, rusty-orange, or burnt
sienna (but sometimes yellowish in old age), blueing quickly when bruised; tubes yellow
becoming greenish-yellow in age. STALK 4-15 cm long, (1.5) 2^1 (5) cm thick, equal or
thicker at either end but not bulbous; yellowish or more often masked by a coating of
minute red to orange dots or granules; blueing quickly when bruised and often dingier
(darker or browner) in age; solid, firm, often curved; not reticulate or only very slightly
so at apex from decurrent tube walls. SPORE PRINT olive-brown to ochraceous-brown;
spores 12-16 * 4-6 microns, elliptical to spindle-shaped, smooth.

HABITAT: Solitary, scattered, or gregarious on ground in woods; widely distributed. It is

fairly common under conifers (especially spruce) from northern California to Alaska, and
under both hardwoods and conifers in eastern North America; also reported from
the Rocky Mountains. In the oak and madrone woodlands of California it is replaced by
B. amygdalinus (see comments).
EDIBILITY: Poisonous to some people—to be avoided. Two books which list B. satanas
as poisonous state flatly that B. erythropus is edible. However, Bill Everson (an intrepid
Californian toadstool-tester) was unaffected by cooked B. satanas but ate a small portion
of sauteed B. erythropus and vomited soon after—an explicit example of why you should
be cautious when trying any mushroom for the first time, even a so-called “edible” one!
COMMENTS: The red to orange pores, non-reticulate and non-bulbous stalk, and brown
to dark brown cap typify this beautiful bolete. It certainly has substance, but is puny in
comparison to the bulky hulk (hulking bulk?) of B. satanas. It bruises blue so rapidly that
Boletus erythropus, a poisonous species with orange or reddish pores that quickly stain blue. Flesh
also stains quickly when cut (see color plate). Note that the stalk is not bulbous as in B. satanas.

a few minutes frenzied handling will render it unrecognizable. A very similar species that is
apparently unique to California, B. amygdalinus, has oranger pores and grows under
hardwoods. Another look-alike, B. subvelutipes (COLOR PLATE 136), is quite common in
the Southwest as well as eastern North America. It can have a somewhat yellower cap and
often has dark red hairs at the base of the stalk. B. hypocarycinus is a very similar south¬
eastern species with a dark red to carmine stalk base and shorter spores. B. orovillus is a
rare but striking bolete with a large brilliant yellow cap, bright red pores that may or may
not bruise blue, and a thick, non-reticulate stalk. It has been taken in a few widely scattered
localities in northern California and the Pacific Northwest. None of these should be eaten.

Boletus satanas (Satan’s Bolete) Color Plate 137

CAP 7-30 cm broad, convex to broadly convex or nearly plane in age; surface dry, smooth
or sometimes cracked; pallid to gray or olive-buff, becoming suffused with pink in age,
especially toward margin (but yellowish where slug-eaten). Flesh white to yellow, blueing
when bruised, especially when young or near the tubes. PORES deep red becoming red to
pink or in old age orange to yellowish, turning blue to blue-black when bruised; tubes
yellow to greenish. ST ALK 6-15 cm long, 2-6 cm thick at apex, with a massive abrupt bulb
at base which is up to 15 cm broad and is especially prominent when young; solid, firm;
upper portion colored more or less like the cap and finely but distinctly reticulate (reti¬
culation usually reddish); bulb usually pink or pinkish-red, but the pink tones usually
more evident in youth and tending to disappear in age. SPORE PRINT brown to olive-
brown; spores 11-15 * 4-6 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary or in groups under oaks (often at the edges of pastures); known only
from California and Europe. In our area it is sometimes abundant in the fall or early
winter. It is usually one of the very first boletes to appear.
EDIBILITY: Poisonous, at least raw! It causes vomiting, diarrhea, and severe cramps.
Thorough cooking reputedly destroys the toxins, and some people eat it regularly. Its
voluminous avoirdupois is certainly inviting—but when so many more delectable and less
dangerous mushrooms abound, why tempt fate?
COMMENTS: The red pores, pallid to olive-buff cap, and bulbous reticulate stem are
diagnostic. Young specimens can be told by their obesity alone—the bulb can be several
inches broad, often larger than the cap and just as round! The red color of the pores
527
528 BOLETACEAE

sometimes fades in old age, but the pronounced bulb and blue-staining tubes signify B.
satanas. I found one specimen at the edge of a meadow that weighed six pounds, and half of
it had been eaten by a cow! B. pulcherrimus is the only other red-pored, reticulate-stalked
bolete in coastal California, but it is not so bulbous and has a brown to reddish-brown cap.

Boletus pulcherrimus (Red-Pored Bolete)

CAP 8-25 cm broad, convex to broadly convex or nearly plane; surface dry, smooth or
minutely velvety when young, sometimes breaking up into small scales in age; brown
to pale olive-brown or reddish-brown( often redder at margin). Flesh thick, yellow, blueing
when exposed. PORES deep red to bright scarlet-red, often duller (reddish-brown) or
paler in old age; quickly staining blue or blue-black when bruised; tubes yellow. STALK
7-20 cm long, 2-5 cm thick at apex and up to 10 cm thick at base; sometimes equal, more
often club-shaped (thicker below), but not abruptly bulbous; solid, firm, yellowish to pale
reddish-brown with dark red reticulation over at least the upper half. SPORE PRINT
olive-brown; spores 13-16 * 5.5-6.5 microns, spindle-shaped to elliptical, smooth.

HABIT AT: S olitary to gregarious in mixed woods and under conifers in the summer and
fall; endemic to western North America, rather rare but sometimes fruiting in quantity.
I’ve seen it in northern California and New Mexico, but have yet to find it in our area.
EDIBILITY: Poisonous! Like B. satanas, it is especially dangerous raw.
COMMENTS: Listed in older books as B. eastwoodiae, this large and beautiful bolete is
easily told by its red to dark red pores, reticulate stalk, and brown to reddish-brown cap. It
is the principal red-pored bolete of the Pacific Northwest, but does not fruit every year.
In our area it is replaced by B. satanas, which is differently colored and much more obese.
A very similar species, B. haematinus (COLOR PLATE 138) is common under western
mountain conifers. It has a yellow-brown to olive-brown or brown cap, sometimes has a
yellower stalk, and often has yellow pores when very young (the pores at the cap margin
of older specimens can also be yellow). A similar but unidentified species with a pallid cap
has been found by Greg Wright under conifers in southern California. Another reticulate-
stalked species, B. luridus, grows under hardwoods and conifers in eastern North America
and the Southwest. It has a slimmer (1-3 cm) stalk, red to orange pores, and a yellowish to
olive-brown, brown, or reddish-brown cap. All of the above species are poisonous and all
lack the exaggerated bulb of B. satanas. For species with bright red caps, see B.frostii.

Boletusfrostii (Apple Bolete; Frost’s Bolete) Color Plate 139

CAP 5-15 cm broad, convex to broadly convex, often becoming plane or with an uplifted
margin in age; surface smooth, viscid when moist and often shiny, dark red (or even
blackish-red) when young, becoming blood-red or apple-red in age, and often developing
yellowish (faded) areas when old. Flesh thick, pallid to yellow, quickly blueing when cut.
PORES small, dark red when fresh and often beaded with yellow droplets when young,
often fading to paler red in age; typically staining dark or dingy bluish when bruised;
tubes yellow to olive. STALK 4-12 cm long, 1-3.5 cm thick, equal or thicker below, dry,
solid, coarsely and deeply reticulate throughout; dark red to red, sometimes with yellowish
areas; reticulation same color or paler; base red to yellow or whitish. SPORE PRINT
brown to olive-brown; spores 11-17 * 4-6 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary, scattered, or in groups under hardwoods; common during the
summer throughout eastern North America, also in Texas, southern Arizona, Mexico,
and Costa Rica. It favors oak, but is said to be frequent under madrone in Mexico.
EDIBILITY: Edible. In Mexico it is often sold in farmer’s markets, but since it has red
pores, be careful! Cook it thoroughly and be sure of your identification!
Left: Boletus fibrillosus (see p. 523) can be confused with B. edulis and B. aereus, but has a hairier
(fibrillose) cap in age and usually has yellow pores. Right: Boletus barrowsii, oak-loving form (also
shown on p. 512). Note whitish cap. See color plate for pine-loving variety from the Southwest.

COMMENTS: One of the most beautiful and memorable of all boletes, this species is
easily told by its viscid red to dark red cap, dark red pores, and coarsely reticulate stalk.
The stalk may be thickened downward, but is never bulbous as in B. satanas. The tendency
of the young pores to exude yellow droplets is also distinctive. In Mexico it is sometimes
called panza agria, which means “sour belly.” Other species: B. flammans and B. rubro-
flammeus, both poisonous, have a non-viscid red to dark red cap and finely to scarcely
reticulate stalk and red pores (at least when young). The first favors eastern conifers, the
second grows with hardwoods in eastern North America and southern Arizona.

Boletus barrowsii (White King Bolete) Color Plate 141

CAP 6-25 cm broad, convex becoming broadly convex to plane; surface smooth or very
slightly velvety, dry, dull white to buff or grayish. Flesh thick, white, not blueing when
exposed (or blueing only very slightly near the tubes). PORES and tubes white or pallid
when young (and appearing stuffed with a pith), becoming yellow to olive-yellow in age;
not blueing when bruised. STALK 6-20 cm long, 2-6 cm thick at apex, equal or thicker
below (especially when young), solid, firm, whitish to buff or colored like cap, sometimes
with brownish stains; finely but distinctly reticulate at least over the upper portion. SPORE
PRINT dark olive-brown; spores 13-15 * 4-5 microns, spindle-shaped toelliptical, smooth.
HABITAT: Solitary or scattered to gregarious under both hardwoods and conifers,
fruiting mainly in the summer and fall; apparently endemic to the drier parts of western
North America and Mexico. It is a warm weather species and more abundant in Arizona
and New Mexico than anywhere else. There it is associated almost exclusively with ponde-
rosa pine, while B. edulis favors spruce. In Colorado, however, I have found it under spruce
and fir, and in coastal California it is usually associated—quite curiously—with live oak,
while B. edulis grows with pine. In our area it fruits in the fall and winter, but is abundant
only after warm early (September-November) rains.

EDIBILITY: Delectably delicious. It is a favorite of collectors in the Southwest, and in

the completely objective opinion of its “discoverer,” Chuck Barrows, it is the best of the
boletes for the table. Unfortunately, the maggots share his opinion, for they attack it even
more voraciously than they do B. edulis\

COMMENTS: For many years this handsome, meaty bolete has passed as a “white”
B. edulis, and is listed as such in the first edition of Mushrooms Demystified. However,
it does not intergrade with B. edulis and lacks the clearly differentiated, often viscid cap
cuticle (skin) of that species. Result: it is now recognized as a distinct species.
 530 BOLETACEAE

Boletus edulis Color Plates 142,143,144

(King Bolete; Cep; Steinpilz; Porcini; etc.)
CAP 8-30 cm broad or more, convex to broadly convex or bun-shaped, orinage becoming
plane; surface dry or more often viscid when moist, smooth to somewhat pitted or in dry
weather sometimes cracking into plaques; color variable: biscuit-brown or warm brown to
yellow-brown, cinnamon-brown, reddish-brown, or dark red (but often with whitish or
pinkish tints while still under the duff); margin sometimes paler or yellower. Flesh thick,
firm, white or sometimes tinged yellowish or dingy reddish inage; not blueing when bruised
or only blueing slightly near the tubes; odor and taste pleasant. PORES at first white or
pallid and appearing stuffed with a pith, then becoming yellow, olive-yellow, or brown in
age, not blueing when bruised; tubes whitish becoming yellow, then olive-yellow. STALK
(3) 8-25 cm long, 2-7 (12) cm thick (often large in relation to cap); usually enlarged below
when young, often becoming more or less equal in age; firm, solid, entirely white, or
whitish at the base and brownish above; finely reticulate over upper portion or throughout;
flesh in base sometimes with dark yellow areas (a parasite?). SPORE PRINT more or less
olive-brown; spores 13-19 * 4-7 microns, spindle-shaped to elliptical, smooth.

HABITAT: Solitary, scattered, or in groups or sometimes troups on ground in woods;

found throughout the world and very common in western North America. It favors coni¬
fers (pine, spruce, hemlock, fir) but also grows with hardwoods such as oak and birch. It
is often abundant in our coastal pine forests in the fall and winter(usually 2-4 weeks after
the first substantial fall rains) as well as under oak, and a smaller flush may appear in the
spring. In coastal northern California and Oregon it grows with pine and spruce in the fall;
in the Sierra Nevada a stout form fruits in large quantities under conifers in the spring,
summer, and fall; and in the high Rockies the variety with a dark red cap fruits in colossal
quantities in August and September, under spruce.
EDIBILITY: One of the finest of fleshy fungi and certainly the best-loved and most sought-
after in Europe, where it has more common names than there are languages. If any
mushroom deserves the dubious title of “king,” this is the one. It is a consummate crea¬
tion, the peerless epitome of earthbound substance, a bald bulbous pillar of thick white
flesh—the one aristocrat the peasantry can eat!
The entire fruiting body is exceptionally delicious, even the tubes if they’re firm enough.
It is delicious raw, but can cause stomach upsets because it’s difficult to digest, so play it
safe and cook it (preferably on high heat in an open pan, so as to drive off the excess mois¬
ture). The odor and taste of dried B. edulis are marvelous—nutty, earthy, and meaty all
at once. But you have to find them before you can eat them, and it isn’t always easy. You
can’t just casually look for them the way you can for chanterelles or blewits—you have to
hunt them down and root them up from under the duff before they’re visible to others.
Timing is of paramount importance, because you face formidable competition from both
maggots and boletivores (see p. 546). One source suggests getting up at the crack of dawn
and wearing your shirt inside out. If that doesn’t work, you can always resort to the dried,
imported version found in delicatessens.
COMMENTS: This magnificent mushroom has several color forms, two of which are
particularly common. The first has a brown to reddish-browncap(Color Plates 142 & 144)
and is the dominant one in coastal California. The second, also called B. edulis var. pinicola
(-B. pinicola, B. pinophilus), has a dark red to red-brown cap. It is prevalent in the Rocky
Mountains and Southwest (Color Plate 143) and Gulf Coast region. Remember: brown to
reddish cap, thick reticulate stalk, mild to nutty (not bitter) taste, and white pores when young
that do not bruise blue. No mushroom is more substantial or satisfying to find! Other edible
species: B. mottii of the Sierra Nevada has a pitted and / or ridged cap; B. separans (reddish to
liver-colored cap when young) and B. pseudoseparans {dark purple cap when young) usually
have a purple or purple-tinged stalk and grow under hardwoods in eastern North America.
Boletus edulis, young specimens with white pores. Some forms have much stouter stalks than these,
which have been trimmed to check for maggots. The caps will broaden as they mature. See p.489 for
a close-up of the reticulate stalk. Also see color plates and photo on p. 512. (Joel Leivick)

Tylopilus species can be somewhat similar, but have pinker pores, pinkish- to reddish-
brown spores, stain dark brown when bruised, and/ or are bitter-tasting. B. fibrillosus is
another look-alike, but its cap is darker and hairier. See also B. aereus and B. barrowsii.

Boletus aereus (Queen Bolete) Color Plate 140

CAP 5-15 (20) cm broad, convex becoming broadly convex to plane in age; surface dry or
moist (or viscid only in age), smooth or somewhat pitted, dark brown to nearly black when
young and covered at least partially with a fine whitish bloom, but inage becoming smooth
and cinnamon or red-brown or blotched with paler (whitish to tan) areas. Flesh thick,
white or tinged reddish, not blueing when exposed (or blueing only slightly near the tubes);
taste mild or pleasant. PORES and tubes white when young and at first stuffed with a pith,
becoming yellow to greenish-yellow in age; not blueing when bruised. STALK 5-15 cm
long, 2-5 cm thick at apex, usually enlarged below when young but often equal in age; firm,
solid, white or often brown in age, finely reticulate at least over upper part. SPORE
PRINT dark olive-brown; spores 12-14 * 4-5 microns, spindle-shaped toelliptical, smooth.
HABITAT: Solitary, scattered, or gregarious in mixed woods and under hardwoods
(especially oaks); found in California, but originally described from Europe. In our area
it is fairly common in the fall in the coastal mountains, usually under tanoak, madrone,
and/ or chinquapin. It also grows with live oak, but not as commonly as B. barrowsii.
EDIBILITY: Delicious! It is every bit as good as B. edulis, and less apt to be wormy!
COMMENTS: This relative of B. edulis often grows with B. appendiculatus, but has white
pores when young, a darker cap, and does not stain blue. The hoary bloom on young caps
(see photos on pp. 50 & 512) disappears as the cap becomes more cinnamon-colored, and
in age there is little to distinguish it from B. edulis other than habitat. Fortunately, both
species are so delectably delicious that it hardly matters if you unwittingly confuse them!
B. fibrillosus is also similar, but has a hairier cap without a whitish bloom. Another cousin
of B. edulis, B. variipes, has a dry, tan to grayish-brown, yellow-brown, or dark brown cap
that lacks the whitish bloom of B. aereus and often becomes areolate (cracked) in age. It is
abundant in the summer under oak and beech in eastern N orth America, but rare or absent
in the West. It is edible, but usually riddled with maggots and sometimes bitterish to boot.

531
532 BOLETACEAE

TYLOPILUS

Medium-sized to large, fleshy, mostly terrestrial boletes. CAP usually dull colored and typically not
viscid. PORES and tubes typically pallid to pinkish, vinaceous, gray, or brown. STALK fleshy,
sometimes reticulate, but not glandular-dotted and not usually with dark scabers. VEIL absent.
SPORE PRINT flesh-colored to pinkish-brown, reddish-brown, cinnamon-brown, vinaceous,
or chocolate-brown. Spores elliptical to spindle-shaped, smooth (except T. gracilis).

TYLOPILUS closely resembles Boletus, but gives a flesh-colored to reddish-brown or

chocolate-brown spore print instead of a brown to olive-brown one. In the field the two
genera can often be told by the color of their pores—in Boletus they are usually orange,
red, yellow, or white and frequently stain blue when bruised, while in Tylopilus they are
typically pallid to flesh-colored, vinaceous, gray, or some shade of brown, and are more
apt to stain reddish, brown, or black than blue.
In eastern North America Tylopilus, like Boletus, is quite diverse and often abundant
in the summer and early fall. The West presents a completely different picture, however—a
mere handful of species occur and only one, T. pseudoscaber (a species “complex”), could
be characterized as “common.” Tylopilus does not offer the plethora of good edibles that
Boletus does. I can find no mention of any Tylopilus being poisonous, but many species
(e.g., T. felleus) have a very bitter taste—even when cooked. Since Tylopilus is such a
conspicuous component of the bolete bounty of eastern North America, I have included
the more common and distinctive eastern species in the key. Four species are described
(three of which occur in California); two additional species are shown in the color plates.

Key to Tylopilus
1. Tubes yellow; pores yellow to brownish or reddish, often blueing when bruised; cap usually
areolate (cracked) in age; western .T. amylosporus(see Boletus chrysenteron, p. 519)
1. Not as above; tubes not yellow, or if so then found in eastern North America .2
2. Stalk white or creamy (but staining dark brown); pores white when fresh but quickly staining
dark red and then dark brown when bruised; flesh in cap and upper stalk often blueing slightly
when cut, then turning reddish or brown; found in eastern North America; rare . . T. snellii
2. Not as above; staining reactions different, or if similar then stalk not whitish; common ... 3
3. Pores and/ or flesh staining blue or blue-black when bruised or at least staining waxed or white
paper blue or blue-green when wrapped in it; fruiting body dark T. pseudoscaber group, p.534
3. Not as above; not staining waxed paper blue or blue-green; fruiting body dark or light .... 4
4. Cap and stalk dark gray to brownish-black or black, but the pores white when youngand pinkish
in age; pores and flesh staining reddish or vinaceous when bruised, then often blackening;
fruiting body usually robust; spore print pinkish or vinaceous; found ineasternNorth America
and the Southwest, usually under hardwoods such as oak; edible .T. alboater
4. Not as above; differently colored or with darker (chocolate-brown, etc.) spores or pores .. 5
5. Pore surface dark gray to dark brown or blackish, even when young; stalk densely scurfy from
numerous minute scales or granules; cap and stalk chocolate-brown to dark brown or with a
purple or bluish tinge; spore print pinkish-brown to vinaceous; found in eastern North
America, mainly under northern conifers. T. eximius
5. Not as above (if dark, then spore print chocolate-brown or stalk not densely scurfy) .6
6. Cap and stalk dark (olive-brown to grayish-brown, dark brown, or blackish); pores usually
gray to brown, at least in age; spore print chocolate-brown . . T. pseudoscaber group, p. 534
6. Not as above; differently colored and/ or spore print pinkish to vinaceous or cinnamon ... 7
7. Found in eastern North America (east of the Rocky Mountains)* .8
7. Found in western North America (from the Rocky Mountains westward)* . 18
8. Stalk base bright yellow; cap pink when fresh (but fading to tan or paler) T. chromapes, p. 533
8. Not as above . 9
9. Cap whitish to buff or pale yellow, sometimes dingy tan in age .21
9. Not as above; cap darker or brighter. 10

*In regions where the eastern and western fungal floras overlap (e.g., southern Arizona), try both choices!
TYLOPILUS 533

10. Cap broadly conical and shaggy-fibrillose (or pitted in age), yellow to bright yellow-brown or
tinged pink, margin often fringed; found in southern bottomlands T. (-Mucilopilus) conicus
10. Not as above; cap not shaggy . 11
n. Cap bright orange to dull orange (but may become cinnamon or browner in age); stalk white to
yellow or orange (or aging brownish) .T. ballouii(see T. chromapes, below)
11. Not as above; cap yellow-brown to cinnamon, brown, purplish, etc., but not truly orange 12
12. Flesh very bitter-tasting (but some people have trouble detecting the bitterness) . 13
12. Taste not bitter or only slightly or sporadically so. 16
13. Stalk purple or purple-and-white when fresh (but may fade in age!); cap purplish to brown;
pores white when young, dingy pinkish in age; stalk not reticulate or only slightly so at apex;
usually found under hardwoods; common . T. plumbeoviolaceus
13. Not as above; cinnamon to brown or dark brown but not purple when fresh . 14
14. Stalk clearly reticulate; usually (not always) under conifers T.felleus(see T. indecisus, p. 535)
14. Stalk not reticulate or only slightly so at apex; usually associated with oak . 15
15. Stalk thick (1-5 cm at apex); cap medium-sized to large; widespread .T. rubrobrunneus
15. Stalk thinner than above; cap often rather small; southern .T. minor
16. Stalk typically long and slender (less than 1 cm thick at apex), often curved at base; cap and stalk
reddish-brown to cinnamon or tawny (see Color Plate 149); found in many habitats but partial
to hemlock; many spores minutely roughened or pitted T. gracilis (see T. chromapes, below)
16. Not as above; stalk usually at least 1 cm thick; spores smooth . 17
17. Cap more or less maroon-red . T. badiceps
17. Not as above .24
18. Stalk distinctly reticulate over upper portion; associated mainly with oak T. indecisus, p. 535
18. Stalk not reticulate or only obscurely so at apex from decurrent tube walls. 19
19. Stalk often short, poorly-developed, and sometimes off-center; margin of cap often incurved
even at maturity; fruiting body often partly buried in soil . T. humilis, p. 535
19. Not as above . 20
20. Cap dark brown .T. ferrugineus(?)
20. Cap tan to dull brown or vinaceous-tinged . T. ammiratii(see T. indecisus, p. 535)
21. Stalk distinctly reticulate or reticulate-ridged; mainly southern . 22
21. Stalk not reticulate or only slightly so at apex; southern or northern .23
22. Reticulation on stalk coarse and raised; stalk usually long in relation to cap; cap often areolate
in age; spores pitted or roughened .(see Boletellus& Austroboletus, p. 508)
22. Not as above; taste bitter or mild; fruiting body often robust .T. rhoadsiae
23. Cap typically wrinkled; northern.T. intermedius
23. Cap not normally wrinkled; southern .T. peralbidus
24. Fruiting body medium-sized to large, tawny to orange-brown to brown (including the pores!);
stalk usually reticulate; taste mild to slightly bitter; mainly southern .T. tabacinus
24. N ot as above; pores usually paler when fresh . T. indecisus & others, p. 535

Tylopilus chromapes (Chrome-Footed Bolete)

CAP (3) 5-15 cm broad, convex, often becoming more or less plane in age; surface smooth
or felty, dry to slightly viscid, bright pink to rose-colored when fresh, but often fading(to
pinkish-tan, tan, or even whitish) in age. Flesh thick, white or tinged pink, not blueing;
taste mild or slightly acidic. PORES minute, white when young becoming pinkish to flesh-
colored to dingy brownish in age, not staining appreciably when bruised, or staining
pinkish; tubes more or less same color or slightly yellower. STALK 4-20 cm long, 1-2.5
cm thick, equal or slightly tapered either way, solid, firm; upper portion white or pallid or
sometimes splotched with pink (especially when young), base bright yellow inside and out;
not reticulate but often scurfy above from small hairs or scales. SPORE PRINT vinaceous-
to pinkish-brown; spores 10-17 * 4-5.5 microns, elliptical to spindle-shaped, smooth.
HABITAT: Solitary to gregarious on ground under both hardwoods and conifers; locally
534 BOLETACEAE

frequent throughout eastern North America in the summer and fall. It is especially
common under birch, aspen, and conifers in northern latitudes in June and July.
EDIBILITY: Edible and quite good, but often riddled with maggots.
COMMENTS: Formerly known as Boletus chromapes and Leccinum chromapes, this
lovely bolete is easily recognized by its pink cap and white pores when young and its
scurfy stalk with a bright yellow base. (The color scheme is reminiscent of Gomphidius
subroseus, a gilled mushroom closely related to the boletes.) It is one of several very
striking species of Tylopilus that are native to eastern North America. Others include:
T. plumbeoviolaceus, a beautiful but bitter-tasting violet to purple-brown species; T.
(-Austroboletus) gracilis, a slender-stemmed species (COLOR PLATE 149); and T.
ballouii, with an orange, Leccinum-like cap when fresh. For more details on these and
other distinctive eastern species, see the key to Tylopilus.

Tylopiluspseudoscaber group (Dark Bolete)

CAP 5-16 cm broad, convex to plane or somewhat irregular; surface dry, minutely velvety
to smooth in age, dark brown to very dark brown or at times grayish-brown to olive-brown,
usually bruising darker brown to blackish; margin sometimes slightly paler. Flesh thick,
whitish, typically bruising blue (but sometimes slowly or erratically), then slowly dis¬
coloring to reddish-brown or brown (or sometimes staining directly to pinkish); odor often
pungent, taste mild or at least not bitter. PORES and tubes dark yellow-brown to coffee-
brown to grayish-brown, chocolate-brown, or very dark brown (but in some forms pallid
to grayish or yellow-gray when young); usually (but not always!) staining blue and then
slowly reddish to dark brown when bruised. STALK (4) 7-16 (20) cm long, 1-3 (4) cm thick,
equal or more often thicker below, firm, dry, minutely velvety and/or scurfy, often
longitudinally lined or streaked; distinctly reticulate above in one form, not reticulate in
others; brown or colored like cap or darker, usually bruising darker brown or sometimes
blue; base typically whitish. SPORE PRINT pale to dark chocolate-brown or reddish-
brown; spores 11-19 (25) * 5-9 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary, scattered, or in groups on ground or rotten wood, usually under
conifers but sometimes with hardwoods; widely distributed but mainly northern. It is
fairly common in the late summer and fall in the Pacific Northwest and northern Cali¬
fornia. It is especially fond of Sitkaspruce—which may explain its absence ornearabsence
south of Mendocino County, California (the southernmost limit of Sitka spruce).
EDIBILITY: Edible, but according to most sources, far from incredible. Its somber
color is unappetizing and makes it difficult to see in the forest gloom. I haven’t tried it.

COMMENTS: Also known as Porphyrelluspseudoscaber, this distinctive bolete is easily

told by its dark color (see photo on p. 535), reddish-brown spore print, dark brown pores in
age, and tendency to stain blue. The latter feature varies considerably in duration and
intensity and may even be absent; however, if the fruiting body is wrapped in white or
waxed paper it will usually stain the paper blue or dark blue-green. The above description
is actually a composite of several closely related “dark boletes” that have passed under
the names T. pseudoscaber, T. porphyrosporus, T. olivaceobrunneus, T. atrofuscus,
T. sordidus, etc. Some mycologists consider them to be forms of one highly variable
species. Others consider them distinct—for instance, they reserve the name T. porphy¬
rosporus for the form with pallid pores when young, and T. olivaceobrunneus for the
variety with a reticulate stalk. Other species in the group include: T. pacificus, which tends
to have a copiously cracked (areolate) cap in age; T.fumosipes, an eastern species with an
areolate cap in age and narrower spores; and T. nebulosus, which typically does not stain
blue. Two other dark species, T. alboater and T. eximius, are described in the key.
Left: Tylopiluspseudoscaber group. Note dark color. Right: Tylopilus indecisus is apt to be mistaken
for a Boletus, but like many species of Tylopilus, it stains brown when handled.

Tylopilus indecisus (Indecisive Bolete)

CAP 5-15 (25) cm broad, convex to plane; surface usually dry and smooth, dark brown to
brown, dingy brown, or dingy cinnamon (usually darker when young, paler in age), stain¬
ing dark brown where bruised. Flesh thick, pallid, often staining pale vinaceous or flesh-
color when bruised; taste mild. PORES and tubes pallid when young, soon becoming dull
pinkish or pale flesh-color; slightly darker, dingier, or more vinaceous in age, staining
brown where bruised. STALK 4-10 cm long, 1-3 (4.5) cm thick, equal or thicker below,
dry, firm, buff to brown, the apex often pallid; darker brown where bruised or handled;
reticulate at least at the apex (but eastern variety sometimes not reticulate), sometimes
nearly throughout. SPORE PRINT flesh-colored to reddish-brown or tinged vinaceous;
spores 10-15 * 3-5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary to scattered or in small groups in mixed woods and under oaks; fairly
common in eastern North America, but rare in the West. I have seen it only a few times in
our area, under live oak in the late fall and early winter, and under white oak inland.
EDIBILITY: Edible, according to the literature. I haven’t fried it.
COMMENTS: There is nothing particularly indecisive about this drab bolete, but there is
nothing particularly decisive about it either. The brown cap and reticulate stalk are remi¬
niscent of Boletus edulis, but the pores are pinker (and never olive-yellow), the fruiting
body stains dark brown when handled or bruised, and the spore print is pinkish. Eastern
material (including var. subpunctipes, which differs microscopically) is less apt to be
reticulate than the western version. T. ammiratiiis similar but has a tan or vinaceous-tinged
cap and whitish stalk that is not reticulate or only slightly so at the apex; it occurs in Califor¬
nia under oak. Similar easterners include: T. subunicolor, a small, slim, tawny-ochre
southern species with yellowish flesh; T. tabacinus and T. rubrobrunneus, both medium to
large (see key to Tylopilus)', and T.felleus (COLOR PLATE 150), with a reticulate stalk
and very bitter taste. T. felleus is especially common under conifers (e.g., hemlock and
pine); it is often mistaken for Boletus edulis. For other easterners, see key to Tylopilus.

Tylopilus humilus (Humble Bolete)

CAP 4-12 cm broad, nearly round to convex when young, sometimes expanding to plane in
age or often somewhat misshapen or irregular; surface dry or slightly tacky, often minutely
velvety, dull brown or sometimes reddish-brown; margin often with vinaceous tints and
frequently remaining incurved through maturity or even covering some of the pores. Flesh
thick, white, bruising pinkish-brown or vinaceous (rarely blue); taste mild. PORES and

535
Tylopilus humilus is a small, rare, often misshapen bolete. Note how stalk is often poorly developed.

tubes whitish when young, becoming pale flesh-color or dull pale pinkish, staining brown
when bruised. STALK 2-5 cm long, 1-3 cm thick, usually rather stocky and sometimes
off-center; equal or swollen below, but the base often pinched; firm, solid, white at apex,
usually brownish- or vinaceous-stained below, especially when handled; not reticulate or
only very slightly so at extreme apex. SPORE PRINT dull reddish-brown; spores 8-12 *
x 3-4 microns, spindle-shaped to elliptical, smooth.
HABITAT: Scattered to gregarious or in small clumps on ground (often partially buried),
fruiting in the fall and winter; known only from California, rare. It appears every year in
sandy soil in a burned-over area near Santa Cruz, apparently in association with manzanita
or live oak, but with knobcone pine also in the general vicinity. It was originally discovered
by Harry Thiers in Mendocino County under pines near a destroyed dormitory.
COMMENTS: This bashful bolete is easily separated from other Tylopilus species by its
semi-underground habit, i.e., it tends not to expose itself until fully mature, and even then
may remain half-buried. The stocky or poorly-developed stem is reminiscent of Gastro-
boletus, but the spores are forcibly discharged and the tubes are arranged vertically.

LECCINUM (Rough-Stemmed Boletes)

Medium-sized to large, fleshy, terrestrial, mycorrhizal, woodland boletes. CAP usually some
shade of orange, red, brown, gray, or white; viscid or dry; often rimmed with a flap of sterile tissue
when young. PORES and tubes usually whitish to dingy buff, grayish, grayish-olive, or pale brown,
only very rarely yellow; not staining blue. STALK fleshy, usually rather tough and fibrous, typically
not reticulate, ornamented with scabers which usually darken at maturity. VEIL typically absent.
SPORE PRINT brown. Spores spindle-shaped to elliptical, smooth.

LECCINUM differs from other boletes by virtue of the tufted hairs or small rough scales
(scabers) on the stalk. The scabers may be pallid when young, but usually darken to brown
or black by maturity. They should not be confused with the resinous or sticky glandular
dots found in Suillus, which do not protrude as scabers do and are not composed of hairs.
Leccinums also have a characteristic appearance which makes them recognizable from a
distance: the cap is typically some shade of orange, reddish-brown, brown, or white; the
pores are usually white or dingy-colored (not yellow or red) and do not stain blue; and the
stalk is usually whitish or pale colored except for the scabers, and usually much tougher
and more fibrous than the cap. Also, the stalk sometimes stains blue when cut or handled,
and in many species is rather long in relation to the cap.
Leccinums are good boletes for beginners because they are widespread, often abundant

536
LECCINUM 537

(in Alaska they are said to outnumber all other boletes), and edible (though a few, like
L. atrostipitatum, reportedly cause stomach upsets in some people). Nearly all Leccinums
have a tendency to blacken when cooked or dried, but in no way does this affect their
flavor. They dry quite nicely and are less apt to be maggoty than other boletes, at least
in our area. Some species, such as L. insigne, are delicious fresh. Others, like L. man-
zanitae, are watery and bland unless dried first. Even the less savory ones, however, have
an important role to fulfill—they help stuff the buckets and bellies of boletivores (see
p. 546) who might otherwise grow despondent and dangerous!
Like most boletes, Leccinums are mycorrhizal. Their favorite hosts are aspen and birch,
but they also occur with conifers and oaks, and in California—where birch is not native
and aspen is restricted to the Sierra Nevada—they link up with madrone and manzanita,
masquerading under the moniker “manzanita boletes.” Over 100 species of Leccinum have
been described from North America (most of them from the northern part), but many are
very similar in appearance and can only be distinguished with a microscope. For instance,
L. aurantiacum differs from L. discolor by the presence of minute pigment globules in the
hyphae of the cap cuticle, while L. discolor differs from L. rufescentoides in its smaller
cuticular hyphal end cells! The staining reactions of the flesh upon exposure to air—
particularly at the juncture of the cap and stalk—are also significant and should be
determined by slicing open the fruiting body lengthwise, rubbing the flesh a couple times,
then observing it at 5-minute intervals for a period of !^-l hour.
Fortunately, Leccinum is such a safe genus that boletivorous bipeds such as you and I
can leave such dubious and esoteric distinctions to professional boletologists, and con¬
centrate on learning a few common “prototypes”—e.g., the L. manzanitae group that is
so common with manzanita and madrone; the orange to reddish-brown-capped L. insigne-
L. aurantiacum group that grows profusely wherever there are aspens; and the white-
capped L. holopus and dingy colored L. scabrum groups that favor birch. Three “proto¬
types” are described here and several others are keyed out.
Key to Leccinum
1. Pores and tubes yellow when fresh .2
1. Pores and tubes not yellow.3
2. Scabers on stalk brown to blackish in age; cap yellow to ochraceous to brown or dark brown;
found under hardwoods (mainly oak) in eastern and southern North America .29
2. Not as above; scabers reddish or paler, not darkening appreciably in age (see Boletus, p. 511)
3. Base of stalk bright yellow and cap pink when fresh (but fading!) or cap and stalk chocolate-
brown to purplish to dark blue-gray and pores chocolate-brown to dark gray to blackish at
maturity; restricted to eastern North America .(see Tylopilus, p. 532)
3. Not as above ..4
4. Margin of cap lacking sterile flap(s) of tissue; cap white to buff, grayish, brown, dingy yellow-
brown, olive-tinged, or even black when fresh . 5
4. Cap pink to apricot-buff, orange, red, reddish-brown, orange-brown, or rusty-brown, or if
colored as above then margin of young cap rimmed with a sterile flap of tissue 2-6 mm wide, the
flap breaking up into segments as the cap expands, or disappearing. 12
5. Cap white or whitish (but sometimes tinged buff, pinkish-buff, brown, or olive in age) .... 6
5. Cap more highly or deeply colored (pink, orange, red, brown, gray, black, etc.) .8
6. Associated with aspen; known only from California L. californicum (see L. scabrum, p. 541)
6. Not as above . 7
7. Cap often olive-tinged in age, sometimes viscid; common under birch in northern and eastern
North America .L. holopus & others (see L. scabrum, p. 541)
7. Not as above; cap not viscid; associated with hardwoods (especially oak) in southern and
eastern North America (especially common in the South) L. albellum(see L. scabrum, p. 541)
8. Cap bluish-black to dark grayish-brown to black when fresh (but often fading in age); asso¬
ciated with aspen and birch in eastern North America .9
8. Not as above; cap dull brown to gray-brown, dingy yellow-brown, olive-tinged, etc.10
 538 BOLETACEAE

9. Found with birch; flesh in stalk apex reddening when bruised L. snellii( see L. scab rum, p. 541)
9. N ot as above; associated with aspen .L. griseonigrum (see L. scab rum, p. 541)
10. Cap yellow-brown when young becoming duller or grayer in age and usually prominently
cracked (areolate); associated with oak and other hardwoods; especially common in south¬
eastern North America .L. griseum(see L. scab rum, p. 541)
10. Not as above; cap not usually areolate; associated with birch or aspen. 11
11. Associated with birch; widely distributed .L. scabrum & others, p. 541
11. Associated with aspen; common in the Sierra Nevada L. montanum{see L. scabrum, p. 541)
12. Veil present, covering the young cap, then breaking up into whitish patches; associated with
aspen in the Great Lakes region.L. potteri
12. Not as above; veil absent(but margin of cap may have sterile flap(s) of tissue) . 13
13. Scabers on stalk dense and coal-black even in the button stage and black or dark brown in age;
associated mainly with birch in eastern and northern North America, but also found with
aspen in Southwest .L. atrostipitatum& L. testaceoscabrum(see L. insigne, p. 540)
13. Not as above . 14
14. Associated with manzanita and madrone(and possibly toyon); cap often (but not always) viscid
when moist; common in west coast states, especially Oregon and California . 15
14. Associated with aspen, conifers, or other trees; cap viscid or dry; widely distributed but not
found in coastal California . 18
15. Cap dark red to brown or sometimes orange-brown .L. manzanitae & others, p. 539
15. Cap bright orange to pink or apricot-buff .16
16. Scabers on stalk orange; bruised or cut flesh usually reddening somewhat before darkening . .
.L. “aurantioscaber” (see L. manzanitae, p. 539)
16. Not as above .17
17. Cap pinkish to apricot; flesh not staining when bruised L. constans (see L. manzanitae, p. 539)
17. Cap orange; flesh usually reddening or darkening L. armeniacum (see L. manzanitae, p. 539)
18. Flesh (especially at juncture of cap and stalk) not changing color appreciably when cut or
bruised, or staining only very slightly (but lower stalk may stain blue); found with conifers 19
18. Flesh (especially at juncture of cap and stalk) staining purple-gray, bluish-gray, smoky, reddish,
vinaceous, etc. when cut or bruised (but often slowly or erratically); lower stalk may stain
blue as well . 20
19. Fruiting body medium-sized to very large (mature cap 10-30 cm broad); found in the Pacific
Northwest .L. ponderosum & others (see L. manzanitae, p. 539)
19. Fruiting body smaller (cap usually less than 10 cm broad); known from the Southwest and
eastern North America .L. vulpinum
20. Flesh staining purple-gray to bluish-gray or smoky (fuscous) directly when bruised or cut (but
often staining slowly or erratically) . 21
20. Flesh staining reddish, vinaceous, or burgundy when bruised or cut (but often staining purple-
gray to bluish-gray or fuscous after that) . 23
21. Cap whitish when young but becoming brownish in age; stalk thick and club-shaped (thicker or
swollen below); associated with western conifers (mainly in mountains) .L. clavatum
21. Not as above . 22
22. Cap orange to rusty-orange to reddish when young (may become browner in age), or if brown
when young then tubes often staining vinaceous or purplish when bruised L. insigne, p. 540
22. Cap dark brown to dull reddish-brown or dull brown; tubes not staining vinaceous or purplish
when bruised .L. brunneum (see L. manzanitae, p. 539)
23. Cap liver-colored to liver-brown and fibrillose to fibrillose-scaly; associated with conifers
in the Pacific Northwest . L. fibrillosum(see L. manzanitae, p. 539)
23. Not as above; cap typically reddish to rusty-orange, brownish, or paler. 24
24. Cap pale tan to pale dingy orange-brown or yellow-brown; associated with aspen and birch in
northern and eastern North America, and in the Southwest .L. cinnamomeum
24. Not as above; cap usually darker or brighter . 25
25. Cap pinkish- or vinaceous-tinged when young; known only from Idaho under whitebark pine
.L. incarnatum
25. Not as above . 26
LECCINUM 539

26. Stalk more or less equal or narrowed near apex; associated mostly with aspen or pine; very
widely distributed and common . 27
26. Stalk distinctly and consistently club-shaped (thicker below); associated with conifers (mainly
spruce and fir) in the Rocky Mountains and Southwest . 28
27. Cap typically reddish to orange to brick-red when young (often somewhat duller in age); asso¬
ciated with aspen and conifers (pine, etc.) .L. aurantiacum(see L. insigne, p. 540)
27. Cap somewhat similar in color to above or often duller even when young (brown to orange-
brown, pale cinnamon, etc.); associated only with aspen . L. discolor(see L. insigne, p. 540)
28. Cap fibrillose or fibrillose-scaly .L. subalpinum{see L. insigne, p. 540)
28. Not as above .L.fallax(see L. insigne, p. 540)
29. Stalk often robust and swollen in the middle or below; cap yellow-brown to dark brown ....
.L. crocipodium
29. Stalk long and more or less equal; cap yellow-brown to ochre, often wrinkled or finely pitted
in age .L. rugosiceps

Leccinum manzanitae (Manzanita Bolete) Color Plate 145

CAP (5) 7-20 (30) cm broad, rounded becoming convex and then broadly convex to plane
or somewhat irregular; surface viscid when moist, practically smooth but usually with
flattened fibrils, sometimes pitted in age; dark red to reddish-brown, or at times rusty-
brown, rusty-orange, or brown (see comments); margin with flaps of sterile tissue when
young, which break up into segments or disappear in age. Flesh thick, firm becoming soft
at maturity, white, usually bruising smoky-gray to purple-gray or bluish-gray when cut, but
often very slowly or only in certain areas. PORES and tubes whitish to pale olive, olive-
gray, or grayish, usually deep dingy olive-buff in age; not blueing when bruised but often
discoloring brown. STALK 8-20 cm long, 1.5-4 cm thick at apex, equal or often thicker or
swollen below, tough, fibrous, solid, white or whitish when fresh, but roughened by
numerous small projecting scabers that are pallid at first but deep brown to black by
maturity; lower portion often bruising (or already stained) bright blue to greenish-blue.
SPORE PRINT brown; spores 13-19 * 3-5 microns, spindle-shaped to elliptical, smooth.
HABITAT: Solitary to widely scattered or gregarious on ground, associated with man¬
zanita and madrone but often growing in mixed woods; known only from the west coast.
It is common in our area in the fall and winter and is easily the most prevalent Leccinum in
coastal California. It is seldom as prolific as B. edulis, but I have seen more than fifty
prime fructifications under a patch of stunted madrones in the middle of a pasture.

EDIBILITY: Edible and esteemed by many boletivores, but disappointingly bland in

my experience. Mature or waterlogged specimens should be dried in order to concentrate
their flavor and get rid of excess moisture. In one blind tasting of several local boletes (all
fresh), it placed next to last—slightly behind the boring Boletus subtomentosus but ahead
of the abominable Suillusfuscotomentosus. Michael Cabaniss, who did the line drawings
for this book, says: “I have tried only dried slices, a form which is not the most visually
stimulating . . . their consistency is rather grainy, giving the gustatory experience of dried
earth or decomposed redwood chips. Dried slices are not even chewy; they break up readily
when inserted into the oral cavity and are swallowed with great difficulty.”
COMMENTS: This large, impressive bolete is easily identified by its reddish to brown
cap, hard tough stalk with dark scabers, and growth with manzanita and madrone (see
photo on p. 39). It is the central species in a cluster of edible Leccinums that appear to
associate only with ericaceous plants such as manzanita and madrone, and are
consequently referred to collectively as “manzanita boletes.” Their ranks include a
common but unidentified (probably unnamed) species in our area with a bright orange to
rusty-brown cap, orange scabers, and flesh that tends to stain reddish before turning
fuscous (it is known locally as L. “aurantioscaber”), L. arbuticola, with reddish-staining
Leccinum manzanitae is a common associate of manzanita and madrone. Note the dark scabers on the
stalk and the white flesh that stains fuscous (smoky to smoky-purple) erratically when cut.

flesh and a buff to pale brown cap; L. aeneum, with a non-viscid, oranger cap;/,, largentii,
possibly associated with toyon, with a dry cap, dark olive-gray pores, and very dense
scabers; L. constans, with a pale pink to apricot-colored cap and unchanging flesh; L.
armeniacum, a common species with an orangish cap and somewhat paler scabers (especially
when young); and L. arctostaphylos, an Alaskan species associated with bearberry (a type of
manzanita). Other species: L.ponderosum(COLOR PLATE 146) is a sometimes massive
species that looks like L. manzanitae, but typically has unchanging (or only slightly
staining) flesh and is associated with conifers in the Pacific Northwest; it is quite tasty.
L. fibrillosum also grows with conifers in the Pacific Northwest, but it has a fibrillose
to fibrillose-scaly, liver-brown cap and its flesh stains reddish or vinaceous when cut.L.
idahoensis has a roughened, liver-colored cap but does not stain appreciably when
bruised, while L. brunneum has a dark brown to reddish-brown cap, stains directly to
fuscous, and is quite common under aspen in the Sierra Nevada and Southwest. See also
L. insigne and the species listed under it.

Leccinum insigne (Aspen Bolete) Color Plate 147

CAP (4) 6-17 cm broad, round to convex becoming broadly convex to nearly plane; surface
dry or only slightly viscid when wet, smooth to minutely fibrillose or sometimes breaking
up into small scales, in age sometimes pitted; color variable: bright orange to rusty-orange,
reddish-orange, orange-brown, reddish-brown, or cinnamon (or in one form brown),
often paler or duller (tan, brown, or dull orange-brown) in age; margin usually with flaps
of sterile tissue, at least when young. Flesh thick, often rather soft in age, white, turning
bluish-gray to purple-gray or fuscous when bruised or cut, but sometimes very slowly or
erratically. PORES and tubes pallid or whitish when young, becoming olive-buff to
grayish or dull yellowish-buff in age; not blueing when bruised (but may stain yellowish to
brown, lavender, or vinaceous). STALK 6-15 cm long, 1-2.5 cm thick at apex, equal or
swollen below, dry, solid, rather tough and fibrous; white or whitish when fresh, but
roughened by numerous small, projecting scabers which are initially pallid but become
reddish-orange to brown and finally blackish by maturity; base or lower portion often
staining or discoloring blue. SPORE PRINT brown to yellow-brown; spores 11-18 * 4-6
microns, spindle-shaped to elongated-elliptical, smooth.

540
LECCINUM 541

HABITAT: Widely scattered to gregarious or in troops on ground in woods and at their

edges; associated primarily if not exclusively with aspen, and common in the summer and
early fall wherever aspen occurs. It is abundant in the Sierra Nevada (along with other
Leccinums—see comments), and I have seen enormous fruitings in the Southwest and
Rocky Mountains shortly after the blue columbines bloom. It is also abundant in Alaska.
EDIBILITY: Edible and good. Though not the equal of Boletus edulis, it is far better in
flavor than its coastal counterpart, L. manzanitae. Like most Leccinums it darkens when
cooked or dried. If this intimidates you, that’s fine with me—I’ll be more than happy to take
them off your hands!
COMMENTS: This common aspen-lover has many look-alikes in the genus Leccinum,
most of which have passed under the name L. aurantiacum. They are very difficult to dis¬
tinguish without a microscope but all are apparently edible, though some may cause
digestive upsets in sensitive individuals. L. insigne is recognized by its rusty-orange to
reddish-brown or brown cap, scabers on the stalk which darken in age, and growth with
aspen. Some of its more common look-alikes include: L. aurantiacum, very similar and
widespread under aspen and conifers, but sometimes larger and more robust and with
flesh that distinctly stains burgundy-red before turning purplish- or bluish-gray; L. dis¬
color, very similar to L. aurantiacum, but growing only with aspen and with a slightly
duller, browner, or more cinnamon-colored cap (see p. 537 for the critical microscopic
difference); L. atrostipitatum andE. testaceoscabrum (the latter more brightly colored),
with scabers that are black even in the button stage, associated mainly with birch and
poisonous to some people; and L. fallax and L. subalpinum, with a rusty to dark reddish
cap and club-shaped stalk, associated with spruce and fir in the Rocky Mountains and
Southwest. Also see L. manzanitae and the species discussed under it.

Leccinum scabrum (Birch Bolete)

CAP 4-10 cm broad, convex to broadly convex to nearly plane or slightly depressed
centrally; surface smooth, dry to slightly viscid when wet, dull brown to grayish-brown to
dingy yellow-brown or tan (but often developing olive tints in age); margin lacking
flaps of sterile tissue. Flesh thick, white, typically not staining when cut or discoloring
slightly pinkish to brownish in the stalk. PORES and tubes dull whitish or pallid becoming
dingy brownish in age; not blueing when bruised but sometimes staining slightly yellowish
or ochre. STALK 7-15 (20) cm long, 0.5-1.5 (3) cm thick at apex, usually thicker below,
firm, solid, white to grayish but roughened by numerous projecting tufted hairs (scabers)
which soon become brownish to black at least over the upper portion; often staining blue-
green below. SPORE PRINT brown; spores 14-20 * 5-7 microns, spindle-shaped to
elongated-elliptical, smooth.
HABITAT: Solitary, scattered, or in groups near or under birch in the summer and fall;
common throughout the northern hemisphere wherever birch occurs. It has not been
found in California, but could conceivably appear on lawns where birches have been
planted (just like Lactarius torminosus).
EDIBILITY: Edible and choice according to some, mediocre according to others; only
firm specimens should be used.
COMMENTS: The dingy brownish cap, dark scabers on the stalk, and association with
birch are the principal fieldmarks of this bolete. There are several similarly colored species
that are best differentiated microscopically, but L. montanum of the Sierra Nevada can
be distinguished by its different mycorrhizal mate—aspen. Other species: L. griseonigrum,
associated with birch, is one of several species with a blackish to bluish-gray to dark brown
cap (which, however, may bleach out in age); L. snellii also has a dark cap but favors aspen; L.
alaskanum, common with birch in Alaska, has a dark and light, streaked or mottled cap.
Leccinum scabrum is one of many birch-loving Leccinums. Cap is dull colored (brown to tan, gray, or
dingy olive) and the stalk is usually rather slender.

There are also several species with a whitish or pallid cap, including: L. holopus and L.
rotundifoliae, both fairly common in northern North America under birch, the former
often developing olive tinges on the cap in age and the latter usually growing in bogs;
and L. albellum, a slender-stemmed, dry-capped eastern species that favors southern oaks.
L. californicum has a whitish to buff cap, but grows with aspen in the Sierra Nevada, while
L. cretaceum has a pale buff cap and flesh that stains vinaceous-pink erratically. L. roseo-
fracta has a dark brown cap and reddish- or pinkish-staining flesh and is associated with
birch. Finally, there is L. griseum, an eastern species that somewhat resembles L. scabrum
but usually has an areolate (cracked) cap in age and favors oak and other hardwoods over
birch. All of these species lack the flaps of sterile tissue on the cap margin characteristic of
L. insigne, L. manzanitae, and the species discussed under them.

Left: Leccinum holopus, a white-capped birch-loving species. Note dark scabers on stalk. Right:
Strobilomyces confusus (see comments under S. floccopus, p. 543, for details).

1
“Old Man of the Woods,” Strobilomyces floccopus, is easily told by its shaggy black or gray fruiting
body. Note cottony veil in young specimen at center and dark gray to black pores in older one at left.

STROBILOMYCES

THIS small genus is unique among the boletes in having a shaggy or scaly gray to black
fruiting body, gray to black pores when mature, and blackish-brown, reticulate or warty
spores. It is so unique, in fact, that some mycologists place it in a separate family. A woolly
veil is present when young, and may form a slight annulus(ring) on the stem. Since only one
species is described here, a key is not included.

Strobilomyces floccopus (Old Man of the W oods)

CAP 4-15 cm broad, convex or cushion-shaped becoming broadly convex to plane; surface
dry, shaggy from numerous soft, coarse, gray to grayish-brown or black scales on a paler
background (but in age sometimes entirely blackish); margin usually hung with pallid to
grayish veil remnants. Flesh whitish, usually turning reddish (often slowly) when bruised,
then dark brownish to black. PORES fairly large, white or gray becoming darker gray to
nearly black in age, usually bruising reddish, then brownish to black; tubes same color.
STALK 5-12 cm long, 1 -2.5 cm thick, equal or somewhat thicker toward base, solid, firm,
dry, sheathed by large gray to grayish-black scales or shaggy material except at apex. VEIL
pallid to grayish, woolly-cottony, leaving tissue on cap margin and usually one or more
shaggy zones on stalk. SPORE PRINT black to blackish-brown; spores 9.5-15 * 8.5-12
microns, broadly elliptical to nearly round, reticulate.
HABIT AT: Solitary to scattered or in groups in humus under hard woods (especially oak),
sometimes also with conifers; common in the summer and fall in eastern North America,
and according to Dr. Robert Gilbertson of the U niversity of Arizona, also fairly common
in southern Arizona. Its dark color makes it hard to see in the forest gloom.
EDIBILITY: Edible, but not rated highly by most mushroom enthusiasts. I found it
rather insipid although the texture was good. Only young specimens should be used.
COMMENTS: This most distinctive of all the boletes is easily told by its shaggy gray to
black fruiting body. In contrast to most boletes, it does not decay readily. As a result, it
often seems more numerous than it actually is. Other species: S. strobilaceus is a synonym.
S’, confusus of eastern North America is quite similar in color and overall aspect, but has
more erect scales on the cap (see photo on p. 542) and spiny-warty to only partly reticulate
spores; S. dryophilus is a southern species that is similarly colored in age but usually paler
(cap pinkish-tan to brown) when young.
543
544 BOLETACEAE

GASTROBOLETUS (Gastroid Boletes)

Small to medium-sized boletes usually buried or partially buried in humus. CAP convex to some¬
what irregular in shape. PORES variously colored; tubes often irregularly arranged, i.e., not all
vertically oriented. STALK usually short, poorly developed, sometimes off-center. VEIL absent
or present as persistent membrane that covers the tubes. SPORE PRINT unobtainable. Spores
elliptical to spindle-shaped, smooth, brown to golden-brown under the microscope.

THESE funky fungi have been modified or “reduced” from normal, everyday, upright
boletes to abnormal, semi-underground, “decadent” ones. They resemble puffballs in that
they do not forcibly discharge their spores, yet they retain a modicum of respectability in
the form of a rudimentary stalk and definite tube layer arranged rather haphazardly
within the misshapen cap.
Gastroboletus is a small genus, with only one species common along the coast. How¬
ever, several species occur in the Sierra Nevada and Cascades—perhaps in response to
the more severe and unpredictable weather. Fruiting underground helps the mushrooms
to withstand the pendulum-like hot-and-cold or warm-and-dry spells. None are known to
be poisonous, yet none are known to be edible, since none are known to have been tried.

Key to Gastroboletus
1. Tubes and flesh blueing when bruised or cut .G. turbinatus & others, below
1. Not as above .2
2. Stalk relatively long (6-10 cm) and roughened by small scales or scabers; known only from
eastern North America . G. scabrosus
2. Not as above . 3
3. Cap whitish to buff or pale tan (sometimes slightly darker in age); tubes whitish to buff to olive-
brown, or grayish-yellow; stalk averaging 2-5 cm thick . G. subalpinus, p. 545
3. Not as above; cap pale or dark; tubes yellow to olive-yellow; flesh often yellow; stalk typically
1.5 cm thick or less .G. suilloides& G. amyloideus (see G. turbinatus, below)

Gastroboletus turbinatus (Bogus Boletus; Gastroid Bolete)

CAP 2-5 (8) cm broad, convex to slightly depressed to irregular (misshapen); surface dry,
minutely velvety, often pitted or wrinkled; color variable, but usually brown to reddish-
brown to bright yellow with red or brown spots and stains. Flesh soft, yellow (but often
reddish in base of stalk), typically bruising blue quickly. PORES pinkish to reddish,
orange, or yellow, typically blueing quickly when bruised; tubes greenish-yellow to yellow,
irregular and sinuous or disoriented (arranged at various angles rather than exclusively
vertical). STALK \ A (7) cm long, 1-2.5 cm thick, equal or narrowed at base, often(but
not always) protruding only a short distance below the tubes; solid, dry, yellow or with
reddish streaks or granules or colored like the cap, sometimes reddish in old age and often
reddish at the base. SPORE PRINT unobtainable; spores(9.5) 13.5-18 * 6.5-9.5 microns,
elliptical to spindle-shaped, smooth, brown or golden under the microscope.

HABITAT: Solitary to scattered or gregarious on ground in woods or at their edges, often

partially buried; widely distributed. It is fairly common in the summer and fall under
conifers in the Pacific Northwest, northern California, and the Sierra Nevada. I have
found it twice in our area under live oak, in July and October (it is not necessarily rare,
however—just easy to overlook).
EDIBILITY: Unknown, like myself, and likely to remain so.

COMMENTS: A curiously wrought fungal afterthought, easily recognized by its irregular

shape, semi-underground growth habit, and disoriented tube layer. Since the spores are
not forcibly discharged, there is no need to elevate the cap above the ground. Just how
the spores are spread is a mystery—perhaps with the help of boletivorous bipeds such as I.
Left: Gastroboletus turbinatus. These two specimens have unusually well-developed stems, but the
misshapen caps are typical. Right: Gastroboletus xerocomoides. This specimen has been sliced open
to show the disoriented (haphazardly arranged) tube layer. (Herb Saylor)

Many agarics have been “reduced” to gastroid forms (see Podaxales & Allies, p. 724);
this is one instance in which a bolete has. Most of the other members of the genus have
limited distributions. Several occur under conifers in the Sierra Nevada, including:
G. xerocomoides, which blues only slightly and has mostly truncate spores, and three
species which do not blue when bruised: G. suil/oides, with a brown cap; G. amyloideus,
with a yellow, buff, or reddish-spotted cap and amyloid spores; and G. subalpinus{bz\ovi).

Gastroboletus subalpinus (Gastroid King Bolete)

CAP 5-12 cm broad, convex becoming plane to depressed or somewhat irregular; surface
smooth to slightly velvety, often pitted and/ or wrinkled, not viscid but often covered with
dirt and debris; whitish to buff or pale tan, sometimes darkening somewhat in age; margin
extending downward and over the tubes as a thin white membrane that often disappears
or breaks up in age. Flesh thick, soft, white or with yellow to olive discolorations; not
blueing when bruised but sometimes staining pinkish. PORES pallid or whitish becoming
grayish-yellow to buff and finally darkening to olive-brown, not blueing when bruised;
tubes occasionally oriented vertically (especially near stalk) but mostly arranged hap¬
hazardly (at various angles); colored more or less like pores. STALK 2-6 cm long, 2-5
cm thick, tapered downward or sometimes equal or even swollen below; solid, dry (but
often dirty), white to buff, sometimes darkening to tan in age; base sometimes orange-
yellow. SPORE PRINT unobtainable; spores 10-16(18) * 4.5-6 (8) microns, elliptical to
spindle-shaped, smooth, yellowish to brown under the microscope.
HABIT AT: S olitary to gregarious in soil or duff under mountain conifers in the late spring
and summer; known only from the Sierra Nevada (where it is fairly common) but to be
expected in other mountain ranges of the West. It develops underground but is often
partially exposed (erumpent) at maturity.
EDIBILITY: Presumably edible; I haven’t tried it.

COMMENTS: This gastroid bolete is easily told from its brethren by its pale color and
white, non-blueing flesh. It is likely to be mistaken for an aborted button of Boletus edulis,
or more likely still, B. barrowsii. However, the disoriented tube layer, tendency to develop
underground, and refusal to give a spore print are distinctive. The thin layer of tissue
which at least partially covers the tubes of young specimens may represent a more elabo¬
rate version of the whitish pith that plugs the tubes of young B. edulis and B. barrowsii.
546

BOLETIVORES
THESE curious creatures, as their name suggests, are a major predator of Boletus edulis.
Since they are likely to be encountered by anyone who looks for mushrooms, it seems
appropriate to describe them and include a few observations on their habits.
Boletivores appear shortly after the onset of the rainy season, not coincidentally, at
the same time Boletus edulis appears. As a group they share a number of telltale traits
which the experienced observer can discern at a glance. Their overall appearance is rather
dingy and disheveled, their color drab. The cap is quite variable, ranging from depressed or
deflated to uplifted, ecstatic, or inflated; its surface may be dry and distinctly tomentose
or smooth and somewhat viscid. The flesh is normally pallid, becoming blue when bruised
and exuding a red latex when cut, but it is most often obscured by a ragged woolly cuticle
which is typically appressed above and baggy below. The stalk may be fleshy or cartilagi¬
nous, and more often than not, bent. The odor varies considerably from individual to
individual; the taste is unpleasant, and a spore print is not obtainable, at least with normal
methods. Boletivores are strictly terrestrial, and are invariably equipped with a long
pointed stick, rapacious eye, and capacious bucket or “universal pail.” At least two
distinct species occur in our area; these are keyed below.

Key to the Boletivores

1. Retreating furtively when approached .Boletivorus clandestinus, below
1. Advancing boldly when approached . Boletivorus brutalosipes, below

Boletivorus clandestinus is the more common of the two species. In addition to

its secretive nature, it can be recognized (if you can get close enough!) by its tough,
wizened stalk with a persistent, even permanent, stoop. The gait is also highly distinctive:
curiously hitched and truncated, marvelously efficient, yet flailing and disjointed. The
overall impression is that of a creature completely immersed in, yet not designed for, its
element—like a kayaker trying to negotiate a rapids without wetting her back. When one
is crept up on unawares, it can be heard alternately cooing to and swearing at its pro¬
spective prey in a vaguely familiar, yet unintelligible, tongue. This species occurs
solitary to scattered or in small groups in the woods, but always near roads. There is usually
a rusty pick-up truck, station wagon, or ’65 Dodge Dart nearby, parked on the side
of the road. Near the vehicle will be found one or more telltale “middens”—neat piles of
discarded tubes and wormy stalks. These resemble the feathers that remain from a freshly
and systematically disemboweled bird—positive proof of boletivores’ singular lust
and unbridled craving for their quarry.
Boletivorus brutalosipes (the “Brutal-Footed Boletivore”), on the other hand, can be
instantly recognized by its inquisitiveness, its bold advance and cavalier stance. In
addition, the stalk is more fleshy than that of Boletivorus clandestinus—often swollen in
the middle or even bulbous. The gait is decidedly more compact, the stride purposeful,
yet completely arbitrary. When one comes rushing toward you, the net effect is that of a
rapids intent upon wettingyoi/r back. Its insatiable greed for Boletus edulis may be cleverly
disguised by an amicable disposition and disarmingly friendly, fibrillose smile. But it will
stop at nothing to achieve its ends, so never leave your basket unattended in the woods!
If there are Boletus edulis in it, they will be gone, and other species will be stepped on,
masticated and regurgitated, or otherwise obliterated. Trampled fly amanitas (Amanita
muscaria), incidentally, are a sure sign that brutal-footed boletivores are in the vicinity!
Boletivorus brutalosipes is found in roughly the same habitats as Boletivorus clandestinus,
A telltale “midden” left by Boletivorus clandestinus.

but fortunately, is not as common. Invariably there is a rusty pick-up truck, station
wagon, or ’65 Dodge Dart nearby, parked in the middle of the road, and the discarded
tubes and wormy stalks of its quarry are apt to be haphazardly strewn about rather than
stacked in the “middens” characteristic of B. clandestinus.
Boletivorus clandestinus is probably harmless, though I haven’t been able to get close
enough to find out. Boletivorus brutalosipes, on the other hand, has a well-deserved
reputation for unprovoked acts of aggression. If you have some Boletus edulis in your
basket, watch out!
Between these extremes, as might be expected, “hybrids” occur, in addition to possibly
autonomous species* (I have even seen a skinny, blonde, bare-stemmed variety). However,
further study—and preferably, a critical comparison with European specimens—is
necessary before a definitive and natural classification can be worked out. Both species
occur throughout the range of Boletus edulis. On weekends they are particularly abundant
on lower Empire Grade above Santa Cruz, California, where their crafty cousin, Pseudo-
boletivorus incognitus, is also in evidence. (The latter closely mimics Boletivorus clandes¬
tinus, but only pretends to be stalking Boletus edulis—the true object of its desires being
Amanita calyptrata.)
As a final note, I might add that what boletivores do during the summer is a mystery. It
has been suggested by one colleague of mine that they hibernate in the trunks of their cars.
Another possibility is that they follow the bolete season up and down the coast. A third is
that they migrate east—or west. But perhaps the most reasonable hypothesis is that they
are like squirrels—they stash away what they gather in hedges, holes, nests, or sheds,
for when the hunting is good, they have neither the time nor the inclination to eat. Then,
in the summer, when they have nothing but time, they do nothing but eat.

*In the newsletter of the Mycological Association of Washington, D.C., Anne Dow announces the discovery of
a third species, Boletivorus europaeus, in the Tetons of Wyoming. Like B. brutalosipes, it “advances boldly when
approached,” but it “fills [rather than empties] the baskets [sic] of other boletivores” (or does she mean other
bolete hunters?). I have never encountered such a species in California, but would welcome its introduction!

547
548

APHYLLOPHORALES
THIS large and diverse assemblage of fleshy—and not-so-fleshy— fungi includes all the
Hymenomycetes (p. 57) with the exception of the agarics, boletes, and jelly fungi. The
fruiting body is not normally gelatinous (as in the jelly fungi), there is no veil (as in many
agarics and boletes), and the basidia are typically simple and clublike. The hymenium
(spore-bearing surface) can be completely smooth and unspecialized as in the crust and
parchment fungi, or it can take the form of tubes (the polypores), spines or “teeth” (the
teeth fungi), upright clubs or branches (the coral fungi), or veins, wrinkles, or even rudi¬
mentary gills (the chanterelles).
More than twenty families in the Aphyllophorales are now recognized, but many of
the distinguishing criteria are esoteric. To facilitate field identification, the Aphyllo¬
phorales have been divided into five major groups (plus one aberrant species), keyed
below. The coral fungi and chanterelles (Clavariaceae and Cantharellaceae) are placed
in a separate order, the Cantharellales, by some taxonomists.

Key to the Aphyllophorales

1. Fruiting body with a layer of downward-pointing spines or “teeth” on underside of cap or with
icicle-like spines hanging from branches or a cushion of tissue .Hydnaceae, p. 611
1. Not as above (but fruiting body may have pores whose walls become torn or toothlike in age) 2
2. Fruiting body with a layer of tubes(usually on underside of cap); tube mouths(pores) large and
sinuous to small or very minute .Polyporaceae & Allies, p. 549
2. Not as above; tubes and pores absent (but spore-bearing surface may have veins that give it a
somewhat poroid appearance) . 3
3. Fruiting body consisting of a small (up to 2.5 cm) hollow, bladderlike cap mounted on a thin
stalk; surface of cap smooth or wavy, whitish to yellowish or pale brownish; found on ground
or wood in eastern North America, often in groups or clusters .Physalacria inflata
3. Not as above; common and widely distributed .4
4. Fruiting body usually bracketlike, crustlike, or sheetlike (i.e., usually without a stalk and often
without a cap); usually growing on wood or herbaceous stems .. Stereaceae & Allies, p. 604
4. N ot as above; fruiting body upright, either simple and clublike or branched (coral- or bushlike)
or with a cap (often vase-shaped) and stalk; usually on ground, sometimes on wood .5
5. Fruiting body with a cap (i.e., a clearly differentiated, usually flattened or depressed upper
sterile surface) and stalk . Cantharellaceae, p. 658
5. Fruiting body clublike or branched or a rosette-like mass of flattened, white to creamy or tan
lobes or segments (i.e., without a well-defined cap) . Clavariaceae, p. 630

Bondarzewia montana (see description on p. 565). When these compound fruiting bodies were
younger, they were whitish and resembled coral fungi (e.g., Sparassis) but for the presence of pores.
As they mature, however, the upper lobes or “branches” broadened into tan or brown caps. This
species is also capable of producing simple fruiting bodies (with just one cap and stalk).
 549

Polypores and Bracket Fungi

POLYPORACEAE & Allies

THIS large and exceedingly diverse group comes in a mind-boggling multiplicity of

shapes and sizes and a pleasing array of gay decorator colors. As the name “bracket fungi”
implies, the fruiting body is most often bracketlike or shelflike. “Polypore” (meaning
“many-pored”) describes the spore-producing tube layer which lines the underside of the
cap. However, the tube mouths or pores are sometimes so minute that they’re virtually
invisible without a hand lens, and in some bracket fungi the pores are replaced by large
mazelike pockets or even gills and in others the tube walls break up to form “teeth.” The
boletes are also equipped with tubes, but they are rapidly-decaying, mostly terrestrial fungi
with a soft fleshy cap, central stem, and tubes that usually peel away easily from the cap.
Polypores, in contrast, grow mostly on wood and have a tube layer which is scarcely
detachable. The most familiar types fruit on logs and stumps in dense, shelving masses
(or in ascending tiers if the tree is still standing). Others, called conks, are those hard,
woody, hooflike growths you see on living trees. Still others are resupinate: with neither
cap nor stem, they lie flat on the wood or incrust it. Those that grow on the ground often
originate from roots or buried wood, and their stem is usually off-center or even lateral.
Polypores are usually described as being tough, leathery, or woody. This is true of old
specimens, but fresh individuals of some species, such as Phaeolus schweinitzii, are so soft
and watery they can be wrung out like a sponge. Others, like the beefsteak fungus (Fistulina
hepatica) and the sulfur shelf (Laetiporus sulphureus) “weep” in wet weather, exuding
colored water droplets. In contrast, Fomitopsis and Ganoderma can be woodier than the
wood they feed on!
Like most fungi, polypores fruit during the rainy season. However, mature fruiting
bodies are so tough they tend to dry out instead of decaying. They may thus persist for
months, even years. Some (e.g., Ganoderma, Fomitopsis, Phellinus) are actually
perennial: rather than going to the trouble of manufacturing new fruiting bodies every
year, they quite sensibly make use of existing resources by adding a new growth layer onto
the old one. In this manner they may attain gargantuan dimensions—a 5 ft.x 3 ft. specimen
of Oxyporus nobilissimus from the Pacific Northwest weighed 300 pounds, undoubtedly
a record for fleshy fungal fructifications!
There is much more to polypores than their fruiting bodies, however. One cannot fully
appreciate them without appreciating their work—that is, the chemical and structural
changes they produce in their hosts. Polypores are absolutely indispensable to the forests
of this world. They are the major group of wood-rotting fungi. Though they wreak
economic havoc (15-20% of all standing timber in this country is said to be defective or
unusable because of fungal decay, more than 90% of it caused by bracket fungi; struc¬
tural timber is also destroyed—in ships, mines, houses, bridges, etc.), the polypores
should not be seen as enemies. Without them there would be no logging industry in the
first place; every cut stump, felled log, and lopped-off limb would lie indefinitely on the
forest floor, the woods would quickly become impenetrable, and new trees would have
neither room nor nutrients to grow.
In living trees the dead tissue (heartwood) in the center of the trunk is more susceptible
to attack than the living tissue (sapwood) beneath the bark. Trees infected by heart rot may
look outwardly sound, even when they are completely hollowed out inside. Sometimes
the only clue to the pernicious presence of the rot is the fruiting body of the fungus! Since
heartwood is largely dead, heart rot fungi are technically not parasites, but they can be
deleterious—the infected trees are steadily weakened until they come crashing down at
550 POLYPORACEAE & ALLIES

the slightest provocation. (So remember not to breathe too hard when picking polypores
from a standing tree!) A few species, such as Heterobasidion annosum, are virulent
parasites, destroying both the sapwood and the heartwood.
Spores generally gain access to living trees through wounds in the bark. Trees bruised or
battered by wind, nearby construction, careless machinery, or mindless vandalism (“Jody
loves Judy”) are especially vulnerable. Sometimes spores enter through holes bored by
grubs or beetles, which may in turn munch on mycelium growing in their tunnels.
Heart rot continues after the tree dries, and sap rot sets in. On fallen logs and branches
you will find large clusters of colorful bracket fungi (e.g., Trametes versicolor and
Trichaptum abietinus) which are digesting the sapwood. Several varieties may simul¬
taneously or successively inhabit the same log. Recently felled timber is also susceptible
to attack. Only complete immersion in water will prevent infection.
Wood is composed largely of dead, empty cells from which the fungus extracts nutrients.
The cell walls have two principal constituents: cellulose, which makes the wood soft and
tough, and lignin, which makes the wood hard and brittle. Fungi which digest the cellulose
and leave the lignin behind are called carbonizing decays or brown rots because they
render the wood dry, brittle, and darker than normal wood. Fungi which digest cellulose
and lignin are called delignifying decays or white rots because they make the wood soft,
spongy, and whiter than normal wood. Some species may completely hollow out the trunk
or produce small, hollow pockets called pocket rot. The mycelial threads of bracket fungi
can often be seen if the wood is broken open and examined closely. But if more than one
species inhabit the same piece of wood, interpretation and diagnosis become complicated.
Only repeated observation can tell you which ones produce which types of rot.
The part of the tree infected should also be noted. Butt rots such as Phaeolus schweinitzii
are confined to the roots and base of the tree and the fruiting bodies are often found on the
ground. Trunk rots like Phellinus pini infect the entire trunk, while top rots such as
Fomitopsis rosea inhabit the top of the trunk.
Practically all of the polypores were originally lumped together in one giant genus,
Polyporus. Dozens of families and genera have subsequently been proposed in a laudable
effort to break the group down. As usual, however, there are widely divergent opinions on
exactly how the task should be accomplished. The result is scholarly chaos, and the goal
of arriving at one name for each kind of organism is still a long way off. T o give you an idea
of the difficulties faced by a prospective polyporologist (or any toadstool taxonomist)
when she sets about consulting the literature on her pet polypore, here is a list of names
given to Trametes occidentalis by various investigators:
Polyporus occidentalis, Coriolus occidentalis, Polystictus occidentalis, Microporus
occidentalis, Coriolopsis occidentalis, Boletus sericeus, Trametes lanata, Polystictus
lanatus, Microporus lanatus, Polyporus lanatus, Polyporus lenis, Polystictus lenis,
Microporus lenis, Trametes wahlenbergii, Trametes scalaris, Polystictus scalaris,
Polystictus cyclodes var. homoporus, Polystictus scorteus, Microporus scorteus,
Polyporus gourliaei, Fomes gourliaei, Scindalma gourliaei, Trametes hispidula,
Trametes devexa, Polystictus malachodermus, Polystictus substrigosus, Polyporus
illotus, Polystictus illotus, Microporus illotus, Daedalea subcongener, Polystictus
subcongener, Trametes heteromalla, Polystictus extensus, Polyporus badiolutescens
. . . (I’m suddenly very grateful that I’m not a taxonomist!)

Several hundred different polypores are known from North America; perhaps half of
them occur in California. Unfortunately, the vast majority are too tough or too bitter
(or too tough and too bitter) to eat. Two happy exceptions are the sulfur shelf (Laetiporus
sulphureus), a succulent and unmistakable treat, and the beefsteak fungus (Fistulina
hepatica), which looks like a slab of steak. The hen of the woods, Grifola frondosa, and its
look-alike, G. umbellata, are also excellent, but do not seem to occur in our area. A few
polypores, however, may actually be poisonous (e.g., Phaeolus schweinitzii), and even
the edible species are difficult to digest unless cooked thoroughly.
POLYPORACEAE & ALLIES 551

Since so few bracket fungi are edible, they are ignored by most people and barely
mentioned in many popular mushroom books. As I am also ignored by most people (and
have yet to be mentioned in any kind of book), I feel I have something vital in common
with these unheralded but indispensable organisms. Therefore, a fairly extensive (but by
no means comprehensive) treatment is offered here, in hopes that readers will at least learn
to notice polypores, if not identify them. In the following key they are broken down into
several groups based on field characters. If you have trouble with some of the choices, you
can take solace in the fact that almost everyone finds the polypores difficult to identify!

Key to the Polyporaceae & Allies

1. Spore-bearing surface composed of closely-packed but discrete (separate) tubes that look like
small pipes (use a hand lens if unsure and see photo at top of p. 553); flesh usually marbled
or streaked when sectioned; fruiting body fleshy (not hard and woody) and often exuding a
bloodlike juice when moist; found on or around hardwoods . Fistulina, p. 553
1. Not as above; spore-bearing surface not composed of tubes, or if composed of tubes then the
tubes forming a united layer (i.e., not discrete); fruiting body fleshy, tough, woody, etc. . . 2
2. Fruiting body with a stalk(s) and cap(s) (but may appear fingerlike when emerging); found on
ground, or if on wood then stalk usually central or off-center; cap and stalk not varnished 3
2. Fruiting body knoblike, hooflike, bracketlike, shelflike, or crustlike; stalk absent, rudimentary,
or attached to side or top of cap (or if central, then varnished); growing on wood or roots 4
3. Flesh yellow-brown to tawny, rusty-brown, or brown (or sometimes dark yellow, but if so then
associated with conifers); flesh darkening (staining red to vinaceous-brown or black) in potas¬
sium hydroxide (KOH); pores often colored like flesh; usually growing on ground (often
around stumps or trees or on roots) .Phaeolus, Inonotus, Coltricia, & Allies, p. 566
3. Not as above; flesh usually white or pallid to pale yellow when fresh (in one case yellow to olive-
yellow, but if so then associated with hardwoods in eastern North America), but sometimes pale
brownish or otherwise discolored in age or where bruised; pores usually colored like flesh when
fresh; on wood or ground .Polyporus, Albatrellus, & Allies, p. 554
4. Pore surface completely enclosed (hidden) by a tough membrane (i.e., fruiting body with a tube-
lined internal cavity) or fruiting body growing on birch and possessing a blunt projecting
margin that forms a “curb” around the pore surface .... Piptoporus& Cryptoporus, p. 584
4. Pore surface exposed; not growing on birch, or if on birch then margin not curblike .5
5. Tube layer elastic and waxy to somewhat gelatinous when fresh, often separating easily from
rest of fruiting body; pores usually pinkish to reddish-purple to blackish-purple (but occa¬
sionally white to buff); cap white and hairy (when present) . 19
5. Not as above; pore surface differently colored and/ or not separable and elastic-gelatinous 6
6. Fruiting body typically resupinate (i.e., lying flat on the wood or incrusting it), consisting of a
simple layer of tubes; cap (sterile surface) absent or rudimentary .... Poria & Allies, p. 602
6. Fruiting body normally with a cap (upper sterile surface), but sometimes resupinate, especially
if growing on the undersides of logs . 7
7. Spore bearing surface composed of shallow veins which may form very broad “pores” or
pits . (see Stereaceae & Allies, p. 604)
7. Spore-bearing surface comprised of a true tube layer which forms minute to fairly large pores
or spore-bearing surface with deep, elongated, mazelike pockets or even gills or “teeth” . . 8
8. U nderside of cap (spore-bearing surface) with gills .9
8. Spore-bearing surface with tubes (pores), but the pores sometimes elongated or mazelike or
breaking up to form small “teeth” . 10
9. Gills appearing to be split lengthwise (along their edges), but actually composed of two adjacent
plates which curl back in dry weather (see photos on p. 591) .Schizophyllum, p. 590
9. Not as above .Lenzites, Daedalea, & Allies, p. 586
10. Fruiting body annual (with only one tube layer), the cap soon dark brown to blackish or bluish-
black with a resinous, often roughened (sandpaper-like) and/or radially wrinkled surface;
fruiting body at first watery and often beaded with droplets, but tougher in age; pore surface
whitish at first but aging or bruising darker (brownish); found on dead trees (logs, stumps,
etc.); widespread but not common .Ischnoderma, p. 573
10. Not with above features (but may have some of them); very common . 11
Fistulina hepatica, a young specimen in which the cap is still velvety. It’s easy to see why it is sometimes
called the “Ox Tongue.” A mature specimen is shown in the color plate. (Ralph Buchsbaum)

11. Fruiting body punky, corky, or hard and woody even when fresh, medium-sized to very large
and often thick; usually perennial, i.e., with more than one tube layer, but if annual then often
with a varnished surface crust . Ganoderma, Fomitopsis, Phellinus, & Allies, p. 574
11. Not as above; fruiting body usually annual, small to medium-sized or if large then usually fleshy
or spongy when young and fresh; if tough when fresh then cap usually fairly thin and lacking
a highly varnished surface crust . 12
12. Flesh soon red to orange, rusty-brown, tawny, yellow-brown, or dark brown and darkening
(staining red to vinaceous-brown to black) in potassium hydroxide (KOH); pores variously
colored (including red, orange, and mustard-yellow), but not rosy, purple, or bright sulfur-
yellow)* .Phaeolus, Inonotus, Coltricia, & Allies, p. 566
12. Not as above; flesh white to yellow, yellowish, beige, or sometimes light brown or salmon-
tinged, not darkening in KOH* . 13
13. Underside of cap with elongated pocketlike or mazelike pores (see photos on pp. 586, 588)
.Lenzites, Daedalea, & Allies, p. 586
13. Not as above (but pores may break up to form “teeth”) . 14
14. Fruiting body usually preceded by a small, often zoned cup- or saucerlike “nest” (0.5-2 cm
broad) from which small whitish to yellowish shelves (fruiting bodies) develop; common on
dead hardwoods, especially elm, in eastern North America ...Poronidulus conchifer
14. Not as above . 15
15. Cap only 1-5 mm broad, with a stubby or knoblike stalk attached to the top; fruiting body white
to brownish; common on hardwoods (rarely conifers) in eastern states Porodisculuspendulus
15. Not as above; usually larger, widely distributed . 16
16. Pore surface white to buff to yellowish or bright sulfur-yellow when fresh (but may stain blackish
or rusty-reddish in age or when bruised) . 17
16. Pore surface red to orange, orange-yellow, lavender- or violet-tinged, rosy, gray, black, or
brown, or soon becoming these colors .Trametes & Allies, p. 592
17. Pore surface bright sulfur-yellow when fresh (or rarely white, and if so then cap salmon-colored);
cap orange-red to bright orange, bright yellow, or salmon-colored (but fading), usually at least
6 cm broad when mature .Laetiporus, p. 572
17. Not as above . 18
18. Cap thin and tough (but pliant) even when fresh, typically tearing easily in a radial direction;
cap surface usually hairy or velvety and often concentrically zoned; often fruiting in large
masses; very common, especially on hardwoods .Trametes & Allies, p. 592
18. Cap soft, watery, spongy, or fleshy when fresh (but often tough in age), or if tough when fresh
then not as above; cap often but not always thick, the surface sometimes hairy but not usually
zoned; usually (but not always) found in smail numbers . Tyromyces & Allies, p. 597
19. Pores averaging 4-8 per mm; found mainly on hardwoods Caloporus (-Gloeoporus) dichrous
19. Pores averaging 2-4 per mm; tubes very shallow; found on dead conifers Skeletocutis amorpha

*Several household cleaning agents (e.g., Drano, 1 teaspoon per % cup water) can be substituted for KOH, but
unless the flesh is borderline in color (i.e., dark yellow or light brown) the test is usually unnecessary. If there’s
any doubt, simply try both choices!
552
Left: A close-up of the tubes in Fistulina hepatica. Note how bruised area is darker. Right: An even
closer view, in which the tubes can be seen as discrete units or “pipes”—a unique feature of Fistulina.

FISTULINA

THIS genus contains only one common species, the beefsteak fungus, F. hepatica. It is not
a “true” polypore because the tubes, though packed together closely, are discrete units
arranged like the bristles of a brush (see above photos). This character, which is best ob¬
served with a hand lens, has led polyporologists to give Fistulina a family of its own.

Fistulina hepatica (Beefsteak Fungus; Ox Tongue) Color Plate 152

FRUITING BODY annual, at first knoblike, then cushion- or shelflike; juicy when
fresh, soft to fairly firm. CAP 7-30 cm broad and 2-6 cm thick when mature, tongue-shaped
or fan-shaped; surface minutely roughened and suedelike or velvety, but often gelatinous
when wet, firm or very soft and spongy, sometimes knobby; orange-buff to reddish-orange
to pinkish, reddish-brown, dark red, or liver-colored; margin often lobed. Flesh thick,
soft and watery when fresh and oozing with dark reddish juice; whitish to grayish, olive-
brown, or reddish, streaked or marbled horizontally with paler veins (usually reddening
after standing); taste rather sour or strongly acidic. PORES 1-3 per mm, circular, whitish
to yellowish, buff, or pale flesh-color, becoming dark reddish-brown when bruised or in
old age; tubes 10-15 mm long, closely packed but discrete (their walls not joined). STALK
absent, or if present then lateral and rather short, 2-6 (10) cm long, 1-4 cm thick; tougher
than the cap but continous with it and colored similarly, firm. SPORE PRINT pinkish-
salmon to pale ochre or pale rusty-brown; spores 4-6 x 3-4 microns, elliptical, smooth.
HABITAT: Solitary or sometimes several together at the bases of hardwoods and on
stumps, especially of chestnut and oak; widely distributed, but most common in Europe
and eastern North America. In coastal California it seems to grow exclusively onchinqua-
pin (a close relative of chestnut) in the fall and winter. In my experience it is not common,
but Chroogomphus-king Ciro Milazzo discovered one large local fruiting of over thirty
individuals on some recently cut chinquapin stumps. It causes a serious carbonizing decay
in its host. The rich brown hue it imparts to the wood is prized by cabinetmakers.
EDIBILITY: Edible. The strong sour taste is displeasing to some, esteemed by others.
Parboiling may remove some of the sourness as well as some of the nutrients (it is rich in
vitamin C). I like to marinate thin, raw slices in seasoned vinegar and olive oil.

553
Fistulina hepatica, sliced open to show the streaked, meatlike flesh. Fresh specimens even exude a
bloodlike juice. For other views of this photogenic fungus, see color plate and photos on pp. 552-553.

COMMENTS: This remarkable fungus looks like a slab of raw meat, especially after it
is sliced open and starts oozing “blood.” The streaked or marbled flesh is very distinctive,
plus it never becomes as tough as other polypores and bracket fungi. The fruiting bodies
seem to develop very slowly and are rarely attacked by maggots. Other species: Pseudo-
fistulina radicata (-F. pallida) of eastern North America and Mexico also has discrete
tubes, but the cap is grayish-brown to pale reddish and the stalk is usually longer(to 15 cm)
and somewhat rooting. It is edible also.

POLYPORUS, ALBATRELLUS,& Allies

(Stalked Polypores)
Fruiting body annual, fleshy to tough, with a cap and stalk or sometimes compound (with many
caps and stalks); growing on ground or on wood. CAP convex to plane, depressed, or misshapen.
PORES large to very minute, often decurrent on the stalk; tubes usually not cleanly separable from
the cap. ST ALK central to off-center or sometimes lateral, usually well-developed but sometimes
stubby. SPORE PRINT usually white (when obtainable), but sometimes brownish in Boletopsis.
Spores smooth, warty, or spiny; amyloid in Bondarzewia, otherwise not amyloid.

THESE are fleshy to rather tough polypores with a clearly defined cap, pore surface,
and stalk. As such they are easy to recognize—the presence of a stalk separates them from
most other polypores, and confusion with the boletes is unlikely because boletes have a soft,
fleshy texture, central stalk, and tubes which are usually longer (deeper) and peel away
easily from the cap (and often from each other).
The genus Polyporus once embraced practically all the polypores or woody pore fungi.
The “splitters” have steadily whittled away at it, however, so that it now contains a modest
number of stalked polypores with white, cylindrical, smooth, non-amyloid spores. Most
of its species grow on dead hardwoods, but some arise from roots or underground “tubers”
and thus may appear terrestrial. Albatrellus, on the other hand, is one of the few groups
of polypores that is truly terrestrial, fruiting in sometimes massive numbers under both
hardwoods and conifers. Microscopically it differs from Polyporus in having elliptical
to nearly round spores which may or may not be amyloid.
To facilitate identification, a number of other stalked polypores are treated here as well,
though their similarity to Polyporus and Albatrellus may be entirely superficial. These
include: Boletopsis, terrestrial like Albatrellus, but with warted spores; Heteroporus, with
a frequently misshapen fruiting body that is often entirely covered with pores; Grifola, with
a compound, intricately branched fruiting body that gives rise to many small overlapping
caps; and Bondarzewia, with a frequently compound fruiting body and warted amyloid

554
POLYPORUS, ALBATRELLUS, & ALLIES 555

spores. The latter two genera can be reminiscent of coral fungi (especially Sparassis), but
are easily differentiated by the presence of pores on the underside of each cap or “lobe.”
Tender specimens of Grifola are delicious when thoroughly cooked, but most of the
other polyporps treated here are too tough or chewy to be more than marginally edible.
None are known to be dangerously poisonous, however. Fourteen species are described
here and several others are keyed out.

Key to Polyporus, Albatrellus, & Allies

1. Fruiting body often misshapen and covered in various areas (or almost entirely) by pores;
pore surface usually reddening when handled .Heteroporus biennis, p. 566
1. Pores confined to underside of cap (or decurrent on stalk), reddening in only a few cases .. 2
2. Fruiting body wood-inhabiting (on wood or roots or near the bases of trees or stumps) ... 3
2. Fruiting body truly terrestrial (growing on ground) . 13
3. Stalk black or with a black base, and only one cap; pores minute (averaging 3-7 per mm) . . 4
3. Stalk not black at base, or if so then pores larger and/or fruiting body compound .5
4. Cap reddish-brown to chestnut-brown to blackish; stalk usually black .... P. badius, p. 562
4. Cap tan to ochre or paler; stalk often black only at base .P. elegans & others, p. 562
5. Flesh very dense and heavy, becoming bone-hard as it dries out; cap(s) white to gray, buff, or
even brownish, 2-10 cm broad, often in fused clusters but not truly compound; pores white to
yellowish; found on dead conifers or rarely birch; not common Osteina obducta (-O. ossea)
5. Not as above (larger or with different habitat or color, etc.); not drying bone-hard .6
6. Fruiting body with one cap, small to medium-sized (cap usually 1-8 cm broad), flesh usually
2 mm thick or less; stalk usually less than 6 mm thick; found on wood above the ground . 7
6. Not as above; if growing on wood above ground then fruiting body larger or compound . . 9
7. Margin of cap often ciliate (fringed with hairs); pores averaging about 1 mm broad, usually
hexagonal; stalk typically central .P. arcularius, p. 563
7. Margin of cap not ciliate; pores usually larger or smaller than above; stalk central to lateral 8
8. Pores large (up to 3 mm), diamond-shaped or hexagonal, usually arranged in radiating rows; stalk
very short, often lateral .P. mori (see P. arcularius, p. 563)
8. Not as above; pores smaller; stalk central to lateral; stalk short or long .
.P. brumalis and Microporellus spp. (see P. arcularius, p. 563)
9. Pores minute (3-7 per mm); pore surface and/ or margin of cap staining gray to dark brown or
black when bruised or dried; fruiting bodies often quite large; found at bases of hardwoods in
eastern North America .Meripilus giganteus (see Bondarzewia montana, p. 565)
9. Not with above features . 10
10. Fruiting body compound, i.e., with many small (mostly 2-7 cm broad) overlapping caps or
segments arising from a common, often branched base; fairly common in eastern North
America, especially with hardwoods, but rare in the West; spores smooth. 11
10. N ot as above; fruiting body not compound, or if so then mature caps larger (5 cm broad or more)
or fewer or habitat different; spores smooth or ornamented . 12
11. Fruiting body white or tinged cream; fertile surface smooth, toothed, or with pores; growing in
rosettes on hardwood stumps or roots in tropics and Gulf Coast Hyduopolyporuspalmatus
11. Not with above features . 28
12. Pores minute (2-4 per mm); pore surface sulfur-yellow, or if not then cap salmon-colored or
pinkish or found in Southeast and cap buff to pinkish- or red-brown (see Laetiporus, p. 572)
12. Pores larger, or if small then not as above . 22
13. Cap blue to blue-gray to bluish-green (but often salmon-,ochre-, or rusty-stained in age) . 14
13. Cap not bluish when fresh (but may be greenish) . 15
14. Pore surface white when fresh; found in western North America .A.flettii, p. 558
14. Pore surface bluish when fresh; found in eastern states A. caeruleoporus (see A.flettii, p. 558)
15. Cap yellowish to yellow-brown to greenish or brownish-stained, with matted hairs or prominent
scales in age; pores usually staining greenish, averaging0.5-2 per mm A. ellisii& others, p. 559
15. Not as above; differently colored and/or cap not hairy-scaly .. 16
16. Spore print olive-brown to brown; flesh and pores not white; pores yellow to greenish, averaging
3 or less per mm; associated with hardwoods (especially ash and alder) (see Boletaceae, p. 488)
16. Not as above; spore print whitish and/ or flesh white before exposure or differently colored 17
556 POLYPORACEAE & ALLIES

17. Found under eastern hardwoods (occasionally conifers); pores 1-3 per mm, yellow to greenish or
if white then cap yellowish, ochre, or greenish; spores smooth A.cristatus{see A. ellisii, p. 559)
17. Not as above; differently colored or pores smaller or spores not smooth . 18
18. Fruiting body with many yellowish petal-shaped caps arising from a common base; found under
western conifers; apparently rare. A. dispansus
18. Not as above . 19
19. Cap and stalk grayish to purple-gray, black, vinaceous, or with olive, pinkish, or brownish tones
(sometimes whitish when young), but not typically yellow-stained; cap often with a somewhat
streaked appearance; pores small(M per mm), white when fresh butoftendiscoloringgrayish,
vinaceous, etc., in age and usually staining olive to black in KOH; spores angular-warty under
the microscope, not amyloid .Boletopsis subsquamosa group, below
19. Not as above; pores larger and/ or cap and stalk differently colored and/ or developing yellowish
tones in age or after handling; spores smooth and/or amyloid . 20
20. Pores small (2-5 per mm); cap white to yellow, pinkish-tan, or pinkish-cinnamon (occasionally
darker), not hairy or scaly (but may be cracked); stalk not blackish at base; tuber absent 21
20. Not as above; pores larger or stalk darker or tuber present or cap hairy or scaly, etc.22
21. Cap or entire fruiting body often developing reddish or orangish stains in age or when dried;
taste usually bitter .A. confluens& others (see A. ovinus, p. 557)
21. Not as above; taste mild or bitter.A. ovinus &. others, p. 557
22. Cap and stalk brown to grayish-brown and covered with small hairs; taste very bitter (chew on a
small piece of cap); odor sometimes (but not always!) like iodine; stalk with neither a tuber nor
a gnarly rooting base; spores smooth; found in western North America . . . . P. hirtus, p. 560
22. Not as above .23
23. Fruiting body arising from a large, swollen, brown to black underground “tuber”(sclerotium);
cap often with fibrils or scales; spores smooth .P. tuberaster, p. 563
23. Not as above (but fruiting body may have a rooting base); spores smooth or warted .24
24. Found at or near bases of trees or stumps; fruiting bodies often clustered or compound (with one
to many caps), usually arising from a gnarly, rooting base; caps pallid to tan, ochre, or brown,
but usually lacking darker scales; spores with amyloid warts or spines .29
24. Not as above; fruiting body typically not compound (but may be clustered); spores smooth 25
25. Stalk with a black rooting base; pores usually 2-3 per mm P. radicatus(see P. tuberaster, p.563)
25. Not with above features . 26
26. Cap dark brown to brown, reddish-brown, or pinkish-brown; terrestrial A. pescaprae, p. 560
26. Found on wood, or if terrestrial then cap usually paler, oranger, or yellower than above . 27
27. Cap with brown scales, often large; pores often more than 1 mm in widest dimension; stalk often
blackish or dark brown at base; found on wood, widespread but especially common in eastern
North America .P. squamosus(see P. decurrens, p. 561)
27. Not as above; usually medium-sized; if found on wood then pores averaging 1 mm wide or
smaller; cap smooth or scaly; stalk base not normally blackish P. decurrens & others, p. 561
28. Caps usually spoon-or fan-shaped; stalks off-center to lateral .Grifolafrondosa, p.564
28. Caps more or less circular; stalks central .Grifola umbellata (see G.frondosa, p. 564)
29. Each fruiting body with one to many caps; favoring conifers . . Bondarzewia montana, p. 565
29. Favoring hardwoods, with one to several caps Bondarzewia berkeleyi(see B. montana, p. 565)

Boletopsis subsquamosa group (Kurokawa)

CAP 4-15 (20) cm broad, broadly convex to plane or slightly depressed; circular in outline
to somewhat irregular; surface dry, smooth to fibrillose or breaking up into small scales,
especially at center; color variable; dingy whitish to gray, purple-gray, bluish-gray,
vinaceous-tinged, or with olive, pinkish, brownish, or even black tones (usually darker
in age), often with a somewhat streaked appearance; margin incurved at first, often wavy or
lobed. Flesh thick, firm, white or tinged variously with cap colors; taste mild to bitter.
PORES small or minute (1-5 per mm), usually white when fresh and young but soon
discolored grayish, vinaceous-buff, brownish, etc.; tubes 2A mm long, usually at least
Boletopsis sub squamosa is variable in color, but usually quite dark (gray, black, purplish, etc.) in age.

somewhat decurrent. ST ALK (2)4-13 cm long, 1 -4 cm thick, central or off-center, equal or

with a narrowed base, solid, smooth; whitish or colored like the cap or pores. SPORE
PRINT white to pale brown; spores 4-7 * 3.5-5 microns, elliptical to nearly round, but
distinctly angular-warty.
HABITAT: Solitary to scattered or gregarious on ground under both hardwoods and
conifers; widely distributed. In our area it is fairly common in the fall and winter under
tanoak and madrone; to the north it often fruits in coastal sand dunes under pine.
EDIBILITY: Edible, but often bitter. It is esteemed by the Japanese, who usually soak it
in brine to remove the bitterness.
COMMENTS: Like Albatrellus ovinus, this polypore is sometimes mistaken for a bolete
because of its frequently central, well-developed stalk and terrestrial growth habit.
However, the tube layer cannot easily be peeled from the cap and the overall shape is quite
different. The color of the fruiting body (especially the cap) is extremely variable, leading
to the naming of several different “species,” including B. leucomelas, with a bluish-black
to black cap, and B. griseus, with a grayish cap. These color forms appear to intergrade,
however, and the common one in our area can be bluish-gray to purplish, pinkish-purple,
or blackish, depending on season, temperature, and exposure to sunlight. The warty-
angular spores are the critical microscopic feature. Albatrellus avellaneus (often listed
as a variety of A. ovinus) can be similar in color but stains yellow and has smooth spores.

A Ibatrellus ovinus (Sheep Polypore)

CAP 4-15 (20) cm broad, circular in outline or irregular; convex becoming plane or slightly
depressed; surface dry, unpolished, smooth or breaking into scales in age; usually whitish
at first, but often yellow, ochre, buff, pinkish, or tan (especially at center) in age, or in some
forms purplish to purple-gray throughout; margin often wavy. Flesh thick, white or yellow,
firm and rather tough. PORES minute (2-4 per mm), white but often becoming yellow
or yellowish in age or when bruised; tubes shallow (1-2 mm long), usually decurrent.
STALK 3-10 cm long, 1-3 (4) cm thick, central or slightly off-center, equal or enlarged
below with a narrowed base; solid, firm, whitish or tinged cap color, but sometimes staining
pinkish. SPORE PRINT white; spores 3-4.5 * 2.5-3.5 microns, elliptical to nearly round,
smooth, not amyloid.
HABITAT: Solitary to scattered or gregarious, sometimes in fused masses, on ground
in mixed woods and under conifers; widely distributed. It is very abundant in the Rocky
Mountains in the late summer under spruce and I have also seen impressive fruitings in
Maine. In our area it is locally common under manzanita in the fall and winter.
EDIBILITY: Edible when cooked well, and according to European sources, fairly good.
Large quantities can have laxative effects, however, and material I tested was rather slimy
(“okraceous”) when cooked.
557
Two farflung terrestrial polypores. Left: Albatrellus ovinus, with a white to tan or yellowish cap.
Right: Albatrelluspescaprae(see p. 560), with a scaly brown cap and well-developed stalk.

COMMENTS: This widespread Albatrellus is variable in color (the purplish-capped

form is often called A. avellaneus), but can usually be recognized by the yellow tints that
develop in age or after handling. The pores are sometimes so tiny that the pore surface may
appear completely smooth. Because of its terrestrial habit this polypore is sometimes
mistaken for a bolete, but is much tougher and does not have detachable tubes. Other
species: A. confluens is a very similar, widespread species which becomes orange to
pinkish-cinnamon in age or upon drying and has slightly larger, weakly amyloid spores
and a bitter taste; A. similis, reported from Arizona, is also similar, but has distinctly amy¬
loid spores; A. peckianus is an eastern beech-lover with a yellow to cinnamon-buff cap.

Albatrellusflettii (Blue-Capped Polypore)

CAP 5-20 or more cm broad, convex becoming plane or centrally depressed; surface dry,
blue to blue-gray or sometimes blue-green, but developing ochraceous, salmon, or rusty
stains in age; margin at first incurved, often lobed or wavy. Flesh thick, firm and rather
tough, white. PORES fairly small (1-4 per mm), white, but often becoming torn and
developing salmon or ochraceous stains in age or upon drying; tubes 1-7 mm long,

Albatrellus flettii. The blue-gray to blue-green cap, white pore surface (the pores are too tiny to see
here), and well-developed stalk distinguish this beautiful species. (Ralph Buchsbaum)
ALBATRELLUS 559

decurrent. STALK 5-15cmlong,(l) 1.5-4cmthick,centraloroff-center,equalorthickerat

either end, white to pale bluish-gray, or aging dingy ochraceous to reddish; solid, firm.
SPORE PRINT white; spores 3.5-4 x 2.5-3 microns, elliptical to nearly round, smooth,
weakly amyloid.
HABITAT: Scattered to gregarious or in fused clusters on ground in mixed woods and
under conifers; known only from western North America. I have found it under knobcone
pine in the fall and winter, but it seems to be quite rare in our area.
EDIBILITY: Edible. It has been sold commercially in northern California, but I haven’t
tried it. If it’s anything like A. ovinus, then it isn’t worth collecting.
COMMENTS: The unusual blue to blue-gray cap color contrasts nicely with the white
pores to set this beautiful terrestrial polypore apart. Dingy ochraceous to rusty-ochraceous
stains develop in age, but there are almost always vestiges of blue-green or blue somewhere
on the cap. Other species: A. caeruleoporus of eastern N orth America is similar, but entirely
indigo-blue to blue-gray (including the pores).

A Ibatrellus ellisii (Greening Goat’s Foot)

CAP 8-25 cm or more broad, convex becoming plane, wavy, or depressed; surface dry,
at first hairy or plushlike but the hairs often matted in age or grouping to form coarse
scales; greenish to sulfur-yellow or yellow-brown (often a mixture of colors), sometimes
with darker brown shades or stains in age; margin often wavy and at first inrolled. Flesh
thick, firm, white, sometimes bruising greenish slowly. PORES 0.5-2 mm in diameter,
white, usually staining greenish or yellow-green when bruised or becoming yellowish to
dingy greenish in old age; tubes 2-6 mm long, often decurrent. STALK 3-12 cm long or
more, 2-6 cm thick, usually off-center or lateral, solid, tough, equal or thicker at either
end; usually colored more or less like cap. SPORE PRINT white; spores 8-9 x 5-7 microns,
elliptical, smooth.
HABITAT: Solitary, scattered, gregarious, or in fused clusters on ground in woods; widely
distributed. It favors conifers and is especially common in the mountains of western N orth
America. I have seen huge fruitings in the late summer and fall in the Cascades and Sierra
Nevada, but in our area it is supplanted by A. pescaprae.
EDIBILITY: Edible, with a mild flavor and pleasantly chewy texture. Care must be taken
to simmer it slowly and thoroughly or it will toughen.

Albatrellus ellisii. This terrestrial species is especially common under conifers in the Pacific North¬
west and Sierra Nevada. Note hairy-scaly cap. The pore surface usually stains greenish when bruised.
560 POLYPORACEAE & ALLIES

COMMENTS: Also known as Scutiger ellisii, this impressive conifer-lover is easily told by
its yellow-green to yellow-brown color, fairly large size, growth on the ground, and ten¬
dency to stain greenish (especially the pores). Its close relative, A. pescaprae, is browner or
redder in color and more southern in distribution. A. sylvestris is a somewhat similar
species with a smoky-olive or darker pore surface and roughened spores. A. cristatus of
eastern North America is similar in color but has a less hairy cap and favors hardwoods.

A Ibatrelluspescaprae (Goat’s Foot)

CAP (2.5) 5-20 cm broad, convex to plane or centrally depressed; surface dry, covered with
fine fibrils which form small, dense scales or sometimes a plush(especially toward center);
dark brown to brown, reddish-brown, or pinkish-brown; margin often deeply indented.
Flesh thick, fairly tough, white, bruising pinkish slowly; taste mild. PORES large, angular,
1 -2 mm or more in diameter (or length); white to yellowish or with greenish stains, some¬
times becoming pinkish in age; tubes 2-5 mm long, often decurrent. STALK 2.5-8 cm
long, 1-3 (4) cm thick, solid, usually enlarged below (sometimes several arising from a
common base); occasionally central, but usually off-center to lateral; white to yellowish or
in age brownish. SPORE PRINT whitish; spores 8-11 x 5-6 microns, elliptical, smooth.
HABITAT: Solitary or in groups or clumps on ground in woods; widely distributed, but
more southern than A. ellisii. In our area it is not uncommon in mixed woods and under
tanoak and madrone, especially in the fall and winter.
EDIBILITY: Edible, at least according to European sources; I haven’t tried it. Young
caps are said to be quite tasty (though chewy) if cooked thoroughly.
COMMENTS: Also known as Scutiger pescaprae, this terrestrial polypore can be told
by its reddish-brown to brown, densely scaly cap plus its large pores, well-developed stem
(see photo at top of p. 558), and mild taste. Bondarzewia species are somewhat similar,
but have spiny spores and less scaly caps and are not normally yellowish- or greenish-
stained. Polyporus hirtus is also somewhat similar, but has a bitter taste.

Polyporus hirtus (Bitter Polypore; Iodine Polypore)

CAP 5-18 cm broad, convex to plane or somewhat irregular; surface dry, covered with
minute short, stiff, erect or matted hairs; uniformly brown to dark brown or grayish-
brown. Flesh thick, white, firm and rather tough; taste very bitter (but often latently).
PORES white to creamy, drying yellowish, 1-2 per mm; tubes 2-6 mm long, adnate to
decurrent. STALK 2-10 cm long, 0.7-3 cm thick, usually lateral or off-center; equal or
tapering downward, hairy and colored like cap; solid, firm, rather tough. SPORE PRINT
white; spores 12-17 * 4.5-6 microns, spindle-shaped or cylindrical (“boletoid”), smooth.
HABITAT: Solitary or in groups on ground, around old stumps and trees (especially
conifers), sometimes also on wood (often buried); northern North America. It is not un¬
common in our area in the fall and winter, particularly in mixed woods of tanoak and
redwood. It is also fairly common in the Pacific Northwest.
EDIBILITY: Inedible. If you boil it for several days and change the water frequently, you
might remove the bitter taste, but you would remove everything else as well!
COMMENTS: Also known as A Ibatrellus hirtus and Scutiger hirtus, this species is the
most common terrestrial polypore in our area, but is by no means numerous. The evenly
colored brown hairy cap plus the white pores, bitter taste, and presence of a stalk are good
fieldmarks (see photos at top of next page). According to mycologist Daniel Stuntz, speci¬
mens in the Pacific Northwest emit an iodine odor soon after being picked, but I have not
detected such an odor in local material. Bondarzewia montana can be similar when it
doesn’t grow in clusters, but has warted amyloid spores and is not as bitter-tasting.
Polyporus hirtus. Left: Top view showing hairy brown cap. Right: Underside, showing white pores.

Polyporus decurrens
CAP 4-14 cm broad, convex to plane or slightly depressed; surface dry, yellowish to yellow-
brown to ochre or dull orange (rarely reddish-brown), with darker (usually brown) erect
fibrillose scales which become flattened in age. Flesh thick, white, tough. PORES fairly
large (0.5-2 mm in diameter), angular or becoming torn or toothlike in age, white, but
sometimes discoloring when dried; tubes 2-6 mm long, usually decurrent. STALK 2-10
cm long, (0.5) 1 -3 cm thick, equal or thicker at either end, solid, tough; white or somewhat
brownish, usually reticulate above from the decurrent tube walls. SPORE PRINT white;
spores 10-18 * 4-6 microns, cylindrical, smooth.
HABITAT: Solitary or in small groups or tufts on the ground (usually originating from
buried wood) or sometimes on dead wood, fall through spring; known from California,
occasional. It favors hardwoods, but I have found it in a variety of habitats. It is the most
common terrestrial polypore of southern California.
EDIBILITY: Unknown. The closely related P. squamosus (see comments) is edible
when thoroughly cooked, but is thoroughly mediocre.
COMMENTS: The dull orange to yellowish-brown to ochre scaly cap with fairly large
decurrent pores distinguish this species from Albatrelluspescaprae and others. It is also
known as P. mcmurphyi. The “Dryad Saddle,” P. squamosus, is a better-known, closely

Polyporus decurrens has large pores, a scaly cap, and may or may not be terrestrial. P. squamosus
(not illustrated) is a similar but larger wood-inhabiting species.
562 POLYPORACEAE & ALLIES

related species. It is larger (cap 6-30 (60) cm broad and stalk up to 5 cm thick) and paler
with dense, flattened brown to dark brown scales (but may become darker brown overall
in age) and very large pores (1 -10 mm each in largest dimension). It usually grows solitary
or clustered on hardwoods (often living) rather than on the ground and its stem, when
well-developed, usually has a black base. In N orth America it is most common east of the
Rocky Mountains, but occurs occasionally in Washington and California. (I’ve seen what
might be a photograph of this species from Fresno.) Another related species,P. fagicola
(-P. lentus) of eastern North America, occurs on hardwoods but is smaller and less scaly.

Polyporus badius (Black-Leg)

CAP 4-20 cm broad, convex becoming depressed or umbilicate; surface tan to chestnut-
brown to dark reddish-brown or often becoming blackish in age; margin usually paler,
often lobed. Flesh tough when fresh, rigid when dry, very thin, white becoming brownish.
PORES minute, 4-7 per mm, white when fresh, brownish in age; tubes shallow, adnate to
somewhat decurrent. STALK 1-6 cm long, 0.3-1.5 cm thick, central or off-center or lateral,
more or less equal; pallid above and black below or more frequently black throughout,
tough. SPORE PRINT white; spores 5-9 * 3-4 microns, cylindrical to elliptical, smooth.
HABITAT: Solitary or in groups on rotting hardwoods (especially aspen) or occasionally
conifers; widely distributed, and occasional in our area in the fall, winter, and spring.
EDIBILITY: Too tough to eat.
COMMENTS: Also known as P. picipes, this species is close to P. elegans but has a darker
and often larger cap (see photo on p. 42). Also, the entire stalk is usually black and it often
grows on logs and stumps whereas P. elegans is more common on branches and sticks.
Both species are retained in the genus Polyporus in its residual (narrowest) sense. For
similar species, see comments under P. elegans.

Polyporus elegans (Elegant Polypore; Black-Foot)

CAP 1.5 -7 cm broad, round to kidney-shaped in outline; convex, soon becoming de¬
pressed, vase-shaped, or umbilicate; surface smooth or finely striate, pale tan to tan or
ochre, often weathering to white; margin often wavy or lobed. Flesh tough when fresh, rigid
when dry, thin, white to pale cinnamon. PORES minute, 4-6 per mm, white when fresh but
often becoming grayish or brownish in age; tubes shallow, usually decurrent. ST ALK 0.5 -5
cm long, 2-6 (10) mm thick, central or off-center or lateral, more or less equal or swollen at
base; pallid or tan above, soon becoming black below; tough. SPORE PRINT white;
spores 6.5-10 * 2.5-4 microns, cylindrical, smooth.
HABITAT: Solitary or several together on decaying hardwoods sticks, branches, and
debris (only rarely on conifers); widely distributed. It is fairly common in our area in the

Polyporus elegans, a common wood-inhabitor. Note pale cap and black lower half of stalk. Compare
it to the photo of P. badius on p. 42.
Polyporus arcularius grows on hardwoods. Note ciliate (hairy) cap margin and stalk that is not black.

winter and spring, especially on willow, alder, poplar (cottonwood), and oak.
EDIBILITY: Too tough to eat, but dries nicely for decorative purposes.
COMMENTS: This dainty polypore is easily recognized by its tan to whitish cap, small
size, and black “foot.” P. varius is sometimes listed as a synonym, but is also applied to a
form with a radially streaked cap that is intermediate in color between P. elegans and P.
badius. T he latter species is usually larger than P. elegans, and has a darker cap and entirely
black stem. Another species, P. melanopus, grows on the ground or from buried wood; it
has a velvety stem and velvety-scurfy cap, at least when young.

Polyporus arcularius (Fringed Polypore)

CAP 1-8 cm broad, round in outline, convex becoming depressed, vase-shaped, or
umbilicate; surface dry, golden-brown to dark brown, usually minutely scaly; margin
often ciliate (fringed with fine hairs). Flesh thin, white, tough. PORES large(l -2 per mm),
angular or hexagonal, white or yellowish; tubes shallow, sometimes decurrent. STALK
2-6 cm long, 2-4 mm thick, usually central or slightly off-center, more or less equal, dark
brown to yellowish-brown, smooth or minutely scaly, the base sometimes hairy. SPORE
PRINT white; spores 7-11 * 2-3 microns, cylindrical, smooth.
HABITAT: Solitary or in small groups on dead hardwoods; widespread, but rare in our
area. I’ve seen it in the Southwest and Pacific Northwest in the summer, and in eastern
North America in the spring. Several closely related species are abundant in the tropics.

EDIBILITY: Much too small and tough to be edible.

COMMENTS: The small size and fairly large pores plus the typically central stem and
frequently ciliate cap margin characterize this attractive polypore. P. mori(also called
Favolus alveolaris), similar but without hairs on cap, and with a stubby stalk and large,
diamond-shaped or hexagonal pores, occurs east of the Rockies. P. brumalis has much
smaller pores and its cap is some shade of brown. It is common on birch and other
hardwoods in eastern North America, rare in the West. Microporellus dealbatus has a
concentrically zoned cap and central stalk; M. obovatus has a whitish, unzoned cap and
central to lateral stalk. Both are common on dead hardwoods in the southeastern United
States. ,None of these have the black or half-black stem of P. elegans and P. badius.

Polyporus tuberaster (Stone Fungus) Color Plate 151

CAP 4-15 cm broad, convex to plane with a depressed or umbilicate center or becoming
funnel-shaped; surface dry, tan to brown or ochre, darker with age and with scattered
darker brown fibrils or fibrillose scales which are often radially arranged; margin often
indented or lobed. Flesh thin, pallid, rigid when dry. PORES 1-3 per mm, white to pale
tan; tubes 1-3 mm long, usually decurrent. STALK 2.5-10 (20) cm long, 1-2.5 (4) cm thick,
more or less equal, central or off-center, brown or colored like cap; solid, tough, arising
from a large brown to black underground sclerotium (“tuber”) which is rather rubbery
when fresh but rock-hard when dry; exterior of “tuber” rough and irregular; interior often
marbled with blackish-brown and paler areas and usually full of dirt and debris. SPORE
PRINT white; spores 10-16 * 3.5-6 microns, cylindrical, smooth.

563
564 POLYPORACEAE & ALLIES

HABITAT: Solitary or in twos or threes on ground in woods; widely distributed, but not
common. I have found it several times in our area in mixed woods and under oak and
madrone. It fruits in the fall, winter, and spring.
EDIBILITY: Edible, but tough unless young, fresh, and thoroughly cooked. Native
Americans ate an underground sclerotium which they called “tuckahoe.” It was once
thought to be this species (as reported in the 1 st edition of Mushrooms Demystified), but is
now believed to be the sclerotium of another polypore (see Poria cocos, p. 604). The
“tubers” of P. tuberaster are inedible because they are full of dirt. However, they are sold
in southern Italy under the name pietra fungaia (“Stone Fungus”). Buyers plant the
“tubers” in flower pots, water them regularly, and then eat the fruiting bodies that result.

COMMENTS: The large underground “tuber” immediately identifies this polypore.

The “tuber” might be mistaken for a piece of buried wood or a parasitized potato (it’s
usually the same size or larger), but the texture when fresh is distinctive. It is thought
to be a resting stage of the fungus, or a secret storage compartment for nutrients.
Bondarzewia berke/eyi may have a thick, rooting stem or“tuber,” but is much larger. Other
species: P. radicatus of eastern North America is somewhat similar but larger, and has a
long, black, narrowed rooting base instead of a sclerotium; P. mylittae of Australia has a
yellower cap and produces edible sclerotia weighing up to 40 lbs. each! The sclerotia are
known as “Blackfellow’s Bread,” a reference to their use as food by the aboriginees.

Grifola frondosa (Hen of the Woods; Sheep’s Head)

FRUITING BODY compound, 15-60 cm broad or more, consisting of a mass of numerous
small, overlapping caps arising from a common, fleshy, repeatedly branched base. CAPS
2-7 (10) cm broad, spoon-shaped, tongue-shaped, or fan-shaped and flattened; surface dry,
smooth or rough to fibrillose, gray to brown or grayish-brown; margin often wavy. Flesh
white and firm, rather tough; taste mild when young. PORES 1-3 per mm, white or
yellowish; tubes shallow (2-3 mm long), decurrent. STALKS (branches) smooth, fleshy
but tough, white or pale grayish, off-center or more often lateral (attached to sides of
caps). SPORE PRINT white; spores 5-7 * 3.5-5 microns, broadly elliptical, smooth.
HABITAT: At or near the bases of trees and stumps (usually oak), fruiting year
after year in the same spots; widely distributed, but common only in eastern North

Left: The “Hen ol the Woods,” Grifola frondosa, is a prized edible mushroom in eastern North
America. The olten massive fruiting body is composed ol numerous petal- or lan-shaped caps arising
Irom a common base. Right: Grifola umbellata mimics G. frondosa, but each of its caps has a more
or less central (rather than lateral) stalk, as shown here. It is also delicious.
GRIFOLA 565

America. It has been reported from Idaho, but is rare in the West and apparently absent in
our area. It causes a delignifying butt rot of both the heartwood and sapwood of its host.
EDIBILITY: Edible and choice—along with the sulfur shelf and beefsteak fungus, the best
of the polypores for the table. Long, slow cooking is recommended and only the young,
tender caps are worth eating. “Allergies” have been reported. It is excellent pickled.
COMMENTS: The numerous overlapping, more or less spoon-shaped caps are remi¬
niscent of a fluffed-up hen, making this one of the safest and most easily recognized of all
edible mushrooms. Clusters weighing 100 pounds have been recorded, but 5-10 pound
specimens are the norm. There are usually more caps per cluster than in Bondarzewia, and
the stems are attached to the sides of the caps rather than being central. Polyporus or
Polypilus frondosus are synonyms. G. umbellata(-P. umbellatus), also edible, is a similar
species with whitish to gray to smoky-brown, circular caps with central stems. Its fruiting
bodies arise from sclerotia (“tubers”) that have been used by the Chinese as an immune
system stimulant. It occurs across the northern half of the continent, but is not common.

Bondarzewia montana
FRUITING BODY simple (with a cap and stalk) or compound (with numerous caps and
stalks arising from a common base), but often fingerlike when first emerging or taking
the form of a lumpy, misshapen, or rosette-like mass (when compound); often massive
when mature (up to 1 m (3 ft.) broad if compound); usually arising from a rooting base.
CAP(s) 5 -25 cm broad when mature, convex to irregular or depressed; surface dry, finely
velvety to fibrillose, or becoming smooth; tan or ochre to brown or dark brown, sometimes
paler when immature. Flesh fairly thick, white, firm but brittle when fresh; taste mild or
sometimes bitter in age. PORES fairly large (0.5-2 mm broad), usually angular or irregular
in shape or breaking up to form “teeth”; pore surface often lumpy or nodulose in compound
fruiting bodies, white to creamy or buff, dingier when dried or in age; tubes 1-6 mm long,
usually decurrent. STALK(s) central to lateral, continuous with individual caps and simi¬
lar in color and texture, usually arising from a tapered, underground, gnarly rooting base
that is 4-12 cm long and 2-5 cm thick. SPORE PRINT white; spores 5-7 microns, round,
with conspicuous amyloid warts.
HABITAT: Solitary or in groups under conifers, usually near stumps or trunks (pre¬
sumably arising from roots or buried wood); locally common in the late summer and fall
in western North America, particularly at higher elevations. I have found it in quantity
near Lake Tahoe under fir and white pine, but have not seen it on the coast. The very similar
B. berkeleyi (see comments) favors living hardwoods, producing a serious delignifying
butt rot of the heartwood (“string and ray rot”) that completely hollows out its host.
EDIBILITY: Tempting because of its size, but rather tough and sometimes bitter to boot.
It is probably harmless, but certainly not the equal of Grifola frondosa.
COMMENTS: The warted amyloid spores, often compound fruiting body, and gnarled
rooting base are characteristic of the genus Bondarzewia. The spores are unique among
the polypores, leading some investigators to suggest a relationship to the agaric genera,
Russula and Lactarius. When B. montana has a massive compound fruiting body (see
photo on p. 548) it is reminiscent of the hen of the woods, Grifola frondosa, an eastern
hardwood-loving species with smooth spores. Fruiting bodies with one to several caps, on
the other hand, are more likely to be confused with Polyporus hirtus, a bitter-tasting,
smooth-spored polypore. Its only close relative, B. berkeleyi, is quite similar but usually
has fewer (one to five), broader, paler (whitish to grayish or yellowish-tan) caps and slightly
larger spores, plus it favors hardwoods (especially oak and maple). It seems to intergrade
with B. montana, and is very widely distributed. Another sometimes gigantic polypore,
Meripilus giganteus, occurs at the bases of hardwoods in eastern North America. It stains,
566 POLYPORACEAE & ALLIES

ages, or dries gray to dark brown or black on the pore surfaceand/ or marginand has much
smaller pores (3-7 per mm). Like Bondarzewia, it often has a compound fruiting body, but
has smooth, non-amyloid spores. The individual caps in all of these species are usually
larger than those of Grifola.

Heteroporus biennis
FRUITING BODY arising from a poorly developed stalk or fleshy base, with one cap or
several fused together in an overlapping rosette, or very distorted with most of the
surface covered with pores. CAP 3-9 (20) cm broad when well-developed, plane to de¬
pressed; surface dry, woolly-hairy or felty, white to tan or aging pinkish to reddish. Flesh
tough, duplex, white or pinkish. PORES 1-3 per mm, angular, irregular, or mazelike, or
becoming toothlike; whitish, but often discoloring or bruising reddish, dingier in age;
tubes 2-6 mm long, usually decurrent. STALK central to off-center, poorly developed or
even absent, continuous with and colored like the cap; hairy. SPORE PRINT white; spores
4-8 x 3-5 microns, elliptical, smooth; thick-walled, nearly round spores (chlamydo-
spores) often present also.
HABITAT: Solitary or in groups on ground around hardwood stumps and trees (pre¬
sumably growing on the roots), occasionally under conifers; widely distributed, but not
common. I have found it in our area under oak and pine in the fall and winter.
EDIBILITY: Unknown; too tough to be worthwhile.
COMMENTS: Misshapen fruiting bodies are usually found with pores covering much or
most of the mushroom—the best fieldmark of this otherwise unimposing, profoundly
forgettable, pitiful excuse for a polypore. The growth habit is sometimes reminiscent of
Phaeolus schxveinitzii. Abortiporus biennis and Polyporus biennis are synonyms.

PHAEOLUS, INONOTUS, COLTRICIA, & Allies

Fruiting body usually annual, small to large, often soft and spongy when fresh but tough or corky-
woody in age; growing shelflike to bracketlike to practically resupinate on wood or growing on
ground and possessing a stalk and one to several caps. CAP usually hairy, fuzzy, or velvety. Flesh
orange to yellow-brown, rusty-brown, or brown. PORES orange to greenish-yellow, rusty-brown,
brown, or grayish (but not white) at maturity; small or large. STALK present or absent. SPORE
PRINT whitish to yellow or brown when obtainable. Spores usually elliptical, smooth. Setae
(large brown sterile cells) often present among basidia. Cap tissue darkening (turning red to black)
in potassium hydroxide (KOH).

THESE orange to brown or rusty-yellow polypores are somewhat intermediate in aspect

between the woody perennial polypores (conks) and the smaller and thinner, annual
bracket fungi and stalked polypores. Phaeolus and Inonotus most closely resemble Phel-
linus (a genus of conks), but are usually softer and fleshier when fresh, and have annual
rather than perennial fruiting bodies. Coltricia, on the other hand, has a central stalk and
is reminiscent of a Polyporus.
All three genera have tissue that darkens or blackens dramatically when touched with a
2% aqueous solution of potassium hydroxide (KOH). This feature is also shared by
Phellinus, leading some mycologists to sequester the four genera in a family of their own.
Phaeolus and Inonotus are wood-inhabiting but appear terrestrial when growing from
roots or buried wood, while Coltricia is nearly always terrestrial. Inonotus species typically
produce white rots and Phaeolus species cause brown ones (P. schxveinitzii is a serious
pest of standing timber). They are not edible because of their tough texture, and P.
schxveinitzii may actually be poisonous. Five species are described here.
PHAEOLUS, INONOTUS, COLTRICIA, & ALLIES 567

Key to Phaeolus, Inonotus, Coltricia, & Allies

l. Fruiting body with a stalk, typically appearing terrestrial (but may be at or near base of tree) 2
1. Fruiting body growing shelflike or bracketlike on logs, stumps, trees, etc.; stalk absent ... 5
2. Underside of cap with concentrically-arranged plates or with very large pores (0.5-2 mm in
diameter); pores whitish to brownish; stalk 2-5 cm long, with only one cap; confined to eastern
North America; not common.Coltricia montagnei(see C. cinnamomea, p. 568)
2. Not as above . 3
3. Stalk well-developed and distinct from cap (only one cap present); flesh corky-tough and thin
(usually less than 2 mm thick), never spongy .4
3. Stalk often present only as a narrowed, often rooting base, with one to several caps; flesh usually
thicker than above and spongy when fresh (but tough in age) . 11
4. Cap shiny or silky, rarely more than 5 cm broad; pores usually not decurrent .
.Coltricia cinnamomea, p. 568
4. Cap often velvety but not usually silky or shiny, sometimes small but up to 11 cm broad ....
.Coltricia perennis (see C. cinnamomea, p. 568)
5. Pores small (2-4 per mm) and bright saffron (orange-yellow) to bright orange or red when fresh;
fruiting body soft or tough; usually found on hardwoods . . (see Trametes & Allies, p. 592)
5. Not with above features; pores differently colored and/or larger .6
6. Flesh and tubes bright orange or soon becoming orange; typically on conifers .7
6. Not as above (but may be rusty-yellow, reddish-brown, etc.). 8
7. Pores large (1 mm or more in diameter); tubes 1 cm or more long; cap rudimentary (usually
present only as a free margin) .Phaeolus alboluteus, p. 571
7. Pores smaller; tubes shorter; cap present .... Phaeolusfibrillosus (see P. alboluteus, p. 571)
8. Typically growing on or under conifers .9
8. Typically growing on hardwoods (but occasionally on conifers) . 12
9. Spore-bearing surface taking the form of gills or very long, meandering mazelike pockets or
if not then fruiting body usually with an anise odor when fresh; flesh corky-tough; pores not
yellowish or greenish when fresh; found on wood . (see Lenzites, Daedalea, & Allies, p. 586)
9. Not as above; found on wood or ground; flesh often spongy when young and fresh . 10
10. Pores minute (3-6 per mm); cap surface soon more or less bald; growing shelflike or bracketlike
on wood .Inonotus dryadeus(see I. tomentosus, p. 569)
10. Pores usually larger (up to 4 per mm); cap surface usually hairy or velvety, at least when young;
growing shelflike or on ground .11
11. Pore surface yellow to greenish-yellow when fresh and young; cap often showing bright (orange
to yellow) zones when young (but fruiting body entirely reddish-brown to dark brown in old
age); stalk (when present) often with more than one cap .Phaeolus schweinitzii, p. 570
11. Pore surface brown to grayish or hoary when fresh; cap yellow-brown to tan when young; stalk
(when present) typically with only one cap; setae present .Inonotus tomentosus, p. 569
12. Fruiting body a roundish to cylindrical, compact mass of numerous small, thin, tough, closely
overlapping caps arising from a solid core or “knot”; pores gray to brown; found on hardwoods
in eastern North America (common name: “Sweet Knot,” because it is sometimes very
fragrant) .Globifomes graveolens
12. Not as above . 13
13. Fruiting body usually resupinate (i.e., lacking a well-defined cap), either dark brown to black
and hard or growing in sheets under oak bark . 14
13. Not as above; fruiting body usually with a cap (upper sterile surface). 15
14. Fruiting body yellow-brown to dull brown; usually growing under the bark of living oaks . . .
.Inonotus undersonii
14. Fruiting body dark brown to black, hard, often cracked and usually irregular or cankerlike in
shape; found mainly on birch.Inonotus obliquus
15. Pores often mazelike or pocketlike (elongated), the walls between them fairly thick; cap thin
(less than 1 cm, the flesh only 1 -4 mm thick), corky or leathery even when fresh .
.(see Lenzites, Daedalea, & Allies, p. 586)
15. Not as above . 16
568 POLYPORACEAE & ALLIES

16. Cap tan to pale brown (or weathering grayish) and distinctly hairy, usually 2 cm thick or less;
pores whitish to gray, grayish-brown, or even blackish; found mainly on dead cottonwood
(Populus) or willow, often in confluent rows . Funalia hispida & relatives
16. Not as above; if growing on cottonwood or willow, then differently colored, etc. 17
17. Fruiting body soft and watery when fresh, entirely tawny-yellow to cinnamon or yellow-brown
and staining red or vinaceous (not black) in KOH; typically found on dead hardwoods in
eastern North America (reported rarely from the West on conifers) . . Hapalopilus nidulans
17. Not as above (but may have some of above features) . 18
18. Fruiting body tough and corky or woody even when fresh; cap surface not hairy or only slightly
so; usually found on dead trees . (ste Phellinus gilvus, p. 582)
18. At least the upper layer of fruiting body usually fleshy (watery or spongy) when young; cap
surface often (but not always) hairy; often on living trees Inonotus hispidus & others, p. 569

Coltricia cinnamomea (Fairy Stool)

FRUITING BODY with a cap and stalk, usually terrestrial. CAP 1-5 cm broad, more or
less circular in outline, centrally depressed or umbilicate; surface dry, bright cinnamon
to reddish-brown, yellow-brown, rusty-brown, or darker, with shiny or silky striations and
narrow or inconspicuous concentric zones; margin often fringed or torn. Flesh very thin
(1 mm thick or less), pliant when fresh; rusty-brown. PORES 2-3 per mm, yellow-brown
to brown or reddish-brown; tubes shallow (0.5-3 mm long), typically not decurrent.
STALK 1 -5 cm long, 1-4 mm thick, usually central and more or less equal, brown to
reddish-brown, hairy or velvety, tough. SPORE PRINT yellowish-brown; spores 6-10
x 4.5-7 microns, elliptical, smooth. Cap tissue staining black in KOH.
HABITAT: Solitary or in small groups on ground or moss in woods, often along well-
beaten paths, roadbanks, and in clearings (rarely on rotten wood); widespread. In our area
this species and C. perennis (see comments) occur year-round, but are not very common.
EDIBILITY: T oo tough to eat, but makes an intriguing addition to seedpod arrangements.
COMMENTS: This beautiful little polypore is easily told by its very thin, silky-shiny,
cinnamon to amber-brown cap, brown pore surface, and reddish-brown to dark brown
(not black!) stem. The rusty-brown flesh distinguishes it from Polyporusand other stalked
polypores. In similar habitats or on charred ground and just as common is the elegant
C. perennis(see photo below). Its cinnamon to yellow-brown or grayish cap is up to 10 cm
broad and usually strongly zoned and finely velvety (rather than silky). Italso tends to have
a thicker stem and brown to grayish-brown pores that are sometimes decurrent. It is widely
distributed, but a third species, C. montagnei, is restricted to eastern North America. It
has large, often mazelike pores or even thick, concentrically arranged plates or “gills.”

Left: Coltricia cinnarpomea has a small silky cap, brownish pores, and thin rusty-brown flesh. Right:
Coltricia perennis is often larger and more strongly zoned. Note tough central stalk in both species.
INONOTUS 569

Inonotus hispidus
FRUITING BODY annual, growing shelflike or bracketlike on wood, at first soft and
spongy but tough in age and rigid when dry. CAP 5-20 (30) cm broad and 2-10 cm thick,
convex or plane; surface at first densely covered with stiff or bristly hairs, the hairs
tending to wear away in age; bright reddish-orange to yellowish-brown to rusty-yellow
when fresh, becoming brown to dark reddish-brown or even blackish in age. Flesh up to
5(10) cm thick, at first watery or spongy (but fibrous), tougher in age; rusty to rusty-yellow
to dark reddish-brown; odor often rather pleasant when fresh. PORES 1-3 (4) per mm,
yellowish to brown or rusty-yellow, sometimes with an olive tinge and sometimes beaded
with droplets, darkening with age or where bruised; tubes 0.5-3 cm long. STALK absent.
SPORE PRINT ochre- to chestnut-brown; spores7.5-11 * 6-9 microns, broadly elliptical
to nearly round, smooth. Flesh darkening (staining red to black) in KOH.
HABITAT: Usually solitary (occasionally several) on living or recently dead hardwoods
or rarely conifers; fruiting mostly in the summer and fall but occurring year-round, widely
distributed. It is a destructive parasite of oak and walnut (causing a white rot of the heart-
wood), but occurs on a wide range of other trees, including mulberry and willow. The
fruiting bodies usually emerge from the wounds of trees, often at a considerable distance
from the ground. I have not seen this species locally but it occurs in southern California.
EDIBILITY: Unknown.
COMMENTS: This species and its close relatives (see below) are reminiscent oil. tomen-
tosus and Phaeolus schweinitzii, but are strictly shelving species with no stalk. They can
also be confused with species of Phellinus, but are soft and spongy when fresh and are not
perennial. Similar species include: I. cuticularis, with a woolly-matted to nearly smooth
(not bristly) cap, found on various hardwoods such as willow, cottonwood, and pepper
trees; /. dryophilus, with a thick granular core between the upper fibrous tissue and the
tube layer, found on hardwoods (especially oak); I. texanus, found on desert legumes
(particularly acacia and mesquite); I. arizonicus, found on sycamore; /. dryadeus (see
comments under I. tomentosus)', and I. radiatus, widespread (but especially common in
eastern North America) on hardwoods such as birch and alder, with a rather thin, firm,
brightly colored (golden to rusty-brown or even yellowish-green) cap that is often zoned
and radially fibrillose plus unpigmented spores. See also I. tomentosus, which some¬
times grows shelflike on conifers.

Inonotus tomentosus
FRUITING BODY usually terrestrial with a cap and a short or rudimentary stalk, but
sometimes lacking a stalk and growing shelflike on wood. CAP 3 -12 (18) cm broad, circular
to fan-shaped in outline, convex to plane to centrally depressed; surface dry, soft and hairy
(tomentose) or velvety, whitish when very young but soon yellowish-brown to tan, dull
ochre, brown, or rusty-brown, sometimes with faint concentric zones. Flesh yellow-brown
to brown, duplex: upper layersoftand spongy when fresh, lower layer rather thin, firm, and
fibrous. PORES round to angular or irregular becoming torn or sometimes toothlike in
age, 2 A per mm, pale buff to grayish or beige or becoming brownish, but often with a hoary
sheen or surface covering; darker brown where bruised; tubes 1.5-7 mm long, usually
decurrent. STALK often rudimentary, when present short(1-5 cm long), 0.5-2 cm thick,
central to off-center or lateral, continuous with cap and more or less same color and texture,
or darker. SPORE PRINT pale yellow to pale brown; spores 4.5-7 * 2.5-4 microns,
elliptical, smooth, hyaline under the microscope. Brown sterile cells (setae) abundant
among basidia, straight and pointed. Cap tissue blackening in KOH (sometimes with a
fleeting red intermediate phase).
Inonotus tomentosus is reminiscent of Phaeolus schweinitzii, but is usually smaller and paler in
color. In addition, the fruiting body is usually simple (with one cap rather than several). Note how
pine needles and sticks are incorporated into the fruiting bodies.

HABITAT: Solitary or in groups on ground under conifers (presumably arising from

roots or buried wood), sometimes also on stumps or bases of trunks; widely distributed.
In our area it is fairly common throughout the mushroom season, especially in coastal
pine forests. It causes a white pocket rot of the roots and butt of a wide variety of conifers
in the pine family, but is especially fond of pine and spruce.
EDIBILITY: Unknown, but too tough to be of value.
COMMENTS: Also known as Onnia tomentosa and Polystictus (or Mucronop or us)
tomentosus, this common polypore can be recognized by its soft hairy or velvety cap and
overall brown to yellow-brown color. It is thicker and spongier than Coltricia; it could be
confused with Phaeolus schweinitzii, but is duller in color, somewhat smaller, and does not
usually have a compound fruiting body. Also, the fresh pore surface is brown to grayish
or hoary rather than yellow or greenish. I. circinatus is a very similar species with hooked
setae; it is also common on or near conifers across North America. /. dryadeus favors
conifers in the West (but oak in the eastern states); it always grows shelflike on wood,
has a bald or nearly bald cap, minute pores, and attains sizes of 30 cm broad or more. For
shelflike (stalkless) species of Inonotus that favor hardwoods and have more highly pig¬
mented (browner) spores, see /. hispidus and the species listed under it.

Phaeolus schweinitzii (Dyer’s Polypore) Color Plate 153

FRUITING BODY usually compound, composed of several caps arising in tiers from a
common base, but sometimes simple (with one cap) and sometimes growing shelflike on
wood. CAP(s) 5-30 or more cm broad, circular to fan-shaped in outline; cushion-shaped
becoming plane or depressed; soft and spongy when fresh and often knobby when actively
growing, tough or corky in age, rigid and brittle when dry; surface covered with a dense felty
or woolly mat of hairs, smoother in age; color variable: orange to ochraceous to yellowish
or greenish-yellow (especially on the margin) when growing, becoming rusty-brown to
dark brown in age (or from the center outward); staining brown to blackish when bruised
and becoming entirely dark brown to blackish in old age; sometimes concentrically zoned
with several of above colors; margin often wavy. Flesh yellowish to rusty-brown or brown,
often appearing zoned. PORES 1-3 per mm or fused together to form larger pores;
mustard-yellow to greenish when fresh, but quickly becoming brown or blackish when
bruised or in age; tubes 2-10 mm long, usually decurrent. STALK (if present) 1-6 cm long
and 1-5 cm thick, usually tapered downward, often rooting, central or off-center, same
color and texture as cap. SPORE PRINT whiteortinged yellow to green; spores5-9 * 3.5-5
microns, elliptical, smooth. Setae absent. Cap surface and tissue staining black in KOH,
often with a fleeting cherry-red intermediate phase.

570
Phaeolus schweinitzii, a common conifer-lover. Note compound fruiting body; these are rather small
specimens.

HABITAT: Solitary or in groups on or around dead and living conifers, usually but not
always appearing terrestrial (originating from the roots); widely distributed and very
common throughout the West. In our area it favors pine and Douglas-fir, the fruiting
bodies usually appearing after the first fall rains but persisting year-round. The mycelium
attacks the roots and heartwood of its host, causing a serious carbonizing decay known as
“red-brown butt rot.” It fractures the wood into cubical blacks as high as fifteen feet up the
trunk, weakening the tree so that it blows over easily. It also occurs on coniferous slash,
in which case it is more apt to be shelflike than terrestrial. In the West it is one of the two or
three most prevalent brown rot fungi. There is one report of it growing on eucalyptus.
EDIBILITY: Possibly poisonous. It contains a stimulant found in the roots of the kava
kava plant, but apparently some toxic substances as well. It is too tough and hairy to be
of food value anyway. However, it is prized by dye-makers for the rich and varied hues
it imparts to yarn.
COMMENTS: One of the most common and conspicuous of the larger polypores, this
cosmopolitan species is apt to confuse the beginner because of the color and texture
changes it undergoes as it ages. However, the yellowish to greenish-yellow pores when
young, yellow to rusty-orange tones in the caps of young specimens, and growth at the base
of or near conifers are diagnostic. (The color plate shows young specimens that are pre¬
dominantly orange rather than greenish-yellow.) It grows gradually, engulfing pine
needles, sticks, plants, and other debris in its way, as shown in the color plate. Old,
weathered fruiting bodies are quite light in weight and entirely dark brown in color.
Inonotus tomentosus is somewhat similar, but has a tan to dull brown cap and brownish or
hoary pore surface. Hydnellum species are superficially similar, but have short, blunt
spines or “teeth” on the underside of the cap instead of pores.

Phaeolus alboluteus (Orange Sponge Polypore)

FRUITING BODY resupinate and spreading or bracketlike, soft and spongy when
fresh, up to 1 meter or more in length. CAP (upper surface) usually present only as a free
margin, soft and spongy; surface bright orange but eventually weathering or fading to
whitish. Flesh thin, orange. PORES large (1-3 mm or more in diameter), angular, the walls
often jagged or splitting to form “teeth”; whitish or yellowish soon becoming orange (or
tubes orange with white edges); tubes long (1-3 cm). STALK absent. SPORE PRINT
white; spores (7) 10-14 * 3-4 microns, cylindrical, smooth. Flesh reddening in KOH.

571
572 POLYPORACEAE & ALLIES

HABITAT: On fallen logs of conifers, usually on the undersides and often developing
in snow; common in the subalpine forests of the West, fruiting mainly in the spring but
persisting into summer, fall, or even winter. It is quite common in the southern Rocky
Mountains, and I have also seen it in the Sierra Nevada. It produces a brown rot(carbon-
izing decay) and is said to occasionally occur on aspen.
EDIBILITY: Unknown.
COMMENTS: This species, which has also been placed in Pycnoporellus and Aurantio-
porellus, is easily told by its large pores and orange spongelike fruiting body that peels off
in one continuous felty layer. P. fibrillosus is a somewhat similar species that is shelflike
(i.e., with a well-defined fiery red to orange or brownish-orange cap). It has creamy to
orange pores and grows on conifer slash or rarely aspen. Pycnoporus, Aurantioporus, and
several Porias are also brightly colored, but are tougher and/ or have much smaller pores.

LAETIPORUS

THIS is a small genus of large, fleshy, shelflike fungi. Only one colorful species—the
well-known and edible sulfur shelf—is described here. A key hardly seems necessary.

Laetiporus sulphureus Color Plates 154,155

(Sulfur Shelf; Chicken of the Woods)
FRUITIN G BODY annual, emerging knoblike or fingerlike, soon becoming shelflike; soft
and fleshy when young, tough in age. CAP 5-50 (70) cm broad and up to 4 cm thick,
fan-shaped to elongated or semi-circular in outline; surface smooth to suedelike, often
uneven or wrinkled; red-orange to bright orange, yellow-orange, sulfur-yellow, or salmon
(the margin usually yellow), fading slowly with age to yellowish, buff, or eventually dull
whitish; margin at first thick and blunt. Flesh thick, soft and watery when very fresh,
becoming tough and eventually crumbly; white to pale yellow or salmon-tinged; when
very fresh often exuding yellow or orange droplets and reminiscent of uncooked chicken;
odor fungal or rather pungent; taste nearly mild or acidic becoming quite sour or un¬
pleasant in age. PORES 2-4 per mm but barely visible when young, bright sulfur-yellow,
but often darkening when bruised and fading slowly in age; tubes shallow (1 -4 mm long).
STALK absent or present only as a narrowed base. SPORE PRINT white; spores
5-7 x 3.5-5 microns, broadly elliptical to nearly round, smooth.
HABITAT: Solitary or more often in overlapping clusters or shelving masses on dead
stumps and logs, sometimes also on living trees or sometimes growing in rosettes from roots
or buried wood; usually appearing on the same stumps year after year; widely distributed
and common. In our area itfruits mainly inthelatesummerandfall, but old fruiting bodies
may persist for months. It grows on a wide range of hardwoods and conifers. In coastal
California it favors eucalyptus (September-October, ushering in the new mushroom
season) and to a lesser extent conifers and oaks (November-December). In the Sierra
Nevada it is common on fir in the summer; in eastern North America it favors oak. It
causes a destructive red-brown carbonizing heart rot that eventually hollows out the tree,
producing cracks in the wood in which thin sheets of white mycelium may appear. It is
said to have caused considerable damage to British sailing vessels.
EDIBILITY: Edible and delectable when thoroughly cooked. However, there have been
several cases of sulfur shelf poisoning on the west coast, so try it cautiously the first time
and never eat it raw. When young the succulent flesh has a mild flavor, tofu-like texture,
and “Candy Corn’Mike color, making it especially attractive and delicious in omelets.
Maturing specimens are tougher and develop a strong sour flavor; their texture when
LAETIPORUS 573

cooked is reminiscent of white chicken meat, so they’re good in sandwiches. If the

specimens you find are mature (but not so old as to be asbestos-like),, you can trim off the
tender, rapidly growing margin (about 5 cm), and perhaps return later for more!
COMMENTS: One ofthe“FoolproofFour”—the brilliant yellow-orange shelving masses
are unmistakable. Actually, nothing is foolproof, but the sulfur shelf is certainly
intelligence-proof, and I trust that no one reading this book is a fool! I always experience
an element of disbelief when I stumble onto a large cluster. It looks like something out of a
Jacques Cousteau movie—you no more expect to find it on an aging eucalyptus stump by
the railroad tracks than you do a freight train at the bottom of the sea! Fresh specimens
can be so soft that it’s difficult to handle them without leaving fingerprints. They are
full of water and weigh far more than their size suggests (I have found clusters weighing
over 50 pounds!). It seems strange that such a large fungus should require so little moisture
to fruit, for in our area it often appears before the arrival of the fall rains. The cap color
ranges from deep orange to yellow or salmon, but the fresh pore surface is always sulfur-
yellow—except in the rare var. semialbinus, which has a salmon-colored cap, white pores,
and a frequently rooting base. Polyporus sulphureus and Grifolasulphureaare synonyms.
Other species: L. persicinus is a southeastern species that usually grows in stalked clumps;
it has whitish to creamy pores and a buff to pinkish-brown or darker cap.

ISCHNODERMA
THIS genus is easily recognized by its roughened, resinous, dark brown to blackish,
shelflike fruiting body. A single species is described here.

Ischnoderma resinosum (Resinous Polypore)

FRUITING BODY annual; shelflike or bracketlike, watery at first and often exuding
droplets, especially near margin of cap, becoming tougher and drier in age. CAP 5-30 cm
broad and 1-3 cm thick, fan- to kidney-shaped or semicircular in outline; surface rough
or velvety (like sandpaper) at first (but often nearly smooth in age), often wrinkled radially
and saturated or incrusted with a resin (the resinous areas often darker and more metallic-
looking and sometimes slightly bluish), often concentrically zoned or ridged in age; color
usually dark brown to blackish, but often overlaid with a thin golden or ochre coating
when very young; margin usually quite thick. Flesh whitish to beige, tan, or brownish,
watery when young becoming tough and corky in age. PORES minute(3-6 per mm), white
or creamy, but often becoming brownish when bruised and ochre-brown to brownish in
age; tubes 1-10 mm long. STALK absent. SPORE PRINT white; spores 4-7 * 1.5-2.5
microns, sausage-shaped to cylindrical, smooth.
HABITAT: Solitary or several together (often overlapping) on dead hardwoods and
conifers, fruiting mostly in the summer and fall; widely distributed but not particularly
common, at least in my experience. In northern California and the Pacific Northwest it
favors larger (first-growth) conifers. It causes a delignifying decay of both the sapwood
and heartwood, and the infected area sometimes has a strong aniselike odor. I have not
seen it ih our area.
EDIBILITY: Said to be edible when young and watery, but soon tough and corky.
COMMENTS: This distinctive shelving polypore is easily identified by its resinous dark
brown to blackish cap that is often radially wrinkled and its tendency to exude droplets
of liquid (especially when young or in wet weather). Some mycologists reserve the name
I. resinosum for the form that grows on hardwoods (particularly elm), while using the
name I. benzoinum for the conifer-loving version with darker (browner) flesh and tubes.
574 POLYPORACEAE & ALLIES

GANODERMA, FOMITOPSIS, PHELLINUS, & Allies

(Conks)
Fruiting body medium-sized to very large; tough, woody, corky, or punky, usually perennial (but
some species annual); often thick; growing on dead or living trees. CAP knoblike to hooflike to
shelflike or bracketlike, sometimes with a hard surface crust; often zoned, ridged, or grooved.
PORES fairly small to minute or barely visible; tubes often stratified (with more than one layer)
as seen in longitudinal section. STALK, usually absent or rudimentary, but sometimes present as a
lateral extension of the cap (see G. lucidum). SPORE PRINT brown to whitish but difficult to
obtain. Spores smooth or minutely prickly.

THESE are the tough, woody, hoof-shaped or shelflike growths you see so often on the
trunks of dead trees or in the wounds and crotches of living ones. They are commonly called
“conks” because of their woody to corky texture and the frequent presence of a hard
surface crust. Conks are larger and thicker than the other tough bracket fungi (e.g.,
Trametes) and usually—but not always—have perennial fruiting bodies (i.e., each year a
new tube layer is added onto the already-existing ones). In many species these tube layers
are distinctly stratified so that the age of the fruiting body can be determined by
counting the number of layers (just like counting the growth rings on a tree). Ages of 50-70
years have been recorded! If the yearly tube layers are not stratified, the age can often be
estimated by the number of growth zones (represented by ridges or furrows) on the cap. A
new zone is “issued” each year as an outer or downward extension of the cap in conjunction
with the new tube layer.
Most conks have at one time or another been classified in the genus Fomes. However
Fomes, like Polyporus, has been obliterated by the “splitters,” so that it now contains
only one common species (F. fomentarius). The principal genera recognized here are
Ganoderma, with a hard, sometimes shiny surface crust, whitish to pallid pore surface
when fresh, and minutely prickly, double-walled spores; Fomitopsis, with whitish to pale-
colored flesh and smooth spores; and Phellinus, with rusty-brown to yellow-brown flesh
that darkens in potassium hydroxide (KOH), and smooth spores. Several small genera
(e.g., Fomes in its residual sense and Heterobasidion) are also described and/ or keyed out.
Conks are responsible for many serious rots. Phellinuspini, Fomitopsis officinalis, and
Heterobasidion annosum are especially destructive to living or standing conifers, Fomi¬
topsis pinicola to dead ones, and the Ganoderma applanatum group to living and dead
hardwoods.
Conks are obviously far too tough or woody to eat (see comments on the edibility of
Fomitopsis pinicola!), but several enjoy other uses (see in particular Ganoderma lucidum,
G. applanatum, and Fomitopsis officinalis). Only some of the more distinctive and/ or
common species in North America are treated here.

Key to Ganoderma, Fomitopsis, Phellinus, & Allies

1. Pore surface rosy or pink when fresh (but often duller or darker in age); flesh usually pinkish also
(when fresh) ... Fomitopsis cajanderiSi others, p. 580
1. Not as above; pore surface not rosy . 2
2. Cap (and stalk if present) with a surface crust that appears varnished, at least when young (unless
covered by spore dust); fruiting body usually annual (with one tube layer); flesh usually light¬
weight or punky;/rcs/r pore surface white, but typically turning brown if rubbed or in age 3
2. Varnished surface crust absent, or if present then not as above; stalk typically absent .6
3. Typically growing on hardwoods . 4
3. Typically growing on conifers . 5
4. Cap ochre to whitish or only partly red; stalk typically present; mainly southern (southeastern
United States), rare northward . Ganoderma curtisii(see G. lucidum, p. 577)
4. Cap usually reddish (or mostly reddish), at least in age; stalk present or absent; widespread in
temperate zone and tropics . Ganoderma lucidum, p. 577
GANODERMA, FOMITOPSIS, PHELLINUS, & ALLIES 575

5. Stalk usually absent; flesh up to 10 cm thick; known only from western North America .
. Ganoderma oregonense (see G. lucidum, p. 577)
5. Stalk usually present; flesh up to 3 cm thick.Ganoderma tsugae(see G. lucidum, p. 577)
6. Flesh bright reddish-orange to rusty-orange; fruiting body often thick (tall) in relation to
diameter; restricted to juniper .Truncospora demidoffii (see Phellinuspini, p. 582)
6. Not as above . 7
7. Flesh bright yellow-brown to rusty-brown, brown, or dark brown when fresh . 8
7. Flesh whitish to yellowish or straw-colored (or light brown to dingy yellow-brown in age) 18
8. Pore surface white when fresh but turning brown when scratched (or in old age); cap with a hard
surface crust; very common .Ganoderma applanatum group, p. 576
8. Not as above . 9
9. Fruiting body annual (though sometimes large), i.e., with only one tube layer; usually soft or
spongy or exuding water droplets when fresh, but usually tough in age; if found on hardwoods,
the hardwoods usually living .(seePhaeolus, Inonotus, Coltricia, & Allies, p. 566)
9. Not as above; fruiting body very tough (corky or woody) even when fresh, usually with more
than one tube layer (perennial), especially if growing on living hardwoods . 10
10. Typically growing on conifers .Phellinus pini 8c others, p. 582
10. Typically growing on hardwoods . 11
11. Fruiting body typically without a cap (resupinate) or with only a rudimentary one . 12
11. Fruiting body with a cap (upper sterile surface), usually hooflike or shelflike . 13
12. Fruiting body hard and woody; free margin (“cap”) when present often with a surface crust
. . . ..Phellinus laevigatus & P. pomaceus (see P. igniarius, p. 581)
12. Fruiting body corky or fibrous-tough but not woody; free margin when present not incrusted
.Phellinus ferruginosus& P.ferreus (see P. gilvus, p. 582)
13. Fruiting body with a clearly differentiated hard surface crust (at least in older specimens), or
if not then the older (buried) tube layers showing whitish flecks or streaks when sectioned; cap
gray, brown, or black; pore surface brown or gray; especially common in northern regions 14
13. Not with above features . 15
14. Older (buried) tube layers typically stuffed with white hyphae that show as white streaks or
flecks when broken open; old caps often black and cracked.Phellinus igniarius, p. 581
14. Not as above .Fomesfomentarius (see Phellinus igniarius, p. 581)
15. Typically growing on locust or other legumes .Phellinus rimosus(see P. gilvus, p. 582)
15. Not as above . 16
16. Cap ochraceous to rusty-brown (sometimes blackish in old age); flesh usually less than 1.5 cm
thick; usually found on dead wood .:.Phellinus gilvus 8c others, p. 582
16. Cap differently colored and/or flesh thicker; usually found on living trees . 17
17. Flesh bright yellow-brown .Phellinus robustus(see P. gilvus, p. 582)
17. Flesh rusty-brown or darker .Phellinus everhartii(see P. gilvus, p. 582)
18. Growing on incense cedar; pore surface usually yellow or yellowish when fresh; typically with
only one layer of tubes ...(see Tyromyces amarus, p. 601)
18. Not as above; usually growing on a different host . 19
19. Cap distinctly hairy and white to pale ochraceous (or greenish from algae and moss) .... 20
19. Not as above . 21
20. Fruiting body often massive; cap surface coarsely hairy “like a doormat” (Stuntz); found on
conifers in western North America; rare .Oxyporus nobilissimus
20. Not as above; found on hardwoods (especially maple) in eastern states . Oxyporuspopulinus
21. Flesh cheesy when young but soon becoming chalky, very bitter-tasting; cap white to yellowish
or becoming grayish or greenish in old age; found on conifers . Fomitopsis officinalis, p. 579
21. Not as above . 22
22. Underside of cap with mazelike or elongated pores or sometimes even gills; cap 5-15 cm broad,
tube walls often thick; found mainly on hardwoods (see Lenzites, Daedalea, & Allies, p. 586)
22. Not as above . 23
23. Cap surface usually (but not always) partly or entirely reddish to reddish-brown or reddish-
black; cap smooth or grooved; tube layers stratified .Fomitopsispinicola, p. 578
23. Cap surface usually roughened, pitted, knobby, ridged, etc., not normally reddish; tube layers
not usually stratified . . Heterobasidion annosum 8c others (see Fomitopsis pinicola, p. 578)
Artist’s Conk (Ganoderma applanatum group), with my wife Judith Mattoon included for scale.
This large specimen shows ridges and furrows characteristic of many perennial polypores (conks).
The underside is white when fresh and turns brown when scratched (see photo on next page).

Ganoderma applanatum group (Artist’s Conk; Artist’s Fungus)

FRUITING BODY emerging whitish and knoblike, becoming hooflike or shelflike to
somewhat irregular; very hard and woody; perennial. CAP 5-75 cm broad or more, 2-20 cm
thick, usually fan-shaped or semi-circular in outline, with a hard surface crust that is
usually cracked, furrowed, ridged, and/ or lumpy or knobby in age, but not varnished;
surface gray to brown or grayish-brown, sometimes aging grayish-black, but often covered
with brown to cocoa-brown spore powder. Flesh punky or corky, 0.5-5 cm thick, brown or
cinnamon-brown, rarely whitish. PORES barely visible (4-6 per mm), white or whitish
when fresh but instantly turning brown when scratched; often dingy yellowish to brown
when dried or in old age; tube layers distinctly stratified and separated by a thin layer of
chocolate-brown tissue; each tube layer 4-12 or more mm deep. STALK usually absent.
SPORE PRINT brown or reddish-brown; spores 6-9.5 * 4.5-7 microns, broadly elliptical
to slightly truncate at apex, thick-walled, appearing minutely spiny.
HABIT AT: S olitary or in groups on hardwood logs and stumps, or growing from wounds
in living trees (usually near the ground); also common on conifers in some areas. Very
widely distributed, found year-round. Its hosts include virtually every hardwood found in
North America, plus numerous conifers. In coastal California it (and close relatives—see
comments) is especially common on bay laurel, but can also be found on oak, magnolia,
pepper trees, acacia, eucalyptus, elm, and Douglas-fir. Along with Fomitopsispinicola, it
is the commonest conk in our area. The only regions where it seems to be absent are those
where there aren’t any trees! It usually attacks dead or dying trees, but can also be parasitic,
producing a whitish, delignifying decay of the sapwood and heartwood. Infected trees blow
over easily, so if one appears on a tree near your house—watch out!
EDIBILITY: Much too tough and woody to be worthwhile (see edibility of Fomitopsis
pinicola)', however, sturdy specimens can be made into stools.
COMMENTS: This is the hard, woody growth you see so often at the bases of bay laurels
and other hardwoods. Since the brown-staining of the pore surface is reasonably per¬
manent, it makes an excellent medium for etching( or better yet, leaving cryptic messages in
the woods)—hence its popular name, artist’s conk. It’s been calculated that a large
specimen (I’ve found one weighing 26 pounds) liberates 30 billion spores a day, 6 months a

576
Ganoderma applanatum group, underside (pore surface). It is sometimes called the “Artist’s Conk”
because the white pore surface turns brown when scratched. This means you can draw pictures on it, or
better yet, leave messages such as this one in the woods!

year—or over 5,000,000,000,000 (5 trillion!) spores annually. Millions of these may be

borne aloft by air currents and deposited on top of the cap, turning it brown. G. applanatum
as discussed here is actually a “collective” species, embracing at least two others: G.
annularis, with unstratified tubes and no flesh; and G. brownii (-G. adspersum? G.
europaeum?), with thicker(3-8 cm), darker flesh. Both have longer spores(9-12 microns)
than the “true” G. applanatum. The latter is especially common in our area on bay laurel,
perhaps more so than G. applanatum. All of these conks used to be placed in Fomes and
have also been put in Elfvingia.

Ganoderma lucidum (Varnished Conk; Ling Chih)

FRUITING BODY annual, corky and tough; often emerging as a whitish or pallid knob
but soon becoming shelflike or developing a cap and stalk. CAP 2-20 (35) cm broad, 4-8
cm thick, circular to semi-circular to fan-shaped or kidney-shaped in outline; surface
usually with a varnished (shiny) surface crust, smooth or often concentrically zoned and
grooved; color variable: dark red to reddish-brown, orange-brown, mahogany, or
reddish-black, but often ochre or yellowish toward the margin (which is often white when
actively growing); surface sometimes covered with brownish spore powder. Flesh ochra-
ceous-brown to dark brown, or pallid near the cap and brownish near the tubes; soft-
corky or punky when fresh, tough when dry or old. PORES minute(4-7 per mm), whitish or
yellowish-white when fresh, usually bruising or aging brown; tubes 2-20 mm long, one
layer only (rarely two). STALK sometimes absent, but often present; usually attached
laterally, but often vertical and well-developed, 3-14 cm long, 0.5-3 (4) cm thick; often
gnarled or twisted, equal or enlarged below; dark red to reddish-black and appearing
varnished like the cap. SPORE PRINT brown; spores 7-13 * 5-8 microns, elliptical,
double-walled, appearing minutely roughened.
HABITAT: Solitary or in small groups at bases of living hardwoods or on stumps and
roots, very rarely on conifers; fruiting during mushroom season but often persisting year-
round. Distribution worldwide—in both the tropical and temperate zones, from Canada
to Argentina and Europe to Siberia, China, India, Australia, and Africa. It is fairly
common in eastern North America but rather infrequent in California. It occurs on a wide

577
578 POLYPORACEAE & ALLIES

range of hosts, but in North America is partial to maple. The very similar G. tsugae and
G. oregonense occur on conifers (see comments). It is sometimes parasitic, but can also
be saprophytic, producing a wet white rot in its host.
EDIBILITY: Too tough to be edible, but in the Orient it is pictured in many classical art
works and has been revered for centuries as a symbol of success and well-being(Ling Chih
means “marvelous herb” or“mushroom of immortality”). It has been used in the treatment
of cancer and various other maladies, and is believed by some to have the power of arousing
the dead. One method of preparation is to soak the fruiting body in wine for several
months. The resulting liquid “essence” or elixir is then drunk or put into candies. The
varnished fruiting bodies are also used for decorative purposes.
COMMENTS: This striking polypore and its close relatives are easily recognized by
their shiny ochre to reddish-black caps that look as if they had been artificially varnished
or shellacked (see Color Plate 156) unless old or covered by spore powder. Like Piptoporus
betulinus, the fruiting bodies are annual (with one tube layer) but persist for months.
T wo extreme growth forms occur: one is fairly large, often sessile( with little or no stalk) and
particularly common in North America. The other is smaller(cap rarely over 15 cm broad),
has a long slender stalk, and appears to be more common in the tropics and Old World.
H owever, a large number of intermediate forms also occur, plus occasional abnormal ones
in which the fruiting body is branched and antler- or tree-like, and these are all currently
thought to be forms of one highly variable, farflung species. G. curtisii is a similar species
with an ochre to whitish or only partly reddish cap that often lacks the varnish in age. It
typically has a stalk and grows on hardwoods in eastern (especially southeastern) North
America. G. tsugae (COLOR PLATE 156) is also very similar to G.lucidum, but has
white flesh and grows only on conifers, particularly hemlock, in northern North America.
Still another similar species, G. oregonense, also grows on conifers, but is usually
larger (cap 5-100 cm broad, 2-20 cm thick!), with larger spores and a somewhat darker or
slightly duller cap. It is the most common “Varnished Conk” of Washington, Oregon, and
California. All of these can be distinguished from Fomitopsis pinicola by their punkier
(softer) flesh, annual fruiting body, and brown-staining pore surface (when fresh).

Fomitopsis pinicola (Red-Belted Conk)

FRUITING BODY emerging as a whitish, pale yellow, or lilac-tinged knob, becoming
hoof-shaped or shelflike or sometimes bracketlike; hard and woody in age. CAP 5-40 (75)
cm broad, 3 -22 cm thick, usually fan-shaped to semi-circular in outline; surface developing
a thin, hard, resinous crust in age which is sometimes slightly varnished; usually at least
partially reddish to dark red, but sometimes brown and often rusty or blackish-brown
toward the base and brightly colored (white, yellow, ochraceous, or reddish) at the margin;
concentrically furrowed and/ or zoned in age; growing margin rounded, thick, and blunt.
Flesh corky or woody, very tough, white to pinkish-buff or yellow when young, pale
brownish or straw-colored in old age; usually bruising pinkish when actively growing; odor
when fresh rather strong and fungal. PORES minute (3-5 per mm), white or pale yellow
becoming brownish in old age; not turning brown when scratched but sometimes bruising
yellow or pinkish-lilac; tube layers distinctly stratified, each layer 2-8 mm deep. STALK
typically absent. SPORE PRINT white or pale yellowish; spores 5-8 x 3.5-5 microns,
cylindrical to elliptical, smooth. Flesh staining reddish to dark reddish-brown in KOH.
HABITAT: Solitary or in groups on dead trees, logs, and stumps or rarely on living trees,
perennial; very common and widely distributed. It has been recorded from a wide range of
hosts but is found mainly on conifers (in our area primarily Douglas-fir and redwood). It
attacks both the heartwood and sapwood of its host, producing a slow carbonizing rot
which fractures the wood and turns it brown (“brown crumbly rot”). Large mycelial mats
can often be seen in the fractured wood.
Left: Fomitopsis pinicola, one of the most common of all the conks (top view). Right: A large, old
specimen of Fomitopsis officinalis (described below). Note how small the penny is!

EDIBILITY: This woody conk is not often eaten, but in a pinch you can use the following
recipe developed by my wife: “Saw into 2-inch cubes, then marinate in olive oil and
dandelion wine for at least 48 hours (be sure to use LOTS of garlic!). Roast slowly on
skewers over charcoal indefinitely (minimum time: 20 hours). Cool. Pound vigorously
with a large mallet between two pieces of thick leather. Pulverize in a meat grinder and
then force through a braced sieve (allow several hours for this step). Wrap the resulting
mess in several thicknesses of cheesecloth and hang up someplace high and out of the way
(on a clothesline or TV antenna). Allow to dangle thus for at least one week. (Aging has
a mellowing effect, so you may want to try one year.) Wring periodically, making sure
to reserve the drippings for gravy or as a motor oil additive. To eat, boil for twenty-four
hours, squeeze thoroughly, garnish with gravel, and serve forth.”

COMMENTS: This beautiful but cosmopolitan conk is a major destroyer of dead coni¬
ferous timber. The reddish or partially reddish cap with a brightly colored margin (when
growing) and pallid pores that do not bruise brown are good fieldmarks. Varnished spe¬
cimens might be confused with Ganoderma tsugae or G. oregonensis{see G. lucidum), but
are perennial and much harder and denser. Fomespinicola and Ungulina marginata are
synonyms. Another common, widespread woody conk with whitish flesh and white to
yellowish pores is Heterobasidion annosum (-Fomes annosus). It has a bracketlike to
shelflike to resupinate or irregularly knobby fruiting body that, when shelflike, is usually
thinner and rougher (knobby, pitted, grooved, etc.) than F. pinicola. The cap is usually
brown to grayish-brown with a pallid growing margin, but is sometimes reddish-brown
(and can be whitish when very young) and its flesh does not redden in KOH. It is a frequent
parasite of conifers (or occasionally hardwoods) and usually grows at the base of the trunk
or from its roots. It is especially common on second-growth mountain conifers(e.g., pon-
derosa and Jeffrey pines). Two other species with whitish to yellow-brown flesh,F.fraxi-
nophilus and F. ellisianus, somewhat resemble H. annosum but have different hosts:
the first favors ash trees, while the second grows on buffalo berry (a shrub).

Fomitopsis officinalis (Quinine Conk; Quinine Fungus)

FRUITING BODY perennial, emerging as a whitish knob, then becoming convex and
finally hoof-shaped to cylindrical; hard and tough in age. CAP 4-30 cm broad and 5-40
cm or more thick (high); surface with a thin crust, often cracked and/ or furrowed in age;

579
580 POLYPORACEAE & ALLIES

white to yellowish, but aging grayish and sometimes with a greenish covering of algae.
Flesh thick, white, cheesy when young but chalky or friable (crumbly) in mature or old
specimens; odor farinaceous; taste very bitter. PORES 34- per mm, white or whitish when
fresh, discoloring in age or drying; tube layers each 3-20 mm long, often stratified. ST ALK
absent. SPORE PRINT whitish; spores 4-5.5 * 3-4 microns, broadly elliptical, smooth.
HABITAT: Solitary or several on living and dead conifers; common year-round
throughout much of the W est, especially on larch and pine, but also on spruce, fir, hemlock,
and Douglas-fir. It apparently does not occur in our area, but I have seen it on sugar pine
and ponderosa pine on the slopes of theSierraNevada.Itis said to be one of the three major
destroyers of standing coniferous timber in the West (Phellinus pini and Phaeolus
schweinitzii are the other two). It causes “felted heart rot,” an extensive carbonizing trunk
rot with thick mycelial mats often several feet long.
EDIBILITY: Friable but not fryable—it is inedible due to the bitter taste and tough
texture, but has been used as a laxative and quinine substitute. (It does not have anti-
malarial properties, but was thought to because of its taste.) Since the fruiting bodies often
grow high above the ground, commercial “quinine” collectors used to dislodge them with
rifles.
COMMENTS: Also knownas Laricifomes officinalis and Forties officinalis, this destruc¬
tive conifer-killer is easily recognized by its pale color, chalky white flesh, bitter taste,
and sometimes massive size. New fruiting bodies are convex, but add layers yearly until
they are hoof-shaped if growing on slash or vertically elongated (cylindrical) if growing
high up on living trees (see photo at top of p. 579). Specimens 60 cm (2 ft.) in height with
more than 70 tube layers have been recorded!

Fomitopsis cajanderi (Rosy Conk)

FRUITING BODY often perennial; shelflike or bracketlike to somewhat hoof-shaped,
tough in age. CAP 2.5-10 (13) cm broad, 0.3-2 cm thick; surface covered with hairs,
becoming nearly bald in age (but often roughened), not incrusted; pinkish-red to pinkish-
brown becoming brown to grayish-brown and in old age blackish except for the margin;
often zoned or grooved concentrically; margin usually thin, acute. Flesh rosy-pink to
reddish-brown or pinkish-brown, rather soft when fresh but corky in age. PORES minute
(3-5 per mm), rosy to pinkish-red or pinkish-brown when fresh, often duller and darker
(reddish-brown) in age; tube layers not distinctly stratified; each layer 1-3 mm deep.
STALK absent. SPORE PRINT whitish; spores 4-8 * 1.5-2.5 microns, cylindrical but
slightly curved (sausage-shaped), smooth.
HABITAT: Usually in colonies on dead conifers, but also reported on madrone and
various fruit trees; widely distributed. It is fairly common in our area, especially on
Douglas-fir, and is usually but not always perennial. I find it most often on the cut ends of
recently felled trees. It produces a carbonizing brown pocket rot but is of minor economic
importance.
EDIBILITY: Inedible (but see comments on edibility of F. pinicola).
COMMENTS: The beautiful rosy pore surface is the outstanding feature of this out¬
standing polypore, which is better known asFomes subroseus. The upper surface is not as
hard and crusty as that of many species of Fomitopsis and Fomes, and the fruiting bodies
are not so obviously perennial—therefore it is apt to be looked for in another genus. Other
species: F. rosea is practically identical, but has paler (silvery-pink to pale rose) flesh,
slightly broader, cylindrical (not curved) spores, and a cap surface that is sometimes
incrusted slightly in older specimens. It also favors conifers, causing a top rot of dead trees.
Phellinus igniarius. The black cracked cap is typical of older specimens. The fruiting body can
be hoof-shaped as well as shelflike.

Phellinus igniarius (False Tinder Polypore; False Tinder Conk)

FRUITING BODY perennial, soon shelllike or bracketlike to hoof-shaped; hard and
woody. CAP 5-20 cm or more broad, 2-12 (20) cm thick; surface usually with a crust, at
least in older specimens; brown and finely hairy or velvety when young, soon becoming
bald and grayish, then finally black; often cracked, furrowed, and/ or knobby in age;
margin brown and velvety when actively growing. Flesh hard, woody, rusty-brown to
brown; taste sour or bitter. PORES minute, 4-5 per mm, grayish-brown to brown; tube
layers each 2-5 mm long but not always stratified; tubes and pores of older(buried) layers
often stuffed with white mycelial threads that show as streaks or flecks when cut open.
ST ALK absent or rudimentary. SPORE PRINT whitish; spores 5-7 * 4-6 microns, round
or nearly round, smooth. Brown sterile cells (setae) sometimes abundant among the
basidia. Cap tissue blackening in potassium hydroxide (KOH).
HABITAT: Solitary or in small groups on hardwood trunks (usually living); widely
distributed. It is especially common on birch and aspen in the northern half of North
America, and occurs year-round. In our area it grows on alder, madrone, manzanita,
maple, and willow, but in my experience is not common. It can be quite destructive, causing
an intensive white heart rot that reduces its host to a “soft, spongy, whitened mass.”
EDIBILITY: Inedible unless you are fond of wood(see recipe under Fomitopsis pinicola).
COMMENTS: This conk is easily recognized by its brown to grayish-black cap and
pores, brown to rusty-brown flesh, velvety brown actively-growing margin, and whitish-
flecked or streaked tubes when old. A smaller variant, often called P. tremulae, is com¬
mon on aspen. P. laevigatus is a closely related resupinate variety that usually grows
on birch; P. pomaceus, often resupinate, causes extensive heart rot in fruit trees (Prunus),
especially wild ones; P. robustus(see comments under P. gilvus) is similar, but has yellow-
brown flesh and favors oak or eucalyptus. Finally, there is the “true” tinder polypore or
“Amadou,” Fortiesfomentarius. This hoof-shaped fungus resembles P. igniarius in color
but has a hard thick surface crust, non-stratified tubes without white mycelial threads,
and cylindrical spores 14-20 microns long. It grows on dead hardwoods (especially birch
and maple) or from wounds in living trees, and is widely distributed. Unlike Ganoderma

581
582 POLYPORACEAE & ALLIES

applanatum, its pore surface is not white and does not turn brown when scratched. As its
name implies, it has been used for centuries to ignite fires. Since the tubes are not stratified,
they are quite long and readily soak up liquids through capillary action. Chunks of the
tube layer were soaked in a salt peter solution, then dried and used as “matches.”

Phellinus gilvus (Oak Conk)

FRUITING BODY shelflike or bracketlike, often perennial; tough and corky when
fresh. CAP 2.5-15 cm broad and 1-3 cm thick; fan-shaped or semi-circular in outline; sur¬
face at first velvety (just the growing margin if perennial), becoming rough to nearly
smooth and/ or somewhat zoned; bright rusty-yellow to ochraceous when young(as is the
growing margin of older specimens), dark rusty-brown and finally blackish in age; not in-
crusted. Flesh tough, bright ochraceous to dark yellow-brown or colored like cap. PORES
minute (4-8 per mm), grayish-brown becoming reddish-brown or dark brown; tubes in 1 -5
layers, each 1-5 mm long. STALK absent. SPORE PRINT whitish; spores 4-5 x 2.5-3.5
microns (but rarely found), oblong-elliptical, smooth. Large brown sterile cells (setae)
abundant among the basidia. Cap tissue blackening in potassium hydroxide (KOH).
HABITAT: Solitary or more often in colonies on dead or occasionally living hardwoods
(rarely on conifers); widely distributed. It occurs year-round in our area on oak and tan-
oak and is our most common perennial polypore of dead hardwoods. It produces a
general delignifying decay of the sapwood, rendering it whitish and very brittle.
EDIBILITY: Unknown.
COMMENTS: The dark rusty-brown color and velvety yellow-ochre growing margin
are the fallible fieldmarks of this common conk. The upper surface is not crusty as in many
conks, and the color and texture are somewhat reminiscent of old Phaeolus schweinitzii,
which grows with conifers( usually on the ground). A specimen of P. gilvus can be seen in the
photograph at the bottom of p. 887. Other Phellinus species that favor hardwoods include:
P. robustus, widespread, especially on oaks, large and woody but not incrusted, with a
gray-brown to blackish cap, bright yellow-brown flesh, and stratified tubes (it also grows
on eucalyptus, pittosporum, walnut, cactus, various other hardwoods, and on conifers);
P. everhartii (see photo at bottom of p. 583), very hard and woody, up to 40 cm broad,
growing mainly on oak, with unstratified tubes, brown spores, and setae;P. rimosus(-P.
robiniae), with a rich brown cap, growing on locust, acacia, and other legumes; and P.
ferruginosus and P.ferreus, common on dead hardwoods and rather similar to P. gilvus,
but with a rusty-brown resupinate (capless) fruiting body. Most of these were once placed
in Pomes, but now belong to Phellinus because they darken in KOH, have rusty-brown to
tawny flesh, and often possess setae. For conifer-loving species, see P. pini, and for a similar
but more brightly colored annual species, see Inonotus radiatus (under I. hispidus).

Phellinus pini (Pine Conk)

FRUITING BODY perennial; shelflike, hootlike, or bracketlike, very tough or woody.
CAP 2-20 cm broad and 1 -15 cm thick; hoof-shaped to convex or fan-shaped; surface hard,
often crusty but not shiny; rough or cracked, minutely hairy or roughened( like sandpaper)
at first, often concentrically grooved in age; tawny to rusty-brown becoming brown to
reddish-brown or brownish-black in age; margin sometimes brighter. Flesh less than 1 cm
thick, tough, tawny to rusty-reddish or ochre. PORES 2-5 per mm, round to irregularly
sinuous, ochraceous-tawny to rusty-brown becoming brown; tubes 2-5 mm long, in one or
several layers. STALK absent or rudimentary. SPORE PRINT brown; spores4-6 * 3.5-5
microns, round or nearly round, smooth. Large brown sterile cells (setae) intermingled
with basidia. Cap tissue blackening in potassium hydroxide (KOH).
HABITAT: Solitary or more often in rows or columns up and down living or recently
Phellinus pini is a major parasite of conifers, particularly pines. The tough fruiting bodies often
form lines along the entire length of the trunk. Note the somewhat sinuous pores.

conifers; widely distributed and common, infecting all important members of the pine
family (pine, Douglas-fir, etc.). Perennial, in our area occurring mainly on pine. It attacks
the heartwood and sometimes the sapwood of living trees, resulting in a delignifying pocket
rot known as “conch rot.” The fruiting bodies are largely confined to older trees, partly
because years of growth must take place before the mycelium can fruit. As a result, the
appearance of a single fruiting body means that extensive heart rot has already taken place
10-15 feet above and below it. This fungus is said to cause more timber loss than any other.
EDIBILITY: Inedible (but see comments on edibility of Fomitopsispinicola).
COMMENTS: The rough unpolished cap, frequently sinuous pores, ochraceous to rust-
colored flesh, and growth on conifers are the fallible fieldmarks of this destructive fungus.
It varies considerably in size, from large hoof-shaped specimens to moderately sized ones
(see photo), to a small thin shelving form that often occurs in fused masses and is considered
a distinct species, P. chrysoloma, by many investigators. Other species: Truncosporademi-
doffii (also known asPulvifomes, Pyrofomes, or Forties juniper inus) is somewhat similar,
but has rusty to reddish-orange flesh and grows only on juniper;/*, texanus resembles a
small P. robustus( see comments under P. gilvus) but also grows on juniper, while a resu-
pinate form of P. robustus with bright yellow-brown flesh often grows on conifers; P.
taxodii causes heart rot in bald cypress; P. nigrolimitatus, common on conifers in the
Rocky Mountains, usually has a spongy upper layer of tissue and fine black lines running
through its flesh; P. torulosus looks like P. gilvus, but has yellow-brown pores and grows
on conifers in Arizona. For hardwood-loving species, see P. gilvus and P. igniarius.

An old Phellinus everhartii(see comments under P. gilvus). Bark protects a tree from infection, but
nailing a signboard to a tree (as shown here) or otherwise defacing it can allow fungal spores to enter!
584 POLYPORACEAE& ALLIES

PIPTOPORUS & CRYPTOPORUS

THESE two oddball genera are each represented in North America by a single farflung
species. Piptoporus has a conklike (but annual) fruiting body whose margin projects
below the pore surface to form a “curb.” In Cryptoporus this theme is carried to its illogical
extreme: the margin of the cap is so ingrown (or overgrown) that it completely covers the
pore surface, thus hiding it from view!

Key to Piptoporus & Cryptoporus

1. Pore surface exposed; cap margin curblike; found on birch .P. betulinus, below
1. Pore surface completely hidden (at least until old age); found on conifers C. volvatus, p. 585

Piptoporus betulinus (Birch Conk; Birch Polypore)

FRUITING BODY annual, nearly round becoming shelflike or hooflike at maturity;
tough or corky when fresh, rigid and hard when dry. CAP (2.5) 5-25 cm broad, 2-6 (10) cm
thick, kidney-shaped to nearly round in outline, convex to nearly plane; surface covered
by a thin, smooth or suedelike, white to buff, tan, brown, or grayish-brown crust that often
breaks up into scales or flat patches or wears away, revealing the whitish undersurface;
margin thick, blunt, inrolled, curblike (projecting below the pore surface), sometimes
wavy. Flesh punky or corky, thick, white. PORES appearing recessed due to curblike
margin, 2-4 per mm; white when fresh, in age becoming pale brown or grayish-brown and
occasionally torn up or toothlike; tubes 2-10 mm long, one layer only. STALK absent or
present only as a stubby extension of the cap; lateral or attached to top of cap. SPORE
PRINT white; spores 3-6 * 1.5-2 microns, cylindrical to sausage-shaped, smooth.
HABITAT: Solitary or in groups or columns on dead or sometimes living birch trees;
common year-round in the northern hemisphere throughout the range of birch. I have not
seen it in California, undoubtedly because birch does not occur naturally. However, it
is found in Washington and Idaho and is very abundant in northeasternNorth America. It
causes a reddish-brown to yellow-brown carbonizing decay.
EDIBILITY: Edible when young (according to Mcllvaine), but tough. The fruiting bodies
are quite attractive, however, and have enjoyed a variety of non-culinary uses, e.g., as

Piptoporus betulinus. Note the thick blunt margin and recessed pore surface of this unique birch-
loving polypore.
PIPTOPORUS 585

tinder, as a razor strop, and as a mounting medium for pinned insect specimens.
COMMENTS: Formerly known as Polyporus betulinus, this common and distinctive
polypore is easily told by its recessed pore surface with a curblike margin (see photo) plus
its white to tan or grayish-tan color and growth on birch. The fruiting bodies are annual
in the sense that they possess only one tube layer; however, they remain intact for many
months and as a result can be found year-round.

Cryptoporus volvatus (Cryptic Globe Fungus; Veiled Polypore)

FRUITING BODY annual, tough or corky, more or less round to oval to slightly com¬
pressed or hooflike; 1.5-8.5 cm broad, with a hollow interior. CAP(upper or outer surface)
with a thin, smooth, glazed or resinous crust; whitish to warm tan or yellowish, drying
darker(ochre-brown to reddish-brown); margin extending down and under to form“veir
which completely covers pore surface; in age the underside perforated by one (rarely two)
holes. Flesh whitish, tough; odor often fragrant (like Sparassis). PORES hidden by the
“veil,”3^4- per mm, white becomingpinkishorbrownishinage;tubes2-5 mmlong.STALK
absent. SPORE PRINT pinkish or flesh-colored; spores often collecting in a heap on the
inner “floor” of fresh specimens), 8-12 * 3-5 microns, cylindrical to elliptical, smooth.
HABITAT: Solitary or more often in groups on dead or occasionally old living conifers
throughout northern North America. Common year-round in our area, especially on
pine. It seems to favor standing or recently felled trees and produces a delignifying decay
that scarcely damages its host.
EDIBILITY: Too tough to be edible. However, Alexander Smith says that “worms”
(insect larvae) found inside the fruiting bodies can be used as fishbait. So can worms
found outside the fruiting bodies.
COMMENTS: This bizarre evolutionary anomaly looks like a cross between a confused
puffball and a bemused oak gall. The smooth, warmly tanned exterior is quite attractive
(often reminding me of a small loaf of bread) and gives no hint of the tube layer within.
Slicing it open, however, reveals a hollow interior with a “ceiling” of tubes. The “floor”
eventually ruptures and tiny bark-boring beetles enter the “trap door” in search of tasty
tube tissue and spores. After feasting they depart to construct brood tunnels in old or
dying conifers, and the spores they carry with them gain entry to a new host. Later, fruiting
bodies may emerge through the very holes bored by the beetles!

Cryptoporus volvatus can be hoof-shaped (like these specimens here) or spherical. The tube layer is
inside the fruiting body, as shown in sectioned specimen at right.
Close-up of the pore surface in Daedaleopsis confragosa (p. 588). Note how the pores range from
round (bottom left) to elongated or mazelike.

LENZITES, DAEDALEA,& Allies

Fruiting body small to medium-sized, tough and leathery or sometimes woody, usually annual
(but often perennial in Daedalea), growing shelflike or bracketlike on wood. CAP zoned or
unzoned, variously colored, hairy to smooth. PORES usually mazelike, sinuous, or meandering, or
forming plate-like gills. STALK absent or rudimentary. SPORE PRINT whitish to brown, but
difficult to obtain. Spores usually cylindrical, smooth.

THESE corky or leathery fungi can be told from Trametes and other bracketlike polypores
by the configuration of their spore-producing surface. In Daedalea and Daedaleopsis
(derived from the Greek word daedalus, meaning “maze”) the pores are usually long and
sinuous or mazelike and have relatively few cross-walls. In Lenzitesand Gloeophyllum, on
the other hand, the cross-walls have all but disappeared, leaving plate-like gills instead of
pores! However, the configuration varies considerably, especially in Daedalea and
Daedaleopsis, leading to confusion with numerous other polypores (when the pores are not
distinctly mazelike), and with species such as Phellinus pini, a “conk” with somewhat
sinuous pores.
Like most polypores, the species treated here are too tough to eat. If your polypore has
gills, it should key out here. If it has mazelike pores, however, it may not key out
convincingly, in which case other groups of polypores should be checked.

Key to Lenzites, Daedalea, & Allies

1. Flesh and gills or pores yellow-brown to reddish-brown or brown when fresh; flesh typically
at least 1 mm thick and usually darkening or blackening in potassium hydroxide (KOH) . 2
1. Not as above; flesh white or pale-colored when fresh (but may become pale brown in old age or
after weathering), or if brown when fresh then typically less than 1 mm thick; pores or gills
variable in color (white, violet-tinged, brown, gray, blackish, etc.) . 5
2. Typically growing on hardwoods (or occasionally on conifers such as bald cypress); flesh thin
. Gloeophyllum trabeum (see G. saepiarium, p. 590)
2. Typically growing on conifers (but occasionally on hardwoods) . 3
3. Fruiting body usually with an anise-odor when fresh; underside of cap usually with pores . . .
.Osmoporus odoratus (see Gloeophyllum saepiarium, p. 590)
3. Not as above .4

586
LENZITES, DAEDALEA, & ALLIES 587

4. Underside of cap usually with gills; cap surface often (but not always) with white, yellow, or
orange tones or zones when fresh; very common Gloeophyllum saepiarium & others, p. 590
4. Underside of cap usually with sinuous or mazelike pores; cap dark or dull brown .
.Daedalea berkeleyi (see D. quercina, below)
5. Cap surface decidedly hairy, woolly, or velvety when fresh (but may be nearly smooth in old
or very weathered specimens) .6
5. Cap surface not hairy or only very slightly so when young . 11
6. Underside of cap typically composed of gills or plates (but sometimes varying to mazelike) 7
6. Underside of cap typically composed of elongated pockets or mazelike pores (but sometimes
forming gills or regular pores) .9
7. Gills crisped (crimped), usually forked or shallow and veinlike; cap up to 2.5 cm broad, typically
not zoned concentrically; found on hardwoods in eastern North America .
. (see Plicaturopsis crispa under Schizophyllum commune, p. 590)
7. Not as above; larger or cap concentrically zoned or found on conifers or in West.8
8. Gills often violet-tinged when fresh; cap often grayish to blackish in old age, zoned or unzoned;
typically found on conifers (see Trichaptum abietinus var. abietis under T. abietinus, p. 593)
8. Not as above; cap usually zoned; usually found on hardwoods .Lenzites betulina, p. 589
9. Fruiting body soft or watery when fresh, discoloring rusty or reddish after handling, or if not,
then often quite thick (3-8 cm); fresh pore surface whitish . (see Tyromyces& Allies, p. 597)
9. Not as above; fruiting body usually quite thin and tough when fresh, not soft and watery; pore
surface often gray to brown or blackish, at least in age . 10
10. Cap brown to blackish and velvety when fresh; tubes often unequal in length or appearing slot¬
like (see photo on p. 596) .(see Datronia mollis, p. 595)
10. Cap white to grayish, brown, greenish, etc., usually hairy; tubes often breaking up in age to
form “teeth” . Cerrena unicolor (see Daedaleopsis confragosa, p. 588)
11. Gills when present thick, blunt, and widely spaced (1 mm or more apart); pores when present
1 mm or more broad, the walls thick and blunt . 12
11. Gills when present thinner and closer (less than 1 mm) than above; pores when present smaller
(averaging 1-3 per mm), the walls thin . 14
12. Typically found on hardwoods .Daedalea quercina, below
12. Typically found on conifers (occasionally on hardwoods) . 13
13. Found on juniper .Daedalea juniperina (see D. quercina, below)
13. Found on other conifers; cap often poorly developed .
.(see Coriolellus heteromorpha under Poria corticola, p. 603)
14. Cap white, at least when fresh .Daedaleopsis ambigua (see D. confragosa, 588)
14. Cap pallid to brown, grayish, reddish-brown, etc.Daedaleopsis confragosa, p. 588

Daedalea quercina (Thick-Walled Maze Polypore)

FRUITING BODY usually perennial; shelflike, rigid and very tough or corky. CAP4-20
cm broad and 1.5-8 cm thick, more or less fan-shaped; convex or plane; surface uneven
or roughened, usually concentrically furrowed or zoned in older specimens (from new
growth layers); whitish to tan, ochraceous, grayish, or brown (usually quite pale when
fresh), often blackening and cracking in old age; margin often thick. Flesh very tough and
corky; white to buff, ochraceous, or pale brown (never dark). PORES usually greatly
elongated and pocket- or mazelike, sometimes even forming gills; whitish to buff, tan,
or dull ochre; tube walls ( or “gills”) thick (1 mm or more), the spaces between them at
least 1 mm broad; tubes 0.5-3 cm long, the layers not distinctly stratified. STALK absent
or rudimentary. SPORE PRINT whitish; spores 5-7.5 * 2-3.5 microns, cylindrical to
elliptical, smooth.

HABITAT: Solitary or sometimes in shelving groups on dead or living hardwoods, espe¬

cially oak, chestnut, and chinquapin; widely distributed, but especially common in eastern
North America. I have seen it in northern California and Oregon, but have yet to find it in
our area. It causes a brown heart rot, and the tough fruiting bodies occur year-round.
Left: Maze like pore surface of Daedalea quercina. Note the thick walls separating the “pores,” and
compare them to the thinner walls of Daedaleopsis confragosa (photo on p. 586). Right: Close-up
of the thick gills of Gloeophyllum saepiarium (see description on p. 590).

EDIBILITY: Much too tough to be edible.

COMMENTS: The color of this species is rather variable, but the configuration of its
spore^bearing surface is relatively constant (i.e., mazelike), and not nearly as polymorphic
as that of Daedaleopsis confragosa. In addition, the tube walls are thicker and often
deeper (see photo) and the fruiting body is usually perennial. Daedalea berkeleyi is
a southern species that grows on dead conifers and has dark brown to dark rusty-brown
flesh. Daedalea juniperina is a widespread but smaller pale-fleshed species that grows
on juniper. Both of the above tend to have mazelike pores and neither is worth eating.

Daedaleopsis confragosa (Thin-Walled Maze Polypore)

FRUITING BODY usually annual, shelflike or sometimes bracketlike, leathery or corky
when fresh, rigid when dry. CAP (3) 5-15 (22) cm broad, fan-shaped to semi-circular in
outline, broadly convex to plane; surface dry, smooth or slightly hairy, usually zoned or
ridged concentrically, often radially wrinkled or bumpy in age; reddish-brown to brown
to grayish, sometimes blackish in old age; margin thin, acute. Flesh white to pinkish or
brownish, tough. PORES 0.5-1.5 mm in diameter, usually elongated and mazelike with
relatively thin walls, but sometimes circular and at other times forming gills or becoming
toothlike in age; white to tan or brown, sometimes bruising pinkish or reddish; tubes up to
1.5 cm long. STALK absent. SPORE PRINT white; spores 7-11 x 2-3 microns, cylin¬
drical, smooth.
HABITAT: Solitary or in groups on dead hardwoods or from wounds in living trees,
occurring year-round (sometimes perennial); widely distributed. Its favorite host is willow,
but it also occurs on birch and other hardwoods and very rarely on conifers. I have yet to
find it in our area, perhaps because it is not fond of oak. However, it occurs in the Pacific
Northwest and is very common in eastern North America. It produces a delignifying decay
of the sapwood.
EDIBILITY: Not edible.
COMMENTS: The maze-like (“daedaloid”) pattern of the pores is characteristic of this
species and several others, all of which were originally placed in a single genus, Daedalea.

588
DAEDALEOPSIS 589

The configuration of the pores is extremely variable, however, leading to confusion with
Lenzites and Gloeophyllum (when they form gills), and dozens of other polypores (when
they are not noticeably daedaloid). This species can usually be recognized, however,
by its relatively thin, corky, fan-shaped, brown to grayish cap with a bald, often zoned
surface. The depth of the tubes and thickness of the cap are also quite variable, depending
on how old the fruiting body is, but the tube walls are never as thick as those of Daedalea
quercina (see photo at top of p. 586). Daedalea confragosa is an older name for it. Other
species: Daedaleopsis ambigua is a similar southern species with a whiter cap. Cerrena
(-Daedalea) unicolor has a thin, hairy, white to grayish (or greenish from a coating of
algae) cap and pallid to grayish or even blackish mazelike pores that often break up to
form small “teeth.” It is common on dead hardwoods in many regions, but not ours. For
other “daedaloid” species, see Daedalea quercina and Gloeophyllum saepiarium.

Lenzites betulina (Gilled Polypore)

FRUITING BODY usually annual; shelflike, bracketlike, or forming rosettes; tough and
leathery. CAP 2-13 cm broad, nearly round to fan-shaped in outline; surface dry, velvety
or hairy, with narrow concentric zones or grooves of various colors: whitish, tan, buff,
gray, brown, yellow-brown, dull orange, etc. (or in old age often greenish from algae).
Flesh thin (1-2 mm), tough, white. GILLS platelike, often branching toward the margin
or forming elongated pockets (especially in young specimens); white or whitish, drying
dingy yellowish or darker, often wavy in age. STALK absent or rudimentary. SPORE
PRINT white; spores 4-7 * 1.5-3 microns, cylindrical to sausage-shaped, smooth.
HABITAT: Scattered or more often in overlapping rows, columns, or shelving masses on
rotting hardwood logs and stumps (rarely on conifers); widely distributed and very
common in our area throughout the year, often sharing logs with Trametes versicolor and
Stereum hirsutum. The species epithet means birch, but it grows on a wide range of hard¬
woods, especially oak and willow. It produces a delignifying decay of the sapwood.
EDIBILITY: Inedible, but dries nicely. If you are adamant about trying it, see comments
on the edibility of Trametes versicolor for cooking suggestions.
COMMENTS: This species is an excellent example of convergent evolution—a gilled
fungus that is not an agaric. In all respects save the gills it is a typical polypore—the multi¬
colored, hairy, zoned cap bearing an uncanny resemblance to Trametes species. In fact, it
sometimes can’t be distinguished from those mushrooms withoutexaminingthe underside
of the cap. It can also be confused with Gloeophyllum, which has brown gills, and Daedalea
and Daedaleopsis, which have“daedaloid”(elongated or mazelike) pores. In eastern North
America, L. betulina often shows orange tones on the cap, whereas in our area it is usually
quite dull in color (grayish, buff, etc.).

Lenzites betulina is a common hardwood-loving polypore with a hairy zoned cap and gills instead
of pores. It is often found with Trametes versicolor.
590 POLYPORACEAE & ALLIES

Gloeophyllum saepiarium (Rusty Gilled Polypore)

FRUITING BODY annual; shelflike or bracketlike, leathery when fresh, rigid when dry.
CAP 2-12 cm broad, more or less fan-shaped in outline; surface dry, hairy to nearly smooth,
concentrically zoned or ridged and often radially wrinkled, rusty-brown to dark brown or
maroon-brown, often with brighter (yellow, orangish, etc.) zones, but sometimes fading or
weathering to grayish; margin orange, yellow, or whitish when actively growing. Flesh 1-5
mm thick, yellow-brown to rusty-brown. GILLS close, ochre to yellow-brown or rusty-
brown becoming brownish in old age, often fused to form elongated pores or sometimes the
underside of cap entirely poroid or even toothlike. ST ALK absent. SPORE PRINT white;
spores 8-13 * 3-5 microns, cylindrical, smooth. Cap tissue blackening in potassium
hydroxide (KOH).
HABITAT: Solitary or in groups or overlapping tiers on dead conifers (or occasionally
dead hardwoods, particularly aspen); very widely distributed. I have yet to find it in our
area, but it is very common in the Southwest, Pacific Northwest, and the mountains of
California. It causes a rapid carbonizing decay (brown rot) of both the heartwood and
sapwood, and along with G. trabeum{see comments), is a pest of telephone poles, struc¬
tural timber in houses and bridges, etc.
EDIBILITY: Inedible.
COMMENTS: Formerly known as Lenzites saepiaria, this common conifer-lover, like
the hardwood-loving Lenzites betulina, typically has gills instead of pores (see photo on
p. 588). However, the gills are not white and the flesh is rusty-brown—a typical feature of
Gloeophyllum. The gills at times join to form mazelike pockets and sometimes there are
pores instead of gills (leading to confusion with Daedalea and Dae dale op sis), but the rusty
to orange-yellow colors of the cap are distinctive. A related species, G. trabeum, hasagray
to brownish, unzoned cap. It usually grows on dead hardwoods (but occasionally on
conifers), and is more apt to have mazelike pores than gills. It is said to be common on the
woodwork of automobiles! Another unzoned species, G. striatum, occurs only on dead
juniper and cypress. Other species: Osmoporus odoratus is rather similar in color to
G. saepiarium, but usually has pores on the underside of the cap and often has an aniselike
odor when fresh. It is widely distributed and produces a brown rot in dead conifers.

SCHIZOPHYLLUM
SCHIZOPHYLLUM is unique by virtue of its longitudinally split or grooved gills (see
description and photographs). It has few close relatives and is placed in its own family,
the Schizophyllaceae, by most taxonomists. It looks like a polypore, however, and is
treated as one here. One cosmopolitan species is described here.

Schizophyllum commune (Split-Gill)

FRUITING BODY shelflike or with a narrowed base, tough and leathery both fresh and
dry. CAP 1 -4 cm broad, more or less fan-shaped (or vase-shaped if stalk central); surface
dry, densely hairy, white to grayish-white, gray, or sometimes brownish-gray when wet;
margin usually lobed and inrolled in dry weather. Flesh tough, leathery, thin, pallid or
grayish. GILLS radiating from point of attachment, well-spaced, white to grayish; edges
appearing split or grooved lengthwise (i.e., cuplike in cross-section), rolling back in dry
weather. ST ALK absent or present only as a narrowed basal point of attachment. SPORE
PRINT white; spores 3^4 (6) x 1 -1.5 (3) microns, cylindrical, smooth.
HABITAT: Scattered or in groups, rows, or fused clusters on hardwood sticks, stumps,
 Schizophyllum commune. This close-up shows how each gill is actually composed of two adjacent
plates. Note hairs on cap.

logs, etc.; distribution worldwide. It survives dry spells by folding back its gills, and hence
can be found practically year-round. In our area it is common on oak, producing a white
rot (delignifying decay) in its host.
EDIBILITY: Too small and tough to be of value. However, some natives of Madagascar
are said to chew them, for reasons unknown.
COMMENTS: The hairy white to grayish cap of this farflung fungus is reminiscent of the
common bracket polypores in the genus Trichaptum. However, the peculiar manner in
which the gills split lengthwise is unique. The “split” gills are actually two adjacent plates
which separate and roll up in dry weather, thus protecting the spore-bearing surface
(see photographs). Specimens sealed in a tube in 1911, then moistened 50 years later,
unrolled their gills and began shedding spores! Schizophyllum has been widely used in
genetic studies because it fruits readily in the laboratory. Trogia(-Plicaturopsis) crispa of
eastern N orth America is a related mushroom with crisped (wavy) gills and a tough, hairy,
tan to yellowish cap.

Schizophyllum commune. Left: Mature specimens. Cap color ranges from pure white to brownish or
grayish. Right: A close-up of the “split” gills.
Pits
592 POLYPORACEAE & ALLIES

TRAMETES & Allies

Fruiting body small to medium-sized, usually annual but persistent; tough and leathery even when
fresh; growing shelflike or bracket like on wood( usually dead wood), often in masses. CAP often
hairy or velvety and zoned concentrically, usually thin. PORES variously colored, often minuteand
barely visible, round to angular or becoming toothlike or rarely gill-like; tubes usually shallow, one
layer only. STALK absent or rudimentary. SPORE PRINT whitish to yellowish (when obtain¬
able). Spores typically oblong to cylindrical or sausage-shaped, smooth.

GROUPED here are a number of smallish tough or leathery bracket fungi. The fruiting
bodies are much thinner than those of the perennial polypores or “conks,” and are not soft
and spongy when fresh as in Tyromyces. In age the tube walls may break up to form small
“teeth,” but do not normally form the mazelike pattern characteristic of Daedalea and
Daedaleopsis.
The various genera treated here are not closely related and can easily be distinguished
by pore (not spore!) color. The most common genus, Trametes, usually has a zoned cap
and white to dingy buff pores. Trichaptum, also abundant, has violet to brownish pores,
while Bjerkandera and Datronia, with brownish to gray or black pores, and Pycnoporus,
with brilliant red to orange pores, are less frequent.
All five genera feed largely on dead wood—logs, branches, even twigs. They are by
far the most numerous polypores, often completely smothering their substrate in hundreds
of shelving fruiting bodies arranged in attractive clusters or rosettes. Unfortunately, they
are not tender enough to eat. Six widespread species are described here.

Key to Trametes & Allies

1. Pore surface bright saffron to orange-red or red . Pycnoporus cinnabarinus & others, p. 597
1. Not as above (but fertile surface may be rosy, violet, or rusty-reddish) .. 2
2. Fertile surface (or pores) typically rosy, violet, or with a distinct violet tinge when fresh ... 3
2. Fertile surface not rosy or violet-tinged, or rarely with a very faint lavender tinge (and then
growing on poplar or willow) . 4
3. Cap pinkish to reddish or brown when fresh (never white), often fairly thick; pores rosy when
fresh .(see Ganoderma, Fomitopsis, & Allies, p. 574)
3. Not as above; pores tinged violet or lavender when fresh; cap thin and differently colored . 8
4. Fruiting body fleshy (watery or spongy) when fresh (but may toughen in age) and either staining
rusty-reddish when handled or in age or with bluish spores and bluish or bluish-gray-tinged
pore surface .(see Tyromyces Si Allies, p. 597)
4. Not as above . 5
5. Pore surface brown (or gray from a hoary coating), the tubes otten ol unequal length or slotlike
(see photo on p. 596) but not usually forming “teeth”; cap very thin, brown to blackish, velvety
(but not coarsely hairy) when fresh, usually zoned .Datronia mollis, p. 595
5. Not as above (but pores may be brown or grayish) . 6
6. Pore surface(tube walls) often breaking up in age to form numerous small“teeth”; cap typically
hairy . 7
6. Not as above . 9
7. Pore surface usually grayish to smoky-brown or even blackish in age; cap usually zoned con¬
centrically .Cerrena unicolor (see Bjerkandera adusta, p. 596)
7. Pore surface violet-tinged, brown, or occasionally pallid; cap zoned or unzoned . 8
8. Cap commonly up to 7 cm broad; found mainly on hardwoods, especially common in eastern
North America . Trichaptum biformis(see T. abietinus, p. 593)
8. Cap usually less than 4 cm broad, or if larger then usually with gills on underside; found mainly
on conifers; common and widespread .Trichaptum abietinus, p. 593
9. Pore surface soon gray to smoky-brown or black; pores small to minute (3-7 per mm), not
pallid when young or if so then tubes usually separated from flesh by a narrow dark line . . .
.Bjerkandera adusta & others, p. 596
9. Not with above features (but pore surface may become beige or slightly grayish in age) .. 10
TRAMETES & ALLIES 593

10. Cap smooth (bald) or nearly so, or if not then growing on living cypress or juniper and causing
a brown rot .(see Tyromyces & Allies, p. 597)
10. Not as above; cap hairy or velvety; very common, usually on dead wood . 11
11. Cap typically lacking marked concentric zones, grayish to ochraceous to tan or tawny; found
mainly on poplar, willow, or birch; not common .(see Tyromyces & Allies, p. 597)
11. Cap usually zoned and/or differently colored; very common on many trees (including poplar,
willow, and birch) . 12
12. Cap typically 2-7 (10) cm broad, usually with zones of contrasting colors; surface velvety or
with velvety zones alternating with silky-smooth zones . Trametes versicolor, p. 594
12. Cap 2.5-10 cm broad or more, usually hairy throughout, the zones not sharply contrasting
in color .Trametes hirsuta & others, p. 595

Trichaptum abietinus (Violet-Pored Bracket Fungus)

FRUITING BODY shelflike orbracketlike, toughand leathery when fresh, rigid when dry.
CAP 1^4 cm broad, fan-shaped to kidney-shaped or elongated in outline; surface dry,
covered with coarse, stiff hairs; usually concentrically zoned or grooved, white to grayish,
but old weathered specimens often greenish from algae or blackish; margin often wavy.
Flesh very thin (up to 1 mm thick), tough, pale gray to brownish or purplish. PORES 2-4
per mm, round to angular but in age often irregularly torn or toothlike; whitish to brownish
but usually tinged bright lavender to purplish when fresh, especially toward cap margin;
duller or browner in age; tubes very shallow (up to 3 mm long). STALK absent. SPORE
PRINT whitish; spores 4-8 * 2-4 microns, cylindrical or sausage-shaped, smooth.
HABITAT: Ingroups, shelving masses, or overlapping tiers on decaying conifers; widely
distributed and common practically year-round. It is said to be the most important de-
lignifier of coniferous slash in North America. In our area it favors Douglas-fir and pine.

EDIBILITY: Boil for 26 hours, squeeze thoroughly, and serve forth.

COMMENTS: Also known as Hirschioporus abietinus, this little polypore is roughly to
rotting conifers what Trametes versicolor and Lenzites betulina are to rotting hardwoods
—overwhelmingly abundant. The violet-tinged pores (when fresh) that break up into
“teeth” are the best field character. The cap is usually zoned, but not multicolored as in
Trametes versicolor. Fomitopsis cajanderi has rosy pores, but is much thicker and has a
reddish to brown cap. Other species: T. biformis (better known as Hirschioporus par-
gamenus) is a hardwood-loving version of T. abietinus with a cap 2-7 cm broad, violet-
tinted pores that quickly become toothlike, and soft, velvety hairs on the cap. It is one of
the most common polypores of eastern North America (even more abundant than
Trametes versicolor!), often growing in huge masses that completely cover rotting logs,
but is not as prevalent in the West. T. abietinus var. abietis resembles the typical variety but
is usually larger, weathers darker, and has “gills” instead of pores on underside of cap.

Trichaptum (-Hirschioporus) abietinus is abundant on rotting conifers. Note hairy white cap. The
fertile surface is violet-tinged when fresh and features pores, tiny spines, or even gills!
Trametes (- Coriolus) versicolor. This picture shows why it is called the “Turkey Tail.” Multi-colored
zones on the cap are distinctive.

Trametes versicolor Color Plate 158

(Turkey Tail; Many-Colored Polypore)
FRUITING BODY shelflike or bracketlike, thin and leathery when fresh, rigid or slightly
flexible when dry. CAP 2-7 (10) cm broad, tongue-shaped becoming fan-shaped, or
growing in circular rosettes; plane or wavy; surface dry, velvety (covered with fine hairs)
or silky, strongly zoned with narrow, concentric bands of contrasting colors (but more
uniformly colored in sheltered situations), hairy zones usually alternating with silky-
smooth ones; colors extremely variable: a mixture of white, gray, brown, yellowish-buff,
bluish, reddish, or black (or even greenish from a coating of algae), or sometimes dark
brown with a white margin; margin often wavy and white or creamy when actively growing.
Flesh very thin (1-2 mm), tough, white. PORES white to dingy yellowish, minute(3-5 per
mm) but visible; tubes shallow (up to 2 mm long). ST ALK absent or rudimentary. SPORE
PRINT white or yellowish; spores 4-6 * 1.5-2.5 microns, cylindrical or sausage-shaped,
smooth.
HABITAT: Typically in groups, rows, tiers, shelving masses, or overlapping clusters on
logs, stumps, and fallen branches of dead hardwoods (particularly oak), sometimes also
on wounds in living trees and rarely on conifers; very common and widely distributed. It is
abundant in our oak woodlands year-round, but fruits mainly in the winter and spring—it
can be seen on almost any jaunt through the woods. It causes a general delignifying decay of
the sapwood, and along with T. hirsuta, is sometimes parasitic on wounded fruit trees
and lilac bushes.
EDIBILITY: Boil for 62 hours, squeeze thoroughly, and serve forth.
COMMENTS: Our most common polypore, this species is well known—by sight if not by
name—to everyone who spends time in the woods. The multicolored, concentrically zoned
caps do not decay readily, making beautiful ornaments, brooch clips, earrings, and
necklaces. The multiplicity of colors is both its most bewildering and most distinctive
characteristic—no two are colored quite alike. Other Trametes species are less radically
colored and/ or hairier (see T. hirsuta). Stereum species are superficially similar, but
lack pores; Trichaptum species have violet-tinged to brownish pores. Synonyms include
Coriolus versicolor, Polystictus versicolor, and Polyporus versicolor.

594
Trametes hirsuta. This species resembles T. versicolor, but is not as colorful and is often larger.

Trametes hirsuta (Hairy Turkey Tail)

FRUITING BODY shelflike or bracketlike, tough and leathery when fresh, fairly rigid
when dry. CAP 2.5-15 (30) cm broad, fan-shaped to nearly circular in outline; surface
dry, densely and conspicuously hairy to coarsely velvety, usually concentrically zoned or
grooved, but the colors of each zone dull and not sharply contrasting; whitish to grayish,
yellowish, dull ochre, buff, beige, or pale brownish, the margin sometimes darker and often
wavy (but entire surface sometimes greenish from algae). Flesh tough, white (or pale
brownish to yellowish in age), 1-5 mm thick. PORES (1) 2-4 per mm, white to dingy
yellowish or buff when fresh, often tinged brownish or gray in age; tubes 1 -3 (5) mm long.
STALK usually absent. SPORE PRINT whitish or pallid; spores 4.5-7.5 * 1.5-3 microns,
cylindrical to sausage-shaped, smooth.

HABIT AT: S olitary or more often in groups, fused rows, or overlapping clusters on dead
hardwoods (or occasionally conifers); widely distributed throughout the northern hemis¬
phere. It occurs year-round in our area on oak, alder, and other hardwoods, but is rather
uncommon, at least in comparison to T. versicolor. Like the latter species, it causes a
general delignifying decay of the sapwood and sometimes parasitizes fruit trees.
EDIBILITY: Tough and hairy (but see T. versicolor for cooking suggestions).
COMMENTS: This lesser known cousin of T. versicolor is thicker and often larger than
that species, but not as brightly colored. Also, the cap is hairier. T. occidentalis is a very
similar, slightly larger species with a southern and tropical distribution. It is so similar,
in fact, that the two may very well be forms of a single wide-ranging species. For a list of
synonyms for T. occidentalis, see p. 550. Lenzites betulina looks quite similar from the top,
but has gills on the underside of the cap. Other species: T. velutina is very similar, but has a
more velvety cap and its pores are often smoky-tinged; T. pubescens is smaller and has a
white to grayish-yellow, silky-hairy cap with a radially striate margin plus a somewhat
fleshier texture when fresh; both occur on hardwoods and are widely distributed.

Datronia mollis
FRUITING BODY annual; bracketlike or at times resupinate; tough when fresh or dry.
CAP (when present) 1-7 cm broad, shelflike and often wavy; surface umber-brown and
velvety when fresh but becoming smooth and dark brown to blackish in age; usually zoned
concentrically. Flesh thin, tough, pale brown, often separated from the velvety surface

595
Left: Close-up of slotlike pores of Datronia mollis. Right: Dark gray pore surface of Bjerkandera
adusta.

by a thin dark line. PORES 1-2 per mm, often becoming elongated and sinuous (slot¬
like); brown, but covered with a hoary bloom when fresh that gives them a grayish appear¬
ance, the bloom rubbing off easily (i.e., surface bruising brown); tubes 0.5-5 mm long.
STALK absent. SPORE PRINT whitish; spores 8-11 * 2.5-4 microns, cylindrical, smooth.
HABITAT: Usually in groups on dead hardwoods, but also reported on conifers; widely
distributed and fairly common, but often overlooked because of its small size. In our area
I have found it several times in the winter and spring on dead oak. It causes a delignifying
decay of the sapwood.
EDIBILITY: Unknown, but like myself, too small, thin, and tough to bother with.
COMMENTS: Formerly known as Trametes mollis and Daedalea mollis, this small,
nondescript hardwood-lover is best recognized by its thin, tough, velvety cap and grayish
to brown, slotlike pores (see photograph). It is most likely to be confused with Bjerkandera
species, but has larger pores and a thinner cap.

Bjerkandera adusta (Smoky Polyp ore)

FRUITING BODY shelflike or bracketlike, several often fused together; tough or corky
when fresh, rigid when dry. CAP(s) 1.5-7 cm broad, plane or wavy, elongated or fan¬
shaped in outline; surface white to tan, smoky-gray, or grayish-brown, dry, finely hairy
(velvety or suede-like) to nearly smooth; margin whitish when young, darkening or black¬
ening in age. Flesh thin (1 -6 mm), tough, white becoming grayish or brown; odor fungal,
taste often rather sour. PORES very minute (5-7 per mm), scarcely visible, at first whitish
and darkening where bruised, but soon becoming gray or smoky throughout, and finally
blackish; tubes shallow (up to 2 mm long). STALK absent. SPORE PRINT white or
yellowish; spores 4-6 * 2.5-3 microns, elliptical, smooth.
HABITAT: On dead hardwoods (or rarely conifers), usually in dense, overlapping or
fused clusters; widely distributed. It is fairly common in our area throughout the mush¬
room season, but not nearly as abundant as T. versicolor. It produces a general deligni¬
fying decay of the sapwood, giving it a whitish-flecked appearance.
EDIBILITY: Unequivocally inedible.
COMMENTS: The gray to black pore surface is the principal fieldmark of this small,
tough bracket fungus. Gloeoporus adustus is a synonym. B. fumosa is a very similar
species that is somewhat larger and thicker; it has slightly larger, pale gray to smoky-
brown (rarely black) pores, larger spores, and a stronger (aniselike to unpleasant) odor.
596
BJERKANDERA 597

In both species, but particularly B.fumosa, a thin dark line separates the pores from the
flesh. Intermediate forms occur, however, and it is questionable whether the two are
distinct. Cerrena unicolor (see comments under Daedaleopsis confragosa) is somewhat
similar, but its grayish to blackish spore-bearing surface usually breaks up into small
flattened “teeth”; it is widely distributed on hardwoods.

Pycnoporus cinnabarinus (Red Polypore)

FRUITING BODY shelflike, tough when fresh, nearly rigid when dry. CAP 2-12 cm
broad, nearly round to elongated or fan-shaped in outline; surface dry, smooth or finely
hairy, wrinkled or warty, bright orange to orange-red, red, or cinnabar-red, but fading
in age. Flesh red to yellowish-red, tough, up to 1.5 cm thick. PORES 2-4 per mm, bright
orange to orange-red or red, scarcely fading; tubes 1-6 mm long. STALK absent. SPORE
PRINT white; spores 5-6 * 2-2.5 microns, oblong-elliptical, smooth. Cap tissue staining
magenta or black in potassium hydroxide (KOH).
HABITAT: Solitary or in groups on dead hardwoods (particularly cherry, but also oak,
birch, etc.) or occasionally on conifers; mainly northern in distribution. I have not found
it in California, but it occurs in the Southwest and Pacific Northwest and is quite common
in eastern North America. The very similar P. sanguineus (see comments) is prevalent in
the southern United States and tropics.
EDIBILITY: Too tough to be edible, but makes a colorful ornament.
COMMENTS: The beautiful bright red to orange color of the pores makes this one of the
easiest of all polypores to recognize. The cap may fade in age, but the pores will retain their
color for years if stored properly. The fruiting body is not spongy when fresh as in Phaeolus,
and has much smaller pores. P. sanguineus is a closely related brilliant red to orange-red
southern species with a thinner (up to 5 mm) cap. Aurantioporus croceus has saffron to
bright orange pores when fresh; it is also larger (cap 5-30 cm broad) and softer when
fresh, and grows on eastern hardwoods.

TYROMYCES & Allies

Fruiting body typically small to medium-sized (occasionally large) and annual; usually soft, watery,
spongy, or cheesy when fresh, becoming tougher in age; growing shelflike or bracketlike on wood.
CAP usually dull colored (white, gray, etc.), cottony or hairy to smooth, usually not zoned. Flesh
usually white, pallid, or pale brown. PORES minute to fairly large, usually white, pallid, or yellow
when fresh, but sometimes bruising darker. STALK absent or rudimentary or present only as a
narrowed, lateral extension of the cap. SPORE PRINT white or bluish (when obtainable); spores
usually smooth and cylindrical or sausage-shaped.

TYROMYCES is an undistinguished genus of fairly forgettable, fan-shaped to hooflike

fungal fructifications with a fleshy rather than leathery texture when fresh. The latter
feature distinguishes them from Trametes and other tough, bracketlike polypores.
Over 20 species of Tyromyces occur in North America. Most cause brown rots in
dead conifers (in contrast to Trametes, which causes white rots in dead hardwoods).
However, a few common species (e.g., T. chioneus) occur on hardwoods. Despite their
fleshiness they are not good edibles, being infrequent, indigestible, and insipid.

Key to Tyromyces & Allies

1. Spore print bluish or bluish-gray; fruiting body often tinged blue or bluish-gray, especially
at cap margin . Tyromyces caesius& others, p. 599
1. Not as above . 2
 POLYPORACEAE & ALLIES

Fruiting body spongy, watery, or cheesy when fresh and quickly staining rusty-reddish or red¬
dish-brown when handled or in age (especially the pore surface) or staining yellow and then
rusty-reddish; usually found on dead conifers . 3
Not as above (but pore surface may stain pinkish-red if found on dead hardwoods) .4
Pores averaging 3-5 per mm, usually staining yellow before reddening .... T.fragilis, p. 600
Pores averaging 1-3 per mm, reddening directly . . . T. mollis & others (see T. fragilis, p. 600)
Upper layer of cap very soft(marshmallow-like) whenfresh(but may toughen in age); cap often
elongated and narrow, its margin often projecting below the pore surface; found only on dead
mountain conifers (usually above 4,000 ft.) .T. leucospongia, p. 600
Not as above . 5
Tube walls soon breaking up to form spines or “teeth”; cap white to ochre .6
Not as above (but tube walls may sometimes be toothlike or partially toothlike in old age) . 7
Found on conifers; “teeth” 5-10 mm long Spongipellis pachyodon(see T. leucospongia, p. 600)
Found mainly on hardwoods (rarely conifers); “teeth” 1-5 mm long; cap hairy, white, leathery;
flesh very thin .Irpex lacteus
Either pore surface staining dark brown to gray or black when bruised or dried and fruiting body
often large or forming white to creamy rosettes on tropical and subtropical hardwood stumps
and roots or fruiting body with very dense, heavy flesh that dries bone-hard; usually growing
in groups or clusters . (see Polyporus, Albatrellus, & Allies, p. 554)
Not as above . 8
Cap small (up to 4 cm), with a narrowed stemlike base T.floriformis (see T. chioneus, p. 599)
Not as above . 9
Found on incense cedar; fruiting body fairly large (7 cm broad or more when mature) and pore
surface usually yellow or yellowish when fresh . T. amarus, p. 601
Not as above (usually found on a different host) . 10
Found on cypress or juniper (usually living); fruiting body very tough or corky, even when
fresh .T. basilaris(see T. chioneus, p. 599)
Not as above; usually found on a different host . 11
Fruiting body leathery or corky when fresh; cap lacking scales and usually bald or nearly bald,
often concentrically zoned or furrowed near margin; pores varying from mazelike to round,
sometimes staining pinkish-red when bruised; found almost exclusively on hardwoods ....
.(see Daedaleopsis confragosa & others, p. 588)
Not as above . 12
Pores fairly large (usually averaging 1 mm broad or more in widest dimension) . 13
Pores smaller (averaging 1-5 per mm) . 15
Pores hexagonal or angular and usually arranged in radiating rows, or if not then cap ochre-buff
to tan, often with darker scales; rudimentary stalk often present; found on or near hardwoods
.. (seePolyporus, Albatrellus, & Allies, p. 554)
Not as above; cap smooth or hairy but not normally with scales . 14
Cap 4-30 cm broad and 3-8 cm thick, spongy or watery when fresh, the margin usually thick;
found on hardwoods. T. unicolor (see T. leucospongia, p. 600)
Not as above . 15
Cap often poorly developed or rudimentary; tubes often unequal in length; pores large or
small .(seePoria& Allies, p. 602)
Not as above; cap usually well-developed and pores small . 16
Fruiting body tough (corky or leathery) even when fresh; pore surface often grayish to smoky-
brown in age or when dry; found mainly on poplar, willow, or birch . 17
Not as above; fruiting body usually watery, spongy, or cheesy whenyoung( but often toughening
in age); found on a variety of hosts . 18
Odor aniselike when fresh; cap hairy or smooth . Coriolellus (-Trametes) suaveolens
Odor not aniselike; cap hairy . Funalia trogii
Odor fragrant when fresh and taste not bitter; usually found on hardwoods T. chioneus, p. 599
Odor not fragrant and/ or taste bitter; found on hardwoods or conifers . 19
TYROMYCES & ALLIES 599

19. Taste bitter; usually found on conifers .. T. guttulatus & T. stipticus (see T. chioneus, below)
19. Taste not bitter; on hardwoods or conifers T. tephroleucus & others (see T. chioneus, below)

Tyromyces caesius (Blue Cheese Polypore)

FRUITING BODY shelflike or bracketlike, soft and spongy or watery when fresh,
tougher when dry. CAP 1-5 (8) cm broad, fan-shaped to semi-circular in outline; surface
white to gray, but usually tinged or mottled bluish or blue-gray, especially toward the
margin; covered with soft, whitish hairs; not zoned. Flesh up to 1 cm thick, white or aging
gray to yellowish, spongy when fresh; odor often sweetish. PORES 2~4 per mm, white or
colored like cap; tubes 2-8 mm long. STALK absent. SPORE PRINT pale ashy-blue;
spores 3-6 * 1-2 microns, cylindrical or sausage-shaped, smooth.
HABIT AT: S olitary or in small groups on decaying wood of both hardwoods and conifers;
widely distributed and fairly common but rarely occurring in large numbers. In our area
it can be found practically year-round. It is associated with a brown carbonizing decay.
EDIBILITY: Unknown.
COMMENTS: The delicate bluish-gray tint to the cap and soft texture when fresh
distinguish this unobtrusive polypore. Other species: T. perdelicatus also has pale bluish
spores, but the cap is smaller (up to 2 cm broad) and not normally bluish, and the taste is
bitter. It is fairly common on dead conifers in the Pacific Northwest.

Tyromyces chioneus (White Cheese Polypore)

FRUITIN G BODY shelflike or bracketlike; fleshy and rather soft or spongy-watery when
fresh, rigid and tough when dry. CAP 2-12 cm broad, fan-shaped to semi-circular, broadly
convex to plane; surface smooth or slightly hairy, pure white to buff, yellowish-buff, or
watery gray, not zoned. Flesh 2-15 mm thick, white; soft and spongy with a fragrant odor
when fresh, crumbly when old and dry. PORES 3-5 per mm, white to creamy oryellowish;
tubes 1.5-3 (7) mm long. STALK absent. SPORE PRINT white; spores 3.5-5 * 1-2
microns, cylindrical to sausage-shaped, smooth.
HABITAT: Solitary or in groups on dead hardwoods or occasionally on dead conifers;
widely distributed and common, but rarely fruiting in large numbers. In our area it is fairly
frequent in the fall and winter, especially on oak. It produces a wet, stringy delignifying
decay of the sapwood.
EDIBILITY: Unknown.
COMMENTS: The pale color, spongy or cheesy texture when fresh, modest size,
fragrant odor, and absence of any staining reactions are the principal characteristics of this
polypore, which is better known as T. albellus. There are a number of more or less similar
white or pallid Tyromyces species that are best differentiated microscopically, including: T.
tephroleucus (-T. lacteus), with a mild odor, on both hardwoods and conifers; T. galac-
tinus, common on hardwoods in eastern North America, with a hairier cap and elliptical
spores; T. balsameus, small, thin, with a faintly zoned cap, on conifers; T. spraguei, with a
cap margin that stains greenish or blackish when young and nearly round to elliptical
spores, on hardwoods; T. guttulatus, often larger (cap up to 16 cm broad), with circular
spots, pits, or droplets on the cap, growing mainly on conifers; T. stipticus (-T. immitis),
even more bitter but lacking spots or pits, on conifers; T. floriformis, usually bitter but
small(cap up to4 cm broad) and petal-like(i.e., the cap has a narrowed, stemlike base), on
conifers; and T. basilaris, tough, attacking living juniper and cypress trees (including
Monterey cypress). T. spumeus should also be mentioned. In contrast to T. chioneus,
nearly all of the above species produce brown rots (carbonizing decays) in their hosts.
Tyromyces fragilis, a nondescript polypore that stains rusty-yellow to reddish when handled.

Tyromyces fragilis (Rusty-Staining Cheese Polypore)

FRUITING BODY shelllike or bracketlike, soft and spongy or watery when fresh; rigid
and brittle when dry. CAP 2-10 cm broad, fan-shaped to elongated in outline; surface
covered with soft white hairs that become matted in age; white, but becoming reddish to
pinkish-red in age; staining yellowish, then rusty to reddish when handled. Flesh soft when
fresh, white, discoloring like the cap surface. PORES 3-5 per mm, white, quickly bruising
yellowish, then rusty-reddish; tubes 2-8 mm long. STALK absent. SPORE PRINT
whitish; spores 4-5 * 1-2 microns, cylindrical to sausage-shaped, smooth.
HABIT AT: S olitary or in groups or fused clusters on rotting conifers; widely distributed.
In our area it is fairly common (along with T. mollis—see comments) in the fall and winter
on pine and Douglas-fir. It produces a brown carbonizing decay in its host.
EDIBILITY: Unknown, and like myself, likely to remain so.
COMMENTS: The tendency of all parts, especially the pores, to stain rusty-red when
handled is the one noteworthy feature of this otherwise unnoteworthy polypore. There
are several very similar reddish-staining species with slightly larger pores (1 -3 per mm) and
thicker flesh (0.5-2 cm) that are best differentiated microscopically, including: T. mollis,
also common in our area, with a larger, smoother cap that stains rusty-red directly; T.
transmutans, with dextrinoid spores; and A mylocystis lapponica, with amyloid cystidia
and spores 7-11 microns long.

Tyromyces leucospongia (Marshmallow Polypore)

FRUITIN G BODY annual, shelflike or bracketlike to somewhat irregular; soft and rather
watery when fresh, but soon becoming tough and rigid except for the soft upper surface.
CAP usually elongated (2-10 cm long and 1-5 cm wide), convex; surface white to buff,
grayish-buff, pinkish, or even brown or cinnamon; usually velvety and very soft or spongy
to the touch, but sometimes papery and fairly firm if the velvety layer wears away; margin
usually extending below the pore surface and sometimes partially covering it. Flesh white
and rather tough, but covered by a thick (up to 1.5 cm), soft, spongy layer of cottony tissue
that is pallid to pinkish-buff or cinnamon. PORES 2-3 per mm, angular and often
becoming torn and markedly toothlike in age( or interspersed with spines or“ teeth”); white
when fresh, often discoloring (buff to tan, brownish, or cinnamon) in age or when dried;
tubes2-6 mm long. STALK absent or rudimentary. SPORE PRINT white; spores4-6 * 1-
1.5 microns, more or less sausage-shaped, smooth.
HABITAT: Solitary to gregarious on dead conifers, usually fruiting from fissures in
the bark; common at higher elevations throughout most of western North America. It
usually fruits in the spring (often beginning its development under the snow), but the
fruiting bodies persist for months without decaying, and can thus be found most anytime.
It causes a brown (carbonizing) rot of both the heartwood and sapwood of its host.
EDIBILITY: Not edible.

600
Tyromyces (-Spongiporus) leucospongia has a unique cottony or marshmallow-like texture. Note
how the margin of the cap extends over the pore surface.

COMMENTS: Also known as Spongiporus (or Spongipellis) leucospongia, this is one of

my very favorite polypores because of the unique texture of its skin. The texture is difficult
to describe, but it is undeniably soft—something like cotton candy (but not sticky) or a
chamois-covered cotton ball, and also reminiscent of a “Nerf ball.” In age it may toughen
and is not so readily identified, but the tendency of the margin to grow over the pore surface
is also distinctive. Other species: Spongipellis pachydon (-Irpex mollis) is a widely
distributed species that grows on conifers and has pores which soon become toothlike; it is
tough, has a spongy skin only when very young, and lacks the extended cap margin of T.
leucospongia. T. (-Spongipellis) unicolor is also widespread and spongy when fresh, but
has larger pores (1 mm or more broad), nearly round spores, a thicker (3-8 cm) and larger
cap, and grows on living or dead hardwoods (usually oak).

Tyromyces amarus (Incense Cedar Polyp ore)

FRUITING BODY more or less hoof-shaped, annual; soft and watery when very young,
hard and rigid in age or when dry. CAP 7-30 cm broad and 4-20 cm thick, convex; surface
downy when young, usually bald in age but often becoming roughened, wrinkled, or
shallowly fissured; whitish or buff to pale brown, sometimes darkening or becoming
more ochraceous as it dries; surface usually quite hard (or even crustlike) in age; margin
thick and blunt. Flesh thick, soft when fresh but very hard when dry; yellowish to pale
brown. PORES 2-3 per mm (but may fuse to form larger pores), yellowish to bright lemon-
yellow when fresh, often discoloring with age or handling; tubes 5-15 mm or more long.
STALK absent. SPORE PRINT white; spores 6-7.5 * 3.5-5 microns, elliptical, smooth.
HABITAT: Solitary or occasionally several together on incense cedar (Libocedrusf,
known only from western North America, fruiting mainly in the late summer, fall, and
early winter (but persisting year-round) wherever incense cedar occurs. It causes a de¬
structive carbonizing trunk rot that creates large tunnels in the wood. One third of all
logged incense cedar is infected by it, and must either be rejected or used as “pecky cedar”
for fencing and other purposes.
EDIBILITY: Unknown, but too tough to be worthwhile.
COMMENTS: This interesting polypore is restricted to incense cedar (although there
is one report of it from Idaho, on fir). It might be mistaken for a conk (Fomitopsis or Phel-
linus) because of its large size and tough texture in age, but the fruiting body is usually
annual and quite watery when actively growing. The yellow pore surface and growth on
incense cedar are the principal fieldmarks.

601
602 POLYPORACEAE & ALLIES

PORIA & Allies

Fruiting body usually annual but sometimes perennial, resupinate (lying flat on the substrate);
soft, tough, or chalky; growing on wood. CAP absent or rudimentary. PORES fairly large to
minute; tube layer always present. STALK absent. SPORE PRINT white or yellowish. Spores
elliptical to cylindrical, smooth.

THIS large group of polypores is distinguished by its resupinate fruiting body: a simple
layer of tubes devoid of cap and stem. In several other genera the fruiting body is some¬
times resupinate—especially when growing on the undersides of logs—but in Poria it is
nearly always resupinate.
Because they lack both a cap and stem, Porias are among the most lackluster of fleshy
fungi—they just lie there on their logs, listless and limpetlike, unobtrusively going about
their business while boletivores and polypores blatantly go about theirs. They usually
grow on dead wood and play an important role in the forest ecosystem. A few are pests of
structural timber in mines, cellars, and other dank places where the dismal, abysmal
conditions are to their liking.
More than 150 species of Poria have been described from North America, but identi¬
fication is a difficult, tedious, and time-consuming task. Like Polyporus and Pomes,
Poria has been split into several smaller genera on the basis of microscopic and chemical
characters beyond the scope of this book. Two representative species are described here,
and a few others are keyed out plus several resupinate species that belong to other genera.
They represent but a fraction of the total number, however, and serious students will want
to consult the technical literature on the subject. If the fruiting body is tough and possibly
perennial, check the key to the conks (Ganoderma, Fomitopsis, Phellinus, & Allies).
Key to Poria & Allies
1. Spore-bearing surface composed of meandering veins that form shallow pits .
. (see Stereaceae & Allies, p. 604)
1. Spore-bearing surface composed of a layer of tubes (pores) or sometimes “teeth” . 2
2. Spore-bearing surface soon breaking up into small “teeth” and/ or violet-tinged when fresh 3
2. Spore-bearing surface not normally violet-tinged or composed of “teeth” .4
3. Spore-bearing surface violet-tinged when fresh . (see Trametes & Allies, p. 592)
3. Spore-bearing surface white or pallid .(see Tyromyces & Allies, p. 597)
4. Pore surface and/or flesh pink to orange, salmon, or reddish . 5
4. Not as above; pore surface white to yellow or some shade of brown or gray . 7
5. Pores large (typically at least 1 mm broad); found on conifers (see Phaeolus alboluteus, p. 571)
5. Pores minute (3-7 per mm); found on hardwoods or conifers . 6
6. Pore surface dark reddish-orange; fruiting body cheesy when fresh; pores scarcely visible
(averaging 7-9 per mm); found mainly on hardwoods .P. spissa
6. Not as above; pore surface pink to salmon or brick-red .Chaetoporus euporus & others
7. Flesh yellow-brown to rusty-brown, dark brown, etc. 8
7. Flesh white to creamy, straw-colored, yellowish, or beige (at least when fresh) . 10
8. Found on charred conifers; pores fairly large (1 -2 per mm) and usually hexagonal; fruiting body
not normally perennial . Coriolellus carbonarius (see Poria corticola, p. 603)
8. Not as above . 9
9. Fruiting body either dark brown to black, hard, often cracked and cankerlike, and usually found
on birch or fruiting body growing in sheets under the bark of living oak .
.(see Phaeolus, Inonotus, Coltricia, & Allies, p. 566)
9. Not as above; fruiting body usually perennial, quite tough or woody .
.(see Ganoderma, Fomitopsis, Phellinus, & Allies, p. 574)
10. Found on living cypress or juniper (causing a brown rot) . . . (see Tyromyces & Allies, p. 597)
10. Not as above . 11
A member of the Poria corticola group. Note the absence of both cap and stem—most Porias consist
solely of a layer of tubes.

11. Fruiting body hard or woody even when fresh, the margin often incrusted; usually perennial
.(see Ganoderma, Fomitopsis, Phellinus, & Allies, p. 574)
11. Not as above . 12
12. Rudimentary cap often present; tubes often unequal in length . 13
12. Not as above . 14
13. Pores large (1-3 mm in diameter) .... Coriolellus heteromorpha (see Poria corticola, below)
13. Pores smaller .Coriolellus sepium & others (see Poria corticola, below)
14. Pore surface bright yellow; found on dead conifers.P. xantha(see P. corticola, below)
14. Not as above (pore surface may be yellowish, but not bright yellow) . 15
15. Fruiting body sometimes growing from an underground “tuber”; uncommon P. cocos, p. 604
15. Not as above; very common . 16
16. Pores averaging 4-6 per mm; fruiting body often perennial (with more than one tube layer)
. Perenniporia subacida (see Poria corticola, below)
16. Pores larger than above and/ or fruiting body not perennial P. corticola & many others, below

Poria corticola (Boring Poria)

FRUITING BODY usually annual, rather tough; resupinate(consisting of a simple layer
of tubes). CAP absent or sometimes present as a free margin. Flesh thin, white, fibrous but
rather soft when fresh. PORES white or creamy, discoloring to pale tan in age or upon
drying, 1-4 permm; tubes3-10 mmlong, rather soft when fresh but drying rigid and tough.
STALK absent. SPORE PRINT white; spores 5-8 * 3-5 microns, broadly elliptical,
smooth. Cystidia present among the basidia.
HABIT AT: S olitary or more often in rows or fused masses on rotting hardwood logs and
branches, less commonly on conifers; widely distributed and fairly common in our area
practically year-round. It produces a white rot; many similar species occur on conifers.
EDIBILITY: The entire group is worthless from an edibility standpoint.
COMMENTS: This species typifies a large number of white to yellowish Porias that can
only be differentiated microscopically. Another common species, Perenniporia subacida,
is often perennial (i.e., with more than one tube layer). It is usually creamy or yellowish,
has minute pores (4-6 per mm), occurs on both hardwoods and conifers, and is particularly
abundant in the Pacific Northwest. There are several similar species that cause brown rots
in structural timber as well as in dead conifers and hardwoods, including. P. vaillantii,
white to yellowish;/>. incrassata, white to yellowish butdryinggrayish-brownto black and
with yellow spores; P. xantha, yellow or yellowish-white with a crumbly or chalky
texture, bitter taste, and minute pores; and P. sequoiae, which infects the butts of living
redwoods. Several Coriolellus species should also be mentioned. They were formerly
placed in Trametes because they often possess a cap, but can also be resupinate or have
only a rudimentary cap. Coriolellus serialis is a corky or woody, white to creamy,
resupinate species that is common on dead conifers and hardwoods; C. carbonarius has
brownish pores and flesh and grows on charred conifers, including redwood; C. hetero-
morphus is whitish and has a rudimentary cap or free margin and large pores( 1 -3 mm each
in diameter); C. sepium also has a rudimentary cap but its pores are smaller (1 -2 per mm);
and C. alaskanus is a resupinate species with very thin (less than 1 mm) flesh.

603
604 POLYPORACEAE & ALLIES

Poria cocos (Tuckahoe)

FRUITING BODY annual, rather tough; resupinate (consisting of a simple tube layer),
but sometimes arising from a buried sclerotium (“tuber”). CAP absent or rudimentary.
Flesh thin, ivory-whitish to tan. PORES yellowish-white to tan or pinkish-buff, 1-3 per
mm. STALK absent. SCLEROTIUM when present often large, somewhat resembling an
oblong coconut; outer surface brown and scaly, interior white and cheesy when fresh.
SPORE PRINT white; spores 7-11 * 3A microns, cylindrical, smooth. Cystidia absent.
HABITAT: Occasionally growing on conifer logs, but more often infecting tree roots and
stumps (of both hardwoods and conifers), and the fruiting bodies appearing terrestrial
if they arise from sclerotia; widely distributed, but most common in southeastern North
America. It is reported from California, but I have not seen it in our area. It causes a brown
cubical root and butt rot in its host.
EDIBILITY: The fruiting bodies, like those of other Porias, are worthless, but the large
underground “tuber” was apparently eaten by various tribes of Native Americans (some
of whom called it “tuckahoe”). The “tubers” are said to be visually unappetizing, but I’ve
seen “Unnative” Americans foraging for even stranger fare in supermarkets—Spam, for
instance, or pickled pig’s feet.
COMMENTS: The ability to form large, coconut-sized sclerotia is the only redeeming
feature of this otherwise boring Poria.

Crust and Parchment Fungi spores

STEREACEAE & Allies

Fruiting body soft and spongy to thin and tough; resupinate, bracketlike, shelflike, or occasionally
stalked; usually growing on wood. CAP present or absent, when present usually small. SPORE-
PRODUCING SURFACE smooth to wrinkled, veined, or warty. STALK usually absent but
occasionally present. SPORE PRINT white to yellow or brown, but often not obtainable. Spores
variously shaped.

THIS is a very difficult and immense group of very simple, less-than-immense fungi with a
smooth to wrinkled, veined, or warty spore-bearing surface. They can be found almost
anywhere, at almost any time, and play an integral role in the breakdown of wood and
plant material. Some forms, such as Stereum, mimic polypores by fruiting in shelving
masses on wood, but the majority (Corticium, Peniophora, etc.) are resupinate, i.e., they
form crust- or parchment-like sheets on logs or stumps. Others form small, tile-like
plaques (Xylobolus), “whitewash” sticks and branches, or cause dark discolorations on the
bark of trees. Still others cause serious damage to structural timber (e.g., rickety bridges
and rotting rafters), and a few, such as Cotylidia, are stalked or branched and/ or grow
on the ground. Whatever their shape and growth habit, however, they all differ
fundamentally from the polypores by virtue of their relatively unspecialized spore-
producing surface—in other words, they lack a layer of tubes and pores.
Most of the crust and parchment fungi were originally classed together in a single
family, the Thelephoraceae. On the basis of various chemical and microscopic features
several families are now recognized: the Stereaceae, Corticiaceae, Coniophoraceae,
Hymenochaetaceae, etc. As relatively few are conspicuous or fleshy enough to warrant
attention from the average mushroom hunter and even fewer can be accurately identified
without painstaking microscopic examination and still fewer are edible, only a very very
few are described here.
STEREACEAE & ALLIES 605

Key to the Stereaceae & Allies

1. Growing in white to creamy rosettes of flattened segments or lobes on hardwood stumps and
roots in tropics and along Gulf Coast .Hydnopolyporuspalmatus
1. Not as above . 2
2. Fruiting body with a cap and stalk, the stalk central to somewhat off-center; usually growing on
ground or on very rotten wood . 3
2. Not as above .4
3. Fruiting body chocolate- to purple-brown or dark brown Thelephora terrestrisgroup, p. 608
3. Fruiting body paler (hazel to buff, whitish, or yellowish) Cotylidia diaphana & others, p. 608
4. Fruiting body hard, black, usually cracked, somewhat irregular in shape or cankerlike; growing
on hardwoods, especially birch .(see Phaeolus, Inonotus, Coltricia, & Allies, p. 566)
4. Not as above . 5
5. Fruiting body bright to dark blue or purple or with a purple tinge when fresh; on hardwoods 6
5. Not as above . 7
6. Fruiting body bright to dark blue, velvety .Pulcherricium caeruleum
6. Fruiting body purple or purple-tinged, at least when fresh.
.Chondrostereumpurpureum (see Stereum hirsutum group, below)
7. Spore-bearing surface smooth to slightly uneven or sometimes cracked . 8
7. Spore-bearing surface conspicuously warty, lumpy, furrowed, ridged, wrinkled, pitted, or
honeycombed with large “pores” . 13
8. Fruiting body terrestrial and branched (coral-like), usually dark brown or purple-brown ....
.Thelephora palmata, p. 609
8. Fruiting body usually found on wood, or if on ground then usually with a cap (upper surface) 9
9. Fruiting body purple- to chocolate-brown or dark grayish-brown, usually growing on herba¬
ceous stems or in rosettes or clusters on the ground; spore print brown; spores warty or angular
under the microscope .Thelephora terrestris group, p. 608
9. Not as above . 10
10. Fruiting body usually “bleeding” when cut or handled (exuding a red juice); found on conifers
. Stereum sanguinolentum (see S. hirsutum group, below)
10. Not as above; found on hardwoods and/or fruiting body not “bleeding”. 11
11. Cap silky-striate, thin; found on hornbeam in eastern North America Stereum striatum, p. 607
11. Not as above . 12
12. Cap very thin, pliant, typically with long, loosely-arranged white hairs pointing toward the
margin .Stereum striatum, p. 607
12. Not as above (but cap usually with fine velvety hairs) Stereum hirsutum group &l others, below
13. Growing on structural timber (in houses, mines, etc.) . 14
13. Growing in the wild (on logs, stumps, etc.) . 15
14. Spore-bearing surface^honeycombed with large pits .Serpula lacrymans, p. 610
14. Spore-bearing surface wrinkled or bumpy Coniophoraputeana(see Serpula lacrymans, p.610)
15. Spore-bearing surface irregularly warty or with radiating or concentric wrinkles or furrows;
cap absent .Phlebia radiata & others, p. 610
15. Not as above; spore-bearing surface usually with a honeycombed or veined appearance; cap
or free margin sometimes present . 16
16. Cap or free margin hairy, white . Merulius tremellosus (see Serpula lacrymans, p. 610)
16. Cap or free margin bright pink to reddish Merulius incarnatus (see Serpula lacrymans, p. 610)

Stereum hirsutum group (False Turkey Tail; Hairy Stereum)

FRUITING BODY thin, leathery, pliant when moist, rigid when dry, annual but per¬
sistent; bracketlike to shelflike or partially resupinate with a free margin. CAP 0.5-4 (5)
cm broad but sometimes fused laterally to form larger, lobed shelves 10 cm long or more;
plane to folded or wavy (crisped); surface dry, often zoned concentrically, with whitish to
brownish or grayish matted hairs (but often smooth toward the margin), the hairs wearing
Stereum hirsutum group. Noted the zoned cap which is finely hairy (under a hand lens) when fresh
but often smooth in old age. The underside is golden to orange or buff when fresh (specimen at top
right), but often darkens in old age (specimen at top center).

away in narrow zones to reveal the reddish-brown to dark chestnut-brown cap cuticle;
margin often orange to golden or tawny, especially when young or growing; overall color
thus variable: orange-brown to reddish-brown to cinnamon when moist, but appearing
buff to grayish or paler from the hairs when dry; in old age sometimes greenish from algae
or even blackish. Flesh thin, tough. UNDERSIDE smooth to slightly bumpy or cracked
(when dry), sometimes exuding a red or yellow liquid when cut (fresh); color variable:
orange to dull orange-buff or tawny to ochraceous, varying to buff or pinkish-buff, some¬
times zoned concentrically, often browner or darker toward the base; in old age often dark
brown to chestnut-brown. STALK absent or present only as a narrowed lateral base.
SPORE PRINT white; spores 5-8 * 2-3.5 microns, cylindrical, smooth.
HABITAT: In groups, rows, fused masses, or dense overlapping clusters on hardwood
sticks, fallen branches, logs, stumps, etc., occasionally on living trees or conifers; widely
distributed and extremely common. Found in our area year-round, especially on dead oak.
Trametes, Lenzites, and/or Tremella species often co-inhabit the same piece of wood.
EDIBILITY: Like myself, too thin and too tough to be edible.
COMMENTS: This omnipresent little bracket fungus can be mistaken for the turkey tail
(Trametes versicolor), but a close inspection of the spore-bearing surface reveals the
complete absence of tubes or pores. The above description is rather lengthy because it
encompasses a number of confusing forms that are sometimes recognized as distinct
species. For instance, the common North American variety with a bright orange to
orange-buff spore-bearing surface has been called S. complicatum and S. rameale.
Another variety, S. gausapatum, “bleeds” red when cut, others exude a yellow juice,
while typical S. hirsutum does not bleed at all. However, these “species” grade into each
other, making a neat, clean separation almost impossible. As a group they can be
recognized easily by their omnipresence (in the hardwood forests of California they
outnumber even Trametes versicolor), the reddish-brown cap cuticle (best seen by sec¬
tioning the fruiting body) beneath the hairs, the frequent orange to yellowish tones on
the cap margin or underside, and the smooth spore-bearing surface.
Other species: S. fasciatum (-S. ostrea, S. lobatum) differs microscopically, but can
usually be told by its slightly larger caps (1 -7 cm broad) that are more prominently zoned
(dark reddish and brown) and usually form individual brackets rather than fusing, and
buff to cinnamon-buff underside; it, too, is common on hardwoods, especially oak.
S. (- Haemalostereum) sangunolentum is one of the few widespread Stereums to grow on
conifers. When fresh its fertile surface “bleeds” dramatically when cut, sometimes even
staining one’s hand. Hymenochaete tabacina resembles the S. hirsutum group in shape
and color, but its tissue blackens in potassium hydroxide (KOH). H. rubiginosa also
blackens in KOH but has a velvety, rusty-brown to chestnut-brown to blackish cap and
rusty-brown underside. Both of these favor oak. Chondrostereum purpureum is purple
when fresh and often resupinate; it parasitizes apple and plum trees, causing “silver leaf’

606
STEREUM 607

disease, but is also frequent on other trees (including oak in our area). Laxitextum bicolor,
found on hardwoods, has a brown cap and white to pale buff fertile surface. Peniophora
gigantea forms a paper-thin crust on dead conifers. Veluticeps berkeleyi has a minutely
bristly fertile surface and grows on both hardwoods and conifers, but favors ponderosa
pine. Dozens of other species also occur.

Stereum striatum
FRUITING BODY very thin, leathery and pliant when fresh, annual but persistent;
bracketlike to cuplike. CAP 0.3-3 (4) cm broad but sometimes fused laterally to form lines
10 cm long or more; flat and circular to fan-shaped in outline or if small, often conical
(inverted cup-shaped); surface dry, whitish to buff or pale brown, sometimes zoned con¬
centrically when moist; covered with long, loosely-arranged white hairs which usually
point toward the margin (var. ochraceoflavum) or the hairs pressed against the cap to give
it a silky-shiny striate appearance (var. striatum). Flesh very thin, tough. UNDERSIDE
(fertile surface) smooth, buff to pale brown or in some forms yellow to yellow-brown,
sometimes fading in age to whitish, sometimes zoned concentrically. STALK absent or
present only as a small knob or “umbo” on top of the cap. SPORE PRINT whitish; spores
5-8.5 * 2-3.5 microns, cylindrical, smooth.
HABITAT: In groups or masses on dead branches and twigs (rarely logs) of hardwoods;
widely distributed. Variety ochraceoflavum can be found year-round in our oak wood¬
lands, but is shrivelled up and inconspicuous in dry weather. Variety striatum occurs in
eastern North America on hornbeam (Carpinus).
EDIBILITY: Not edible.
COMMENTS: Like myself, this species is too tough, too thin, and too small to be edible.
Unlike myself, it contents itself with unambitious undertakings—decomposing branches,
sticks, and lopped-off limbs—while leaving the larger stumps, logs, and standing trees
to the polypores and other bracket fungi. Variety ochraceoflavum (-S. ochraceoflavum)
can be distinguished from the more common S. hirsutum group by its duller, paler color
and thinner, more pliant cap with long white hairs pointing toward the margin, plus the
absence of a red-brown cap cuticle. V ariety striatum(-S. sericeum) is easily told by its silky-
striate cap. At least two growth forms of var. ochraceoflavum occur in our area: a small
one with a concave spore-bearing surface and a somewhat conical cap less than 1 cm
broad that is usually attached by its top to small twigs; and a larger one with a flatter cap
that usually inhabits larger sticks and branches and is often partially resupinate.

Stereum striatum is common on sticks and branches of various hardwoods. Note the long hairs that
protrude from cap.
Left: Top view of Cotylidia diaphana (this specimen has several caps). Center: Underside of Cotylidia
diaphana. Right: Thelephora terrestris group. Note how underside is darker than that of Cotylidia.

Cotylidia diaphana (Stalked Stereum)

FRUITING BODY annual, erect, thin and tough, with a cap and stalk. CAP 0.5-3 cm
broad, vase-shaped or funnel-shaped or often split into petal-like lobes; surface dry, with
fine radiating silky fibrils, whitish to buff to pale hazel-brown, sometimes with obscure
concentric zones. UNDERSIDE (fertile surface) smooth or somewhat uneven but without
pores, whitish to buff, pinkish-buff, or tinged cap color. STALK 0.5-3.5 cm long, 1-2 mm
thick, more or less central, solid, smooth, colored like rest of fruiting body; base usually
with white mycelial down. SPORE PRINT whitish; spores 4-6 (8) * 2.5-4 microns, ellip¬
tical, smooth. Long, narrow, projecting cystidia present among the basidia.
HABITAT: Solitary or in groups among humus and debris in woods; widely distributed.
Occasional in our area in the fall and winter, but easily overlooked.
EDIBILITY: A worthless, miniscule morsel.
COMMENTS: Formerly known as Stereum diaphanum, this species differs from Stereum
by its well-developed, more or less central stalk, and from Thelephora terrestris by its
paler color and whitish spores. It is considered by some to be a variety of C. aurantiaca,
a widely distributed, often larger and yellower species. Other species: C. decolorans
(-Stereum burtianum) of eastern North America is similar but lacks cystidia.

Thelephora terrestris group (Earth Fan; Fiber Vase)

FRUITING BODY annual but persistent, tough; usually vase-shaped to fanlike, often
clustered or in confluent masses, sometimes bracketlike or shelflike on plant stems. CAP
2-5 cm broad or forming rosettes or clusters up to 12 cm broad; surface dry, with radiating
silky fibrils or small scales; brown to reddish- or chocolate-brown to grayish-brown or
fuscous, often darker in age; margin usually fringed, splitting, often paler or whitish. Flesh
very thin, tough; odor mild or earthy. UNDERSIDE smooth or wrinkled, without pores;
some shade of brown. STALK when present lateral to central, thin and tough, colored like
cap or paler, short. SPORE PRINT purplish-brown; spores 8-12 * 6-9 microns,
elliptical-angular, warted (often minutely so).
HABIT AT: Occasionally solitary but more often in groups or clusters in humus, sandy soil,
and decomposing vegetable matter; sometimes on old stumps or climbing up herbaceous
stems or tree seedlings; widespread and common. In our area it grows year-round, but is
easily overlooked. I’ve seen it several times on potted plants in nurseries.

608
Left: Thelephora terrestris group, top view of a compound fruiting body. Right: At top is Thelephora
pa/mata—note its flattened branches. At bottom are small specimens of the Thelephora terrestris
group.

EDIBILITY: I can find no information on it.

COMMENTS: The size and shape of this species is fairly variable but the color is quite
constant (quite variable? fairly constant?). When growing on the ground in small, erect
clusters it might be mistaken for an emaciated Craterellus, but is smaller and differently
colored. When growing on herbaceous stems, on the other hand, it looks more like a
Stereum. T. laciniata is apparently a synonym. Other species: T. multipartita is a small
(1-3 cm high), widespread, terrestrial species with a vase-shaped cap that splits into several
lobes or “branches”; T. vialis of eastern North America is a terrestrial species with a larger
fruiting body and a fetid odor in age, and smaller spores; T. spiculosa encrusts twigs and
stems, but has spiky protuberances and a whitish growing margin; Sebacina incrustansalso
grows on herbaceous stems (usually at their bases) but is paler in color and has white spores.
None of these are worth eating.

Thelephorapalmata (Fetid False Coral)

FRUITING BODY annual, erect, usually profusely branched from a common base;
2-10 cm high and just as broad or broader. BRANCHES purplish-brown to chocolate-
brown or darker, flattened; tips also flattened (palmlike) and usually paler (whitish)
when actively growing. STALK present only as a common base or short “trunk” below
the branches. Flesh tough, leathery; odor garliclike becoming fetid (unpleasant) in age.
SPORE PRINT dark reddish-brown; spores 8-11 * 7-8 microns, elliptical-angular, spiny.
HABITAT: Solitary or in groups on moist ground in woods and at their edges; widely
distributed. It often grows along woodland paths but blends uncannily into its sur¬
roundings. In our area it fruits in the late fall, winter, and spring, but is not particularly
common.
EDIBILITY: Unknown
COMMENTS: This mushroom looks like a coral fungus and is keyed out under that
group. However, the flattened branches, dark color, fetid garlic odor, and angular-
elliptical spores are distinctive. Other species: T. vialis of eastern North America is more
variable in color and has smaller spores.

609
Left: Radulum orbiculare (see comments below) forms sheets of irregularly warted or lumpy tissue.
Right: The veined underside of Merulius tremellosus (see comments under Serpula lacrymans).

Phlebia radiata
FRUITING BODY annual, resupinate (lying flat on substrate), sometimes with a free
margin but no cap or stalk; soft when fresh, tough in age. FERTILE SURFACE usually
fused to form patches 30 cm or more long, but often with smaller, discrete systems of
radiating wrinkles or warty veins 1 -4 cm broad; flesh-colored to bright orange to pinkish-
red, fading to whitish in old age. U nderside of margin (if free) with white woolly hairs. Flesh
thin, rather soft or slightly gelatinous when fresh, tough in age or when dry. SPORE
PRINT whitish; spores 3.5-7 * 1-3 microns, sausage-shaped or elliptical, smooth.
HABITAT: On fallen logs and branches of both hardwoods and conifers; widely dis¬
tributed. I find it occasionally on oak logs in the fall and winter.
EDIBILITY: Inedible. It looks as if it has already been eaten (see comments).
COMMENTS: Also known as P. merismoides, this species is unique by virtue of its
resupinate, orange to pinkish fruiting body with radiating wrinkles. Its overall appearance
is somewhat reminiscent of regurgitated dog food, and a similar species, Radulum
orbiculare, has an irregularly lumpy or warty spore-producing surface that is distinctly
reminiscent of regurgitated dog food. Other species: Punclularia strigoso-zonata is also
similar, but has concentric wrinkles and furrows instead of radiating ones; it grows on
dead hardwoods.

Serpula lacrymans (Dry Rot Fungus)

FRUITING BODY annual, forming widely-spreading, nearly flat, fanlike sheets on
horizontal substrates, but sometimes bracketlike ifgrowingonverticalsubstrates; softand
spongy when fresh, 5 cm-1 m (3 ft.) broad or more. Upper surface (or free margin) silvery-
white to gray, hairy. Flesh thin, dingy yellowish; odor often unpleasant, musty. FERTILE
SURFACE consisting of very shallow (1 mm deep), large, irregular pits or “pores” formed
by a honeycomb-like network of folds and ridges rather than by tubes; olive-yellow to
brownish-yellow, rusty-brown, orange-brown, orcinnamon. STALK typically absent, but
white or grayish mycelial strands (by which it spreads) often present. SPORE PRINT
orange-brown to orange-yellow to brownish; spores 8-12.5 * 4-6 microns, elliptical,
smooth, thick-walled.
HABITAT: A serious pest of structural wood in old houses, buildings, ships, etc., usually
developing indoors or in poorly ventilated situations, often hiding under floorboards.
Bulging wood and a musty odor are telltale signs of its presence. Common in Europe, where
the ventilation in many houses was sealed off during the war; less common in North
America. It is called dry rot because it extracts water from the wood and cracks it into

610
SERPULA 611

cubical blocks, eventually reducing it to a fine, dark powder. A related species, S. himan-
tioides, turns up occasionally in the wild (on dead conifers), as do Merulius species (see
comments).
EDIBILITY: Utterly and indisputably inedible.
COMMENTS: One of the few fleshy fungi that lives up to the label fungus in its most
pejorative sense—odious, insidious, hideous, obnoxious, downright abominable. Once it
gains a foothold it is hard to eradicate because the often gigantic, padlike fruiting bodies
exude great quantities of water, stimulating further fungal growth. The mycelial strands
spread with astonishing rapidity. Like a horde of hungry army ants in search of food, they
will overrun anything and everything in their way: bricks, stones, tiles, plaster, drain
pipes, wires, leather boots, cement floors, books, tea kettles, evencorpses. Forafascinating
account (and pictures) of some of its more heroic feats, see John Ramsbottom’s
mycological treasure trove, Mushrooms and Toadstools.
The veined or honeycombed network of large, irregular, shallow “pores” is characteristic
of Serpula (colored spores) and Merulius (white spores). Both are now quite rightfully
placed in families of their own, apart from other crust and parchment fungi. Other species:
Cortiophora puleana (“Wet Rot”) is another pest of structural timber; it has a similar
growth habit but has an irregularly wrinkled to bumpy fertile surface. Merulius tremellosus
(see photo on p. 610) has an orange to orange-buff to pinkish, veined or honeycombed fer¬
tile surface and a hairy white cap or free margin; it grows in the wild, mainly on dead hard¬
woods. M. incarnatus is an eastern species with a bright coral-pink cap and paler, duller,
honeycombed or veined fertile surface. Several other species differ microscopically.

Teeth Fungi

HYDNACEAE spores

As their name implies, the teeth fungi produce their spores on pendant spines or “teeth”
(see Color Plate 159). The fruiting body is usually stipitate (equipped with a cap and stem),
with the spines lining the underside of the cap. Hericium, however, grows on wood and has
spines which hang like icicles from a rooting base or network of branches, and several
others are shelflike, i.e., with a cap but no stalk. Those that grow on the ground can be tough
and quite reminiscent of polypores (Hydnellum and Phellodon) or fleshy, brittle, and
agaric-like (Hydnum and Dentinum).
The teeth fungi include many highly distinctive mushrooms and some strikingly
beautiful ones. They are most common and diverse in northern pine and spruce forests, but
relatively sparse in our area. Hericium and Dentinum are excellent eating and have the
added advantage of being virtually unmistakable. Most of the others are either too tough or
too bitter (or too tough and too bitter) to eat.
Just as all the boletes were once lumped together in Boletus, so all the teeth fungi were
originally placed in a single genus, Hydnum. Several families and genera are now
recognized based on differences in the shape and texture of the fruiting body and color and
ornamentation of the spores. However, to facilitate identification all of the teeth fungi
have been retained here in a single family, divided into the five groups keyed below.

Key to the Hydnaceae

1. Growing on decaying conifer cones or sometimes in mats or debris made up partly of cones;
cap and stalk hairy, brown to dark brown; stalk only 0.5-3 mm thick . Auriscalpium, p. 629
1. Not as above; not growing on cones; stalk if present usually thicker.2
612 HYDNACEAE

2. Fruiting body rubbery and flexible, small (cap typically 5 cm broad or less), translucent white
to watery gray or with a brownish cap; stalk lateral (attached to side of cap); spines minute
and very short . (see Pseudohydnum gelatinosum, p. 671)
2. Not as above . 3
3. Growing on wood .4
3. Growing on ground (or rarely on very rotten wood) . 5
4. Fruiting body a branched framework or unbranched cushion of tissue from which spines are
suspended (i.e., icicle-like); lacking a distinct cap .Hericium, p. 613
4. Fruiting body resupinate (i.e, crustlike or sheetlike) or more often with a cap (clearly defined
upper sterile surface), the spines lining the underside of the cap Echinodontium Sc Allies, below
5. Fruiting body tough and fibrous or woody or if soft and spongy then with a tough, fibrous inner
core (especially in stalk); fruiting body sometimes encompassing needles and other debris as it
grows; surface of cap often roughened irregularly by projecting spikes, lumps, warts, ridges,
pits, etc.; stalk continuous with cap and sometimes not well-defined; flesh sometimes showing
zones or lines when fruiting body is sliced open lengthwise Hydnellum& Phellodon, p. 622
5. Not as above; fruiting body fleshy and usually brittle (not spongy or woody), not typically
absorbing needles and other debris as it grows; cap smooth, cracked, or scaly but not normally
roughened by projecting spikes, lumps, pits, or ridges; stalk usually well-defined; flesh not
often zoned by lines.Hydnum, Dentinum, & Allies, p. 616

ECHINODONTIUM & Allies

Medium-sized to large, fleshy to very tough or woody fungi growing on wood. FRUITING BODY
typically shelflike or hooflike. CAP clearly defined, smooth to hairy, rough, or cracked. SPINES
long or short, variously colored. STALK typically absent (except in Mycorraphium). SPORE
PRINT typically white (when obtainable). Spores smooth or minutely spiny, sometimes amyloid.

THESE shelving mushrooms are more likely to be mistaken for polypores than for other
teeth fungi. However, they bear their spores on numerous downward-pointing spines
instead of in tubes or pores. They differ from the genus Hericium in possessing a clearly
defined, sterile upper surface (cap) and fertile lower surface (spine layer). Several diverse
genera are keyed below. They are not closely related but share a similar growth habit. Only
one species is described, the others being rare or absent in the West. None are wortheating.

Key to Echinodontium & Allies

1. Stalk often present; cap more or less kidney-shaped, less than8 cm broad, white to tan(but may
blacken when handled); spines only 1 -3 mm long, whitish becoming pinkish, brown, or cinna¬
mon in age; found on dead hardwoods in eastern North America .
.Mycorraphium (-Steccherinum) adustum
1. Not as above . 2
2. Entire fruiting body white to yellowish or pale ochre or pale gray when fresh (but may discolor
in old age), fleshy or tough but not woody; flesh white; growing mostly on hardwoods ... 3
2. Not as above . 5
3. U sually growing in overlapping, shelving masses high up on living hard woods (especially maple);
caps usually 10-30 cm broad, several arising from a common base; fairly common in northern
North America (see photo at top of p. 613) ... Climacodon (-Steccherinum) septentrionale
3. Not as above .4
4. Cap(s) usually roughened or with sterile upright spines; spines on underside usually quite long
(up to 2 cm); spores amyloid; usually on hardwood stumps; rare .... Creolophus cirrhatus
4. Not with above features; pores often present when very young (the tube walls that form the pores
often breaking up in age to form spines); common .(see Polyporaceae & Allies, p. 549)
5. Fruiting body woody, more or less hoof-shaped; flesh bright orange to rusty-red or cinnamon;
growing on conifers in western North America .Echinodontium tinctorium, p. 613
5. Not as above; pores often present when young (their walls breaking up to form spines or “teeth”
in age) .(see Polyporaceae & Allies, p. 549)
Climacodon septentrionale looks like a polypore but has spines under the cap instead of pores (see
key at bottom of p. 612). It forms large shelving masses on northern hardwoods such as maple.

Echinodontium tinctorium (Indian Paint Fungus; Toothed Conk)

FRUITING BODY shelflike, very tough or woody, often perennial, up to 15 cm thick.
CAP 4-25 cm broad, more or less hoof-shaped; surface dry, finely hairy to rough, often
covered with moss. Flesh very tough or woody, bright orange to rusty-red or cinnamon,
zoned. SPINES brittle, blunt, thick, flattened, long (1-3 cm); grayish to pale olive-buff, the
tips sometimes darker. STALK absent. SPORE PRINT white (when obtainable); spores
5.5-8 * 3.5-6 microns, elliptical, minutely spiny, amyloid.
HABITAT: Solitary or several on living or occasionally downed conifers; known only
from western North America, where it favors mountain conifers such as fir and hemlock
(I haven’t seen it on the coast). It causes an extensive white heart rot in its host.
EDIBILITY: Unequivocally inedible, but can be used as a red dye. Its common
name is a tribute to its use by Native Americans in the preparation of war paint.
COMMENTS: The woody hoof-shaped fruiting body looks like a conk, but has long spines
or “teeth” on its underside. The bright orange to reddish-orange flesh is also distinctive. It
is not closely related to other teeth fungi and is now sequestered in a family of its own.

HERICIUM
Medium-sized to very large fleshy fungi growing on wood. FRUITING BODY usually white to
yellowish or pale salmon, branched or unbranched but lacking a distinct cap; spores borne on
clusters or rows of delicate hanging spines. SPORE PRINT white. Spores more or less round,
smooth or minutely roughened, amyloid.

A PRISTINE full-grown Hericium is a breathtakingly beautiful sight. The fruiting body

is unmistakable: a mass of fragile “icicles” suspended from a branched supporting frame¬
work, or in the case of H. erinaceus, from a tough, unbranched cushion of tissue.
Hericiums are as delectable as they are beautiful, providing they are not too old or too
tough—and providing you can bear to pick them in the first place. They are excellent fresh
as well as marinated or pickled, and they have absolutely no poisonous look-alikes. They
grow exclusively on wood and are easily cultivated. Their amyloid spores and novel
fruiting body have led some taxonomists to reward them with a family of their own.
Four well-known species occur in North America. Three favor hardwoods, while H.
abietis grows on conifers. In most regions, including California, they are infrequent to
rare, so you should consider yourself fortunate to find one.
614 HYDNACEAE

Key to Hericium
1. Fruiting body unbranched, consisting of a tough cushion of tissue from which long (2-7 cm)
spines are suspended (but spines short in one eastern variety); found on hardwoods, especially
oak .H. erinaceus, 615
1. Fruiting body branched, the spines hanging from the branches or branch tips (sometimes
scarcely branched and very compact, but if so, then usually growing on conifers) .2
2. Growing on conifers in western North America; fruiting body white to salmon- or yellowish-
tinged when fresh .H. abietis, below
2. Growing mainly on hardwoods; fruiting body white when fresh (but may turn yellowish in
age); widely distributed . 3
3. Spines rather short (3-10 mm), arranged in rows along the branches (like teeth on a comb);
branching usually open rather than compact; fruiting body often delicate H. ramosum, p. 615
3. Not as above; spines often long (up to 4 cm), arranged mostly in tufts or clusters, especially
at the branch tips; branching open or compact .H. coralloides {see H. abietis, below)

Hericium abietis (Conifer Coral Hericium) Color Plate 163

FRUITIN G BODY 10-75 cm or more broad and high at maturity, consisting of an open to
compact branched framework from which tufts of icicle-like spines hang; branches arising
from a thick, tough, rooting base; color variable; white to creamy, yellowish-buff, pale
ochraceous, or salmon-buff. SPINES up to 25 mm long but usually 5-10 mm; soft but
brittle, arranged in tufts or clusters that are mainly grouped at the branch tips. SPORE
PRINT white; spores 4.5-5.5 * 4-5 microns, round or nearly round, smooth or minutely
roughened, amyloid.
HABITAT: Solitary or sometimes several together on dead conifers (especially fir and
Douglas-fir); known only from the Pacific Northwest and northern California, fruiting
mainly in the fall. It is rather infrequent but locally common, particularly at higher eleva¬
tions (the largest fruitings I’ve seen were in the Cascades). It appears year after year on the
same logs, causing a conspicuous white pocket rot. The closely related H. coralloides
(see comments) favors hardwoods and is more widely distributed.
EDIBILITY: Eminently edible, delectably delicious. When thoroughly cooked it is
reminiscent of fish and is excellent sauteed, curried, or marinated. Its large size (a 100-lb.
specimen was wheeled into one mushroom show!) plus its distinctive appearance make it an
excellent mushroom for beginners. However, its breathtaking beauty poses a minor moral
dilemma: should one ruthlessly uproot it for the sake of a meal, or leave it for others to see?
COMMENTS: It is hard to believe that this astonishing Hericium, with its cascading
clusters of pristine “icicles,” is a fungus. It is easily distinguished from other Hericiums by its
white to salmon-buff color, clustered spines, and growth on conifers. It has long been a
favorite among nature-lovers and photographers in the Pacific Northwest, but only
recently has it been recognized as distinct from the better known, equally beautiful and
delicious H. coralloides. Some investigators, in fact, still consider the two to be the same
species. Others reserve the name H. coralloides for a pure white (when fresh) hardwood-
loving form with slightly longer spines (up to 4 cm) and slightly larger spores. This form
occurs in eastern North America, but reports of it from the West are based at least in part
on H. abietis. To complicate matters, it has now been suggested that the name H. coral¬
loides is better applied to H. ramosum, and that the eastern form traditionally called
H. coralloides should be given a new name, H. americanum! To muddle matters even
more, there are several growth forms of H. abietis, including a very compact, scarcely
branched one (formerly H. weirii) that somewhat resembles H. erinaceus, and an exten¬
sively branched one with very short spines (1-5 mm) that mimics H. ramosum (both forms
grow on conifers, however). Since all of these Hericiums are equally edible, their exact
identities needn’t concern you—at least, they don’t concern me!
HERICIUM 615

Hericium ramosum (Comb Hericium) Color Plate 164

FRUITING BODY 8-35 cm broad and 6-15 cm high when mature, comprised of an open
framework of rather delicate, toothed (spine-laden) branches arising from a tough,
repeatedly branched rooting base or “trunk”; pure white when fresh, discoloring creamy to
buff or yellowish-tan in old age. Flesh white. SPINES more or less evenly distributed in
lines along the branches (like teeth on a comb), sometimes also in small tufts at the branch
tips; spines rather short (3-10 mm long or up to 25 mm long in the tufts). SPORE PRINT
white; spores 3-5 * 3^1 microns, nearly round, smooth or minutely roughened, amyloid.
HABITAT: Solitary or in small groups on fallen hardwood branches, logs, and stumps;
widely distributed. It is said to be the most common Hericium in North America, but like
the others, is uncommon in our area. I have found it in the fall, winter, and early spring on
dead oak, and I have seen it in the summer on aspen and poplar in New Mexico.
EDIBILITY: Edible and delicious when cooked slowly, but not as fleshy as the other
Hericiums.
COMMENTS: Formerly known as H. laciniatum, this lovely species is smaller and more
delicate than H. abietis and H. coralloides, with slightly shorter spines and smaller spores.
Also, the branching is more open and the spines are arranged in lines lengthwise along the
branches, rather than exclusively in tufts. A compact form of this species occurs, but is rare.

Hericium erinaceus (Lion’s Mane Hericium; Old Man’s Beard)

FRUITING BODY an unbranched mass of numerous long, closely-packed, icicle-like
spines hanging from a tough, solid, hairy, rooting cushion of tissue; 8-40 cm or more broad
and high when mature; entirely white when fresh, discoloring yellowish to tan or dingy
ochre in age. Flesh white. SPINES (1) 2-5 (7) cm long, soft and pliant when fresh, with
pointed tips. SPORE PRINT white; spores 5-6.5 * 4-5.5 microns, broadly elliptical to
nearly round, smooth to minutely roughened, amyloid.
HABITAT: Solitary (rarely several together) on wounds of living hardwoods oronthecut
ends of recently felled logs; widely distributed. In our area it favors oak and is fairly
common (for a Hericium) in the fall, winter, and spring; farther north it grows on maple.
EDIBILITY: Excellent when fresh, but tougher than other Hericiums and sometimes

Left: Hericium ramosum is the most delicate species in its genus. It favors dead hardwoods. Right:
Hericium erinaceus has long spines hanging from an unbranched cushion of tissue. It favors living or
recently killed hardwoods.
Hericium erinaceus. Note how long the spines are in this rather small specimen.

developing a rather sour, unpleasant taste in age. Slow cooking is called for and the base
should not be eaten—it’s so tough that it’s difficult to remove from the tree without a knife!
COMMENTS: The unbranched white to yellowish fruiting body and long, slender spines
distinguish this impressive fungus from its relatives. Though hardly common, it is more
numerous in our area than the branched Hericiums and is one of our most distinctive
wood-inhabiting fungi. Gigantic specimens weighing several pounds each are not un¬
common. An unbranched version completely covered with small short spines occurs on
hardwoods in eastern North America, but I have not seen it in the West.

HYDNUM, DENTINUM,& Allies

(Hedgehog Mushrooms)
Medium-sized to fairly large, fleshy, terrestrial fungi with a cap and stalk. CAP smooth or cracked
to conspicuously scaly. Flesh firm, usually brittle. SPINES soft, brittle. STALK central or off-
center, usually well-developed, fleshy but not woody. SPORE PRINT white (Dentinum) or
brown (Hydnum). Spores smooth (Dentinum); rough to warty or angular (Hydnum).

HEDGEHOG mushrooms are the best known and most common of the teeth fungi, and
as a group are easy to recognize. The fruiting body has a well-defined cap and stalk and
might be mistaken for a gilled mushroom but for the layer of delicate spines or “teeth”
on the underside of the cap. Also significant is the texture of the flesh: fleshy or firm but
brittle, rather than tough and pliant or leathery or woody as in the other major genera of
terrestrial teeth fungi (Hydnellum and Phellodon).
In the small but common genus Dentinum, the spores are white and smooth, the spines
white to pale orange, and the cap usually smooth. In the larger genus Hydnum, the spores
are brown and warty, the spines are variously colored but usually darker than in Dentinum,
and the cap is often scaly. A third genus, Bankera, has a brownish fruiting body and white
roughened spores but is relatively rare, at least in the West.
Although there is no doubt as to what hedgehog mushrooms are, there is, as usual,
considerable controversy as to what hedgehog mushrooms should be called. Some
mycologists campaign for the use of Hydnum instead of Dentinum, and the creation
of the genus Sarcodon to account for the Hydnums of this book. A “correct” classi¬
fication of the teeth fungi may not be essential to your well-being (it certainly isn’t to

616
HYDNUM, DENTINUM, & ALLIES 617

mine), but please bear in mind that the exacting specialists owe their livelihoods to the
resolution of such matters—and that they are doing their best. Giving even tacit approval
to one system of classification at the expense of another is thus transformed into an act of
inordinate importance, with the taxonomist’s professional reputation at stake. Why they
can’t arrive at a consensus is anyone’s guess—but since when do human beings agree on
anything? I for one find them (human beings, that is) even more perplexing and mystifying
than mushrooms!
Dentinums are delicious and a good choice for beginners since nothing poisonous
remotely resembles them. Hydnums, on the other hand, though sometimes beautiful,
are mostly bitter-tasting or of unknown edibility.
Both Dentinum and Hydnum species are strictly woodland fungi with rather erratic
fruiting habits. Lor instance, in Lebruary and March of 1975 Dentinums were out¬
rageously abundant on the poison-oak-shrouded hillsides of our coastal pine forests.
While barely denting the crop I managed to harvest and can over 200 pounds, which I am
still enjoying today. Lor several years thereafter, however, they were practically absent
in the same area, then produced another stupendous crop in 1981. Similarly, I did not find a
single Hydnum fuscoindicum in our area until 1979, when it fruited by the dozens under
tanoak and madrone. Live distinctive hedgehog mushrooms are described here and
several others are keyed out.

Key to Hydnum, Dentinum, & Allies

1. Fruiting body (including spines) white to pale orange to dull orange, the cap sometimes slightly
darker; spore print white; spores smooth .2
1. Not as above; some part of fruiting body usually darker (gray, brown, etc.) . 3
2. Cap convex to plane or depressed, 2-15 cm broad or more; stalk usually at least 1 cm thick
.D. repandum & others, p. 618
2. Cap usually umbilicate (with a navel-like central depression) and typically less than 5 cm
broad; stalk usually less than 1 cm thick . D. umbilicatum (see D. repandum, p. 618)
3. Spore print white; spines usually pale gray or grayish in age; found under conifers (usually pine)
in eastern North America, but rare in the West . 15
3. Spore print brown; spines variously colored but often dark in age; widespread and common 4
4. Flesh showing distinct violet tints (deep violet to lilac-gray) . 5
4. Not as above; flesh white to brownish, vinaceous-buff, etc. (but may stain purple when cut) 6
5. Entire fruiting body (including spines!) violet to deep violet or blackish-violet .
.H. fuscoindicum, p. 622
5. Not as above; spines brownish to cinnamon-brown with paler tips .
.H. cyanellum & H. fuligineo-violaceum (see H. fuscoindicum, p. 622)
6. Base (tip) of stalk and flesh within it black to olive-black, olive-gray, or bluish-green; taste very
bitter or bitter-farinaceous (chew on a piece of the cap) . 7
6. Not as above; taste mild or bitter. 8
7. Flesh becoming pinkish or lilac-tinted when cut; spines whitish or pallid until old age.
. H. subincarnatum (see H. scabrosum group, p. 620)
7. Not as above .H. scabrosum group, p. 620
8. Scales on cap brown to blackish, usually large and conspicuous even when young and often
raised or upturned; spines usually brownish'at maturity; common . . H. imbricatum, p. 619
8. Not as above; cap usually developing scales only in age and/ or differently colored .9
9. Cap white and plushlike when young, yellowish or tan in age; spines dull yellowish or brown
becoming almost blackish in old age; growing under hardwoods in eastern North America
. H. cristatum
9. Not as above . 10
10. Base of stalk usually white from coating of mycelium; fruiting body often large, thick-fie shed,
buff to tan to yellow-brown, orange-brown, or cinnamon; cap usually with small flattened
scales.H. calvatum group Sl others, p. 621
10. Not as above . 11
618 HYDNACEAE

11. Fruiting body developing greenish-olive tones in old age or when dried; flesh often becoming
greenish also; not common.H. fumosum
11. Not as above . 12
12. Flesh staining lilac, purplish, purple-gray, pinkish, or vinaceous when cut (quickly or quite
slowly) and/ or fruiting body often with vinaceous or purplish tints; widely distributed but
especially common in the West ... 13
12. Not as above; cap pale to dull brown or dark reddish-brown, quite hard and woody when dried;
fairly common under conifers in eastern North America, also reported from the Pacific
Northwest and California.H. stereosarcinon
13. Cap smooth to felty or conspicuously cracked but not truly scaly, reddish-brown to grayish-
brown, vinaceous, grayish, or darker; taste mild to somewhat farinaceous or occasionally
bitter . 14
13. Cap usually with scales, brownish-orange to dark brown, grayish-brown, or pinkish-brown;
taste usually bitter .H. subincarnatum (see H. scabrosum group, p. 620)
14. Cap and stalk often with vinaceous or purplish tints; cap often conspicuously cracked(areolate)
in age; common in western North America . . H. rimosum (see H. scabrosum group, p. 620)
14. Cap grayish to grayish-brown or reddish-brown, smooth to minutely areolate (cracked) in age;
widespread but not common (at least in West) H. laevigatum{ see H. scabrosum group, p. 620)
15. Cap some shade of brown at maturity (the margin often paler), not normally scaly but usually
with needles and other debris adhering to the surface; found under conifers (usually pine) in
eastern North America, but rare in West .Bankera fuligineo-alba
15. Not as above; cap more or less grayish-brown, often cracked or scaly in age; widespread but
not common. Bankera carnosa(-B. violescens?)

Dentinum repandum Color Plates 161,162

(Hedgehog Mushroom; Pig’s Trotter)
CAP 2-17 (25) cm broad, broadly convex to plane or depressed, the margin often wavy or
deeply indented and at first inrolled; surface dry, more or less smooth, but sometimes
cracking into scales in age; pale flesh-color to pale or dull orange, orange-tan, salmon, tan,
or pale cinnamon to reddish-tawny (but white to creamy in var. album)] bruised areas often
darker orange. Flesh thick, occasionally zoned concentrically, firm, brittle, white, usually
discoloring yellow to yellow-ochre or orange-brown when bruised; odor mild, taste mild to
somewhat bitter or peppery. SPINES 2-7 mm long, whitish to yellowish, salmon-buff, or
pale orange, bruising dark orange to ochraceous; slender, brittle but soft, usually
decurrent. STALK 3-10 cm long, (0.5) 1-3 (5)) cm thick, central or off-center, equal or
enlarged below or occasionally tapered downward, firm, white or colored like cap but
usually paler; bruising ochre to dark orange-brown; smooth or downy at base. SPORE
PRINT white; spores 6.5-9 * 5.5-8 microns, broadly elliptical to nearly round, smooth.
HABITAT: Solitary, scattered, gregarious, or in troops on ground under both hard¬
woods and conifers; widely distributed throughout the north temperate zone and probably
the most common of all the teeth fungi. In California it fruits throughout the mushroom
season but develops slowly and normally does not peak until the late winter or early spring,
when the wild irises are in bloom (see Color Plate 161) and most other fungi have long
since rotted away. In our area it favors fern, bramble, and poison oak thickets under pine;
in the Pacific Northwest and Rocky Mountains it grows under a wide range of conifers,
and in eastern North America it is often common under oaks in the summer and fall.
EDIBILITY: Edible and choice! It is comparable to the chanterelle in color, texture, and
flavor, and like that species is usually maggot-free. It is easier to clean, however, and is also
easier to recognize.The peppery taste (if present) disappears in cooking. It’s superb in
casseroles, tomato sauces, or sauteed with sour cream, but should be cooked slowly and
lengthily to make it tender. Be careful to keep the spines clean while picking them!
COMMENTS: Also known as Hydnum repandum, this late bloomer resembles a dull-
DENTINUM 619

colored chanterelle when first spotted amongst the needles and humus. However, the
white to pale orange spines on the underside of the cap immediately distinguish it, making it
one of the safest of all edible mushrooms. The cap color, though variable, is typically some
shade of pale orange to pinkish-orange to reddish-tan, with wounded areas darker orange.
The white form (var. album) does not seem to occur in our area, but is common farther
north under conifers, especially Sitka spruce. Other species: Var. macrosporum has larger
spores but is otherwise similar; D. umbilicatum is a closely related, equally edible conifer-
lover with a smaller (usually less than 5 cm) umbilicate cap, a slimmer stalk (typically less
than 1 cm), and larger spores. It is similar in color and widely distributed, and sometimes
mingles with D. repandum. Two edible white or whitish southern species should also be
mentioned: D. albomagnum and D. albidum. The first has a mild taste and non-staining
flesh, while the latter has an acrid taste and smaller spores.

Hydnum imbricatum (Shingled Hedgehog) Color Plate 159

CAP 5-20 cm broad, convex to plane or centrally depressed; surface dry, buff to pale brown
or dull reddish-brown, but covered with large, coarse, broad, raised or shingle-like,
darker brown to nearly black scales that are often upturned in age; becoming darker
brown throughout in age and sometimes cracking, with the scales sometimes wearing off
except at the center. Flesh thick, firm but brittle; pallid to grayish, tan, or brownish; odor
mild or when dry somewhat smoky or chocolate-like; taste mild to bitter. SPINES pale
brown or grayish or pallid becoming dark brown in age, 2-15 mm long; soft, brittle, often
slightly decurrent. STALK 4-10 cm long, 1.5-3.5 (5) cm thick, central or off-center, often
enlarged below; some shade of brown, often hollow toward the top in age; usually more or
less smooth. SPORE PRINT brown; spores 6-8 x 5-7 microns, nearly round but promi¬
nently warted (angular-nodulose or shaped like a Maltese cross).
HABITAT: Solitary to gregarious onground in woods; widely distributed—it is probably
the most common Hydnum in North America. In many regions it is abundant under
conifers in the late spring, summer, and fall. In our area it occurs under hardwoods in
the late fall and winter, but is rare. I have seen enormous fruitings under spruce and fir.

EDIBILITY: Edible, but of poor quality. Many collections have a bitter taste and par¬
boiling does not necessarily help, plus it causes indigestion in some people. The European
version is apparently better because it is often sold in markets there.

Hydnum imbricatum is easily told by the large brown to blackish scales on the cap, which make it look
like a charred macaroon. See color plate for close-up of spines, which range from grayish to brown.
620 HYDNACEAE

COMMENTS: Also known as Sarcodon imbricatum, this arresting hedgehog is easily

identified by the prominent brown to blackish scales on its cap. The stalk base is not olive-
black as in the H. scabrosum group, and the cap is scalier—even when young. In old age the
central depression of the cap may become perforated, i.e., join up with the hollow in the
upper portion of the stalk. For similar species, see the H. scabrosum group.

Hydnum scabrosum group (Bitter Hedgehog) Color Plate 165

CAP 4-14 (20) cm broad, convex to plane or slightly depressed; surface dry, at first smooth
but soon becoming cracked and scaly, the scales cinnamon-brown to reddish-brown
becoming dark chocolate-brown to vinaceous-brown at maturity and the background
dingy yellowish-brown or darker. Flesh thick, brittle, firm, white or buff, but olive-gray
to olive-black in base of stalk; odor farinaceous or smoky; taste strongly bitter and/ or acrid
to farinaceous. SPINES often unequal in length, 2-10 mm long, pallid or buff, darken¬
ing to tan with paler tips, then darker brown in old age; usually slightly decurrent. STALK
2.5-10 cm long, 1-3.5 cm thick, usually tapered below, central or off-center, often curved,
flesh-color becoming brown or dark brown in age, the base blackish-olive to grayish-olive,
olive, or dark bluish-green; firm, solid. SPORE PRINT brown; spores 6-7.5 x 4-5.5
microns, elliptical to round, prominently warted. Cap tissue staining blue-green in KOH.
HABITAT: Solitary to scattered or gregarious on ground under conifers or sometimes
hardwoods; widely distributed. It is fairly common in the late summer and fall in the
Pacific Northwest and northern California, but rare in our area.
EDIBILITY: Unequivocally and indisputably inedible due to the awful taste.
COMMENTS: The scaly reddish-brown to brown cap, blackish-olive to blue-green stem
base, and bitter taste distinguish this species from most other Hydnums. The scales are
usually quite conspicuous, but not as large as those of H. imbricatum. H. fennicum is a
very similar species with a scaly reddish-brown to brown cap and blackish stem base.
However, it does not stain blue-green in potassium hydroxide (KOH) and it has an even
more intensely bitter taste. It grows under both hardwoods and conifers and is wide¬
spread. Three other somewhat similar species are quite common in the West: H. sub-
incarnatum is usually bitter-tasting and similarly colored or more vinaceous, but its flesh
usually stains pinkish or lilac when cut, its spines are pallid or whitish becoming buff or pale
brown only in old age, and its stalk base may or may not be olive-tinted. In California it
grows under both hardwoods and conifers, sometimes in the company of H. scabrosum.
H. rimosum (see photo below) is also similar, but has a more or less mild (or rarely slightly

Hydnum rimosum (see comments under H. scabrosum group) is not nearly as scaly as H. imbricatum,
but the surface of its cap often cracks in age.
HYDNUM 621

acrid) taste, is not normally olive at the stem base, and has a vinaceous- or purple-tinged
cap that usually develops cracks in age but is not scaly when young. It is fairly common
under conifers in the Pacific Northwest, and I have found it in our area in the winter and
in the Sierra Nevada in the spring. Finally, there is H. laevigatum, a widespread but
infrequently-encountered species that has a smooth to minutely cracked cap and, like
H. rimosum, often stains purplish when cut open. None of these are worth eating.

Hydnum calvatum group (Robust Hedgehog)

CAP (5) 10-25 (35) cm broad, convex to nearly plane or somewhat irregular; surface dry,
soon breaking up into small, flattened scales and usually cracking in age; creamy-buff to
yellowish-tan or pale cinnamon-brown, the scales slightly darker and becoming brownish
where bruised; margin often lobed. Flesh very thick, firm, pallid to pinkish-buff or pale
brownish, or grayer near the spines; odor and taste variable: mild to spicy-fragrant to
farinaceous. SPINES pallid, soon darkening to brown or grayish-brown, the tips usually
paler; 2-12(15) mm long, usually very uneven in length or many aborted or fused together
or forked or with small swellings; sometimes decurrent. STALK 2-9 cm long, 2-4 (6) cm
thick, central or off-center, usually narrowed at the base and sometimes rooting; solid,
firm, colored like cap or slightly paler, the base often whitish from mycelium. SPORE
PRINT brown; spores 4-5.5 * 3-5 microns, nearly round to elliptical but prominently
warted (angular-nodulose). Cap surface staining blue-green to olive-black in KOH.
HABITAT: In groups or clumps, often with small or aborted fruiting bodies present, on
ground in mixed woods and under conifers; known only from western North America.
I’ve found it in August under spruce in New Mexico; it is also fairly common in Oregon.
EDIBILITY: Unknown—but fleshy enough to warrant cautious experimentation.
COMMENTS: The large size, thick flesh, tendency to grow in groups or clumps, whitish
stalk base, and presence of small appressed (flattened) scales on the cap distinguish this
species from most other Hydnums. The color is never as dark as that of H. imbricatum, nor
the cap as coarsely scaly; the stalk is not olive or blue at the base as in the H. scabrosum
group. A similar species, H. crassum, is slightly brighter in color, has larger spores, and
its flesh may stain yellow-green when cut. Its cap surface stains brownish in KOH and
it also occurs under conifers. A somewhat similar and widespread species, H. mar-
tioflavum, has bright cinnamon-orange spines when young and a cinnamon to tawny cap.

Left: Hydnum calvatum group. Note small flattened scales on cap. Right: Underside of Hydnum
fuscoindicum (see description on p. 622). Entire fruiting body is deep purple to black in this beautiful
and unmistakable hedgehog mushroom.
622 HYDNACEAE

Hydnum fuscoindicum (Violet Hedgehog)

CAP 4-18 cm broad, convex to plane or centrally depressed; surface dry, at first smooth
but usually cracking to form scales in age; violet-black to bluish-black, black, or raisin-
colored; margin often somewhat paler or purpler and wavy. Flesh thick, firm but brittle,
deep slate-purple or violet; odor and taste mild to somewhat farinaceous or cinnamon-like.
SPINES deep violet to deep bluish-violet to deep lavender, the tips usually paler or lilac;
soft, brittle, 2-6 (15) mm long, usually decurrent. STALK 2-10 cm long, 1-2 (3.5) cm thick,
equal or more often tapered below, central or off-center, firm, colored more or less like the
spines (deep purplish). SPORE PRINT brown; spores 5-7 * 4.5-6.5 microns, broadly
elliptical to nearly round, prominently warted.
HABITAT: Widely scattered to gregarious on ground in woods; locally common but
erratic in its fruiting behavior—absent some years and abundant others; known only from
western North America. In our area it fruits in the fall and winter under tanoak and
madrone at higher elevations in the coastal mountains; in the Pacific Northwest, however,
it favors conifers, especially hemlock and pine. It is yet another example of a conifer-lover
that crosses over to tanoak-madrone (others include Ar miliaria ponder osa, Cantharellus
subalbidus, Hygrophorus chrysodon, and Tricholoma aurantium).
EDIBILITY: Not recommended. Although not exactly bitter-tasting, small pieces which
I sampled caused a peculiar burning sensation in the back of my throat. It would be
interesting to see what color it yields as a dye.
COMMENTS: The striking deep violet color, which is reminiscent of Cortinarius vio-
laceus, sets apart this attractive Hydnum (see photo at bottom of p. 621). Its color makes it
hard to pick out in the forest gloom, and even the most eagle-eyed Hydnum-hound is likely
to miss it or else mistake it for an old, blackened Russula albonigra. Phellodon atratus is
somewhat similar, but is bluish-black and much smaller and tougher. Other purplish
Hydnums include: H. cyanellum, with a similarly colored cap but paler (“lilac-gray”) flesh,
a bitter taste, and cinnamon-brown spines with whitish tips (it was originally described
from California, but I have not seen it); H. fuligineo-violaceum, with a vinaceous-brown
cap, brown or cinnamon-brown spines, an acrid taste, and gray flesh in the base of the
stalk; and H. rimosum(see comments under the H. scabrosum group), which is vinaceous-
brown to vinaceous-buff rather than deep purple.

HYDNELLUM & PHELLODON

Small to medium-large, terrestrial fungi with a cap and stalk (or sometimes several caps). CAP
columnar to top-shaped becoming plane, depressed, or irregular; often spongy or felty when young
and often lumpy, pitted, or ridged in age. Flesh tough; spongy to fibrous; pliant or woody, often
duplex. SPINES typically short, often blunt; variously colored. STALK central or off-center,
sometimes nearly absent; tough or woody, continuous with cap. SPORE PRINT brown (Hyd-
nellum) or white (Phellodon). Spores roughened by warts or spines.

THESE tough or woody, terrestrial teeth fungi occur primarily in coniferous forests.
They have both a cap and stalk, but the latter is sometimes present only as a poorly-defined
tapered base. Many species have the general aspect of a polypore, but a closer look reveals
the presence of spines rather than pores on the underside of the cap, although the spines
are sometimes so short and blunt that they look like minute warts.
The fruiting body develops and decays over a long period of time, frequently engulfing
needles, twigs, and other debris in the growth process. If there is a dry spell, growth may
cease, then begin anew around the margin of the cap when it is damp again. As a result the
cap frequently has an irregular or somewhat misshapen appearance; several caps may fuse
HYDNELLUM & PHELLODON 623

together or arise in a rosette from a common stem, and their surfaces are often pitted,
ridged, or lumpy. This indeterminate growth pattern plus the tough and pliant to fibrous
or woody texture separate Hydnellum and Phellodon from the fleshier, more brittle
hedgehog mushrooms (Hydnum and Dentinum). In many Hydnellums and some Phello-
dons the cap is spongy or felty to the touch when young or actively growing, and is
sometimes beaded with colored droplets. However, beneath or within the spongy-felty
outer layer there is a tougher, corky or woody-fibrous core, particularly in the stalk. The
flesh thus has two distinct layers of different textures, i.e., it is duplex.
Hydnellum contains about 50 species in North America. They are medium-sized to
fairly large and have brown spores. Phellodon includes only a handful of species, but some
of them are quite common. They are distinguished from Hydnellum by their white spores
and smaller size. Neither genus is common in our area but both are prominent farther
north. Because of their indeterminate growth most species vary tremendously in their
appearance according to environmental conditions. It is therefore imperative, when using
the following key, to have several specimens in hand if at all possible. Seven species are
described fully and others are keyed out. All are much too tough and/or bitter to eat.

Key to Hydnellum & Phellodon

1. Flesh in the cap and/ or stalk black or tinted or zoned (lined) with blue or violet (purple-black,
blue-gray, etc.) when fruiting body is cut in half lengthwise .2
1. Not as above; flesh orange, cinnamon, brown, or shades thereof .9
2. Flesh in lower half of stalk reddish-orange to bright rusty-brown or salmon; flesh in cap usually
zoned with brown and blue or mauve (sometimes faintly) . . . H. caeruleum & others, p. 625
2. Not as above ...3
3. Spines bright blue to dark blue; stalk with a swollen buried base or “tuber”; found in eastern
North America; rare .H. scleropodium
3. Not as above .4
4. Flesh in lower half of stalk bluish-black to purple-black; cap whitish to yellowish to tan or
violet-tinged when fresh; odor often fragrant. H. suaveolens & others, p. 624
4. Not as above . 5
5. Flesh black to grayish-black (not blue or violet-tinted) in both cap and stalk; spore print white
(when obtainable); found mainly from the Rockies eastward P. niger(see P. atratus, p. 629)
5. Not as above .6
6. Cap rather small (5 cm broad or less); stalk slender (averaging 3-5 mm thick); spore print
white . 7
6. Cap medium-sized to rather large (usually 4 cm broad or more); stalk usually thicker; spore
print brown . 8
7. Flesh purple-gray to purple-black; cap typically dark brown to purplish-gray with a pale margin
.P. melaleucus(see P. atratus, p. 629)
7. Flesh purple-black to bluish-black; cap more or less same color .P. atratus, p. 629
8. Found in eastern North America; fruiting body slate-gray to black .
.H. nigellum(see P. atratus, p. 629)
8. Found mainly from the Rocky Mountains westward; flesh and/or cap usually with some
blue or violet tones .H. cyanopodium & H. regium (see H. caeruleum, p. 625)
9. Spines bright yellow at least at the tips; cap usually with bright yellow to olive-yellow tones at
least at the margin; found in eastern North America.H. geogenium
9. Not as above . 10
10. Flesh yellow-orange to bright orange, rusty-orange, or reddish-orange, at least in the stalk; cap
and/or spines often showing same colors . 11
10. Not as above . 12
11. Flesh in cap and stalk orange to orange-red or rusty-cinnamon; cap surface with cinnamon,
orange, or rusty-cinnamon tones, at least in age .H. aurantiacum & others, p. 626
11. Flesh in cap duller (mauve, grayish, brownish, etc.); cap surface also typically duller .
.H. caeruleum & others, p. 625
624 HYDNACEAE

12. Odor sweet and/or taste of flesh very acrid (peppery); cap beaded with bright red to dark red
droplets in wet weather .H. peckii &l others, p. 627
12. Cap not beaded with red droplets, or if so then taste not typically acrid nor odor sweet ... 13

13. Cap usually small (6 cm or less); stalk often slender (less than 1 cm thick); spore print white 14
13. Mature cap often more than 5 cm broad; stalk often thick; spore print brown or brownish 15
14. Spines pale cinnamon to brown at maturity; cap usually zoned .P. tomentosus, p. 628
14. Spines grayish at maturity .P. confluens{see P. tomentosus, p. 628)
15. Stalk exuding yellow juice when broken or crushed (if fresh); cap pale to dark brown, or some¬
times also with yellow-brown tones and exuding a brownish juice when fresh . . H. mirabile
15. Not as above . 16
16. Cap smooth or radially ridged but not lumpy, warty, or spongy, often zoned concentrically (see
Color Plate 157) with various shades of brown, cinnamon-brown, pinkish-brown, etc.; stalk
not spongy or bulbous .H. zonatum (see H. scrobiculatum, p. 627)
16. Not as above . 12
17. Associated with hardwoods (mainly oak) in eastern North America . 18
17. Associated principally with conifers; widely distributed . 19
18. Stalk very spongy and swollen or bulbous; cap and stalk brown to rusty-brown or cinnamon-
brown and finely hairy or velvety; common .H. spongiosipes
18. Not as above; cap very dark brown to blackish in old age .H. piperatum
19. Cap sometimes beaded with pinkish droplets in wet weather; odor typically mild or faint but
not farinaceous; common under pines in northeastern North America .
. H. pineticola (see H. peckii, p. 627)
19. Cap sometimes beaded with red to dark red droplets in wet weather; odor usually farinaceous;
widely distributed under various conifers . H. scrobiculatum & others, p. 627

Hydnellum suaveolens (Fragrant Hydnellum)

CAP (3) 5-15 (30) cm broad when mature, top-shaped becoming plane to somewhat
depressed, often with needles or other debris incorporated into it; surface white and thinly
felty or velvety at first, often bumpy and/ or pitted in age, soon becoming yellowish to tan,
brownish, olive-brown, or violet-gray from the center outward (margin often paler);
usually staining brown where bruised. Flesh duplex, whitish to yellowish-buff zoned with
blue lines in the cap, entirely deep blue to purple-black in the stalk; odor often strongly
fragrant (like anise or peppermint). SPINES whitish to creamy when young, becoming
grayish-brown with pallid tips in age; short (up to 3 mm long), irregularly decurrent.
STALK 1 -5 cm long, 1-3 cm thick, central or slightly off-center, very tough, the base
usually swollen, rooting; grayish-blue to bluish-black. SPORE PRINT brown; spores
4-6 * 3-4 microns, elliptical to nearly round, prominently warted.
HABITAT: Solitary to gregarious or in fused clusters under northern and montane
conifers, late summer through early winter; widespread but especially common in the
Rocky Mountains. I have seen large fruitings under spruce and fir in Idaho and New
Mexico, but have yet to find it in our area.
EDIBILITY: Unknown, but much too fibrous to be worthwhile.
COMMENTS: The blue-lined flesh in the cap and blue-black flesh in the stalk plus the
flagrantly fragrant (fragrantly flagrant?) odor make this an easy species to identify. In
some collections the fragrance is absent, but in others it is overpowering—I once had to
remove two specimens from my car because their aroma was so heady! H. peckii is often
fragrant but much different in color, while H. caeruleum shows orange-red flesh in the
stalk. Other species: H. cruentum, described from Nova Scotia, has a menthol odor,
but when young and fresh its cap has red droplets and it spines are lilac- or bluish-tinged.
Hydnellum caeruleum. Note the conspicuously zoned flesh in the sliced specimen at left.

Hydnellum caeruleum (Blue-Gray Hydnellum)

CAP 3-12 (17) cm broad, often with leaves or needles incorporated into it; top-shaped
becoming plane or slightly depressed; surface felty or velvety at first, often bumpy and/ or
pitted in age and with matted hairs; mauve to pale blue, whitish, or tan when young,
becoming light to dull dark brown or even blackish from the center outward (often a
mixture of these colors), the soft, felty growing margin white or pale blue to bluish-gray.
Flesh duplex: upper or outer layer spongy, inner core tough and fibrous; zoned variously
with bluish, blue-gray, mauve, and brown in the cap; bright rusty-colored to orange-red
in stalk; odor and taste farinaceous. SPINES whitish when young or tinged blue, becoming
brown to dark brown with pallid tips in age; rather short (1-5 mm long), often decurrent.
STALK 2-9 (12) cm long, 1-3 cm thick, central or off-center, very tough, often rooting
deeply in humus; equal or thicker at either end, buff to brown or orange-brown, but usually
covered with debris. SPORE PRINT brown; spores 4.5-7 * 3.5-5 microns, nearly round
to elliptical and irregularly lobed or warted.
HABITAT: Solitary to gregarious or in fused clusters on ground in woods; widely dis¬
tributed. It is common in the Pacific Northwest under pines and other conifers, but in our
area I find it under oak, tanoak, and madrone, usually early in the fall.
EDIBILITY: Indisputably inedible.
COMMENTS: Like most Hydnellums this species develops over a period of several weeks
and undergoes a number of confusing color changes. Y oung specimens are more or less top-
shaped, and have a thick, felty cap margin that shows white and/or bluish tints. In age,
however, or after being battered by rain, the cap becomes depressed (wouldn’t you?), the
margin thins out, and the color becomes darker or duller brown. However, the bluish lines
in the flesh of the cap (though sometimes faint) and the bright rusty- to orange-red flesh
in the stalk (which distinguishes it from H. suaveolens) are fairly constant characters.
Other species: H.ferrugipes of eastern North America has an orange-red stalk, but favors
hardwoods. H. cyanopodium has a vinaceous-blue cap shading toward lavender (or
whitish at margin) that is often beaded with red droplets in wet weather, flesh that is
zoned bluish-black, and spores shaped like jacks. H. regium often forms compound
fruiting bodies (with several caps); its cap is violet-black with a paler margin and the flesh
is brown to pale orange in the stalk and brownish to grayish with violet tones in the cap.
The latter two species occur under conifers (mainly spruce and pine) in northern Cali¬
fornia, the Pacific Northwest, and the Rocky Mountain region.

625
Hydnellum aurantiacum looks like a terrestrial polypore, but has minute spines on underside of cap
instead of pores. Lumpy specimen on right is young and actively growing.

Hydnellum aurantiacum (Orange Hydnellum)

CAP 3-15 cm broad, columnar or somewhat top-shaped becoming plane or depressed in
age; often with pine needles and other debris incorporated into it; surface velvety and
suedelike when fresh and often roughened by projecting knobs and lumps when mature, or
in some forms with radiating ridges; white when young or on actively growing margin,
otherwise orange to rusty-orange to rusty-cinnamon (or mixture of these colors), and
eventually darker (brown) in old age. Flesh thick, tough and corky except for frequent
presence of a spongier outer or upper layer; orange to rusty-cinnamon to orange-red in
both the cap and stalk; odor mild, taste bitter to farinaceous. SPINES short and blunt
(1^1 mm long), whitish to grayish or orange, becoming brown in age with the tips often
paler. STALK 2-6 cm long, 1-3 cm thick, usually central, very tough or woody, equal or
tapered downward or in one form enlarged at base; orange to bright rusty-cinnamon
becoming dark brown in age, with a large mat of pine needles and debris usually stuck to
the base. SPORE PRINT brown; spores 5.5-7.5 * 5-6 microns, nearly round, promi¬
nently warted.
HABITAT: Solitary oringroups onground underpinesandotherconifers, sometimes also
in fused clusters; widely distributed. It is one of the two most common Hydnellums in our
area (from late fall through early spring), but does not often occur in large numbers.
EDIBILITY: Unequivocally inedible.
COMMENTS: The tough texture, knobby or lumpy cap surface, and bright orange to
rusty-cinnamon colors of the cap and flesh are the fallible fieldmarks of this species and its
close relatives. It never exudes the red droplets characteristic of H. peckii, but might be
casually mistaken for a polypore such as Phaeolus schweinitzii. However, a close look
at its underside reveals the presence of small “teeth” instead of pores. Closely related
species include: H. complectipes, usually with many caps fused to form large, complicated
masses or rosettes, found in the Pacific Northwest under conifers;//, conigenum, with very
thin flesh in the cap, also found under conifers; and H.ferrugipes and H. earlianum, found
under hardwoods in eastern North America, the latter with a nearly smooth cap.

626
HYDNELLUM 627

Hydnellum peckii Color Plate 160

(Strawberries and Cream; Bleeding Hydnellum)
CAP 2.5-15 cm broad, often with needles and other debris incorporated into it; top-shaped
becoming broadly convex to plane or finally depressed; surface felty or velvety and white
to pink in young specimens or on actively-growing margin; in age becoming nearly hairless
and lumpy or jagged with projecting nodules, and often ridged and/or pitted; darkening
to brown, dark brown, or vinaceous-brown from the center outward and almost entirely
these colors in old age; beaded with or exuding bright ruby-red to dark red droplets when
fresh and moist. Flesh in both cap and stalk tough and fibrous-corky, faintly zoned;
pinkish-buff to cinnamon-brown, dark reddish-brown, or dingy brown; odor mild to
fragrant or pungent; taste typically extremely acrid (peppery). SPINES rather short (1-6
mm long), dull pinkish becoming brown or purplish-brown, often with paler tips; some¬
times decurrent. STALK 0.5-7.5 cm long, 1-2 (3) cm thick, central or off-center, equal or
tapered below and sometimes rooting, or occasionally swollen at base; felty or velvety
and colored more or less like cap or darker; solid, tough or woody. SPORE PRINT brown;
spores 4.5-5.5 * 3.5-4.5 microns, round or nearly round, prominently warted.
HABITAT: S olitary to scattered, gregarious, or in fused clusters on ground under conifers;
widely distributed, but particularly common in the Pacific Northwest in the late summer
and fall. I’ve recorded only one questionable collection from our area, but have seen it
farther north under pine and fir.
EDIBILITY: Indubitably inedible due to the burning-acrid taste and tough, corky texture.
COMMENTS: The bright red droplets that cling to the surface of the cap in moist weather
make this a striking and easily-identified mushroom (see color plate!). As in other Hyd-
nellums, the cap varies considerably in color and texture according to age and environ¬
mental conditions. When young, white, and beaded with droplets it looks like a Danish
pastry topped with strawberry jam. Older, battered specimens, on the other hand, are
scarcely recognizable and easily confused with other species. Intermediate stages are
typically brown or dark reddish-brown with a white to pink, beaded margin. Other species
exuding red droplets in wet weather include: H. diabolum, very similar if not the same,
with a hairier cap at maturity and stronger odor; and H. pineticola, common under pines
in northeastern North America, with a rather unpleasant but not acrid taste and a cap that
sometimes has pink droplets. See also H. scrobiculatum.

Hydnellum scrobiculatum (Rough Hydnellum)

CAP 3-10 cm broad, more or less top-shaped when young becoming plane to depressed in
age, sometimes with smaller caps on top; surface usually roughened and irregular from
numerous pits and projecting warts or blunt spikes, often also with radial ridges; pallid
to pale salmon-buff or pinkish and plushlike or felty when young or on growing margin of
older specimens; darkening in age from the center outward to buffy-brown, then dull
cinnamon and finally darker brown, but lackingconspicuousconcentriczones; sometimes
beaded with dark red droplets when young and moist, and bruising reddish-black to black
when rubbed, especially at margin. Flesh zoned, sometimes also with white dots; usually
duplex, the upper layer spongy when fresh and colored like cap, the lower layer or core
brown to dark reddish-brown; often exuding dark red juice when squeezed (if moist and
fresh); odor and taste mild to farinaceous. SPINES short (1 -5 mm long), usually decurrent,
sometimes fused; pallid or colored like cap margin, becoming buffy-brown to cinnamon-
brown and finally purple- or dark brown in old age. STALK 1-4 cm long, 0.3-1.5 cm thick,
usually tapered downward but often with a swollen, buried, spongy base; central or off-
center; tough; cinnamon-brown or colored more or less like cap. SPORE PRINT
brownish; 4.5-5.5 (7) * 3.5-5 microns, elliptical to nearly round, prominently warted.
628 HYDNACEAE

HABITAT: Scattered to gregarious or in fused clusters in woods, usually associated

with conifers; widely distributed and sporadically common in some regions. It can be found
nearly every fall in the Pacific Northwest and extends into northern California, but occurs
very rarely if at all in our area.
EDIBILITY: Incomparably inedible.
COMMENTS: This species is rather hard to characterize, as evidenced by the lengthy
description. It is best recognized by its irregularly roughened cap with pits, radial ridges,
and spikelike projections, plus the overall cinnamon to brown color with a pinkish growing
margin that darkens when bruised. When beaded with red droplets it can be confused with
H. peckii, but is not as peppery-tasting as that species and usually has a buried, swollen
“tuber” on the stalk. When older it can be mistaken for various other brownish to reddish
Hydnellums. H. zonatum (-H. scrobiculatum var. zonatum, H. concrescens— COLOR
PLATE 157) is a closely related, similarly colored species with a smooth or radially cor¬
rugated (but not lumpy) cap that is usually zoned concentrically (at least somewhat). It is
quite common under hardwoods in eastern North America but also occurs on the west
coast both in its normal form and a diminutive one whose cap is 4 cm broad or less. Both
H. zonatum and H. scrobiculatum are quite distinct in their typical forms, but appear to
intergrade. Other similar species: H. subsuccosum resembles H. zonatum, but is more
irregular in shape and juicier when fresh, and exudes a pinkish to reddish juice when
squeezed; it also has yellowish-gray to yellow-green mycelium at the base of the stem and its
cap is often flecked with yellowish spots or particles. H. cumulatum has many caps built on
top of one another, and little or no stalk. Both of these occur under conifers.

Phellodon tomentosus (Zoned Phellodon)

CAP 1.5-4 (5.5) cm broad, often fused with others, plane to depressed or broadly funnel-
shaped; surface dry, smooth to ridged or corrugated, minutely hairy (tomentose), white
when very young, soon becoming concentrically zoned with yellow-brown, cinnamon-
brown, darker brown, etc., the growing margin usually remaining white and felty to the
touch, but bruising brownish. Flesh thin, leathery and fibrous, brownish (darker brown
in stalk); odor usually fragrant (like fenugreek); taste mild or slightly bitter. SPINES short
(1-3 (5) mm), crowded, delicate, whitish becoming pale cinnamontobrownwithpalertips;
slightly decurrent. STALK 1-5 cm long, 2-5 (8) mm thick, usually central, equal ortapering
downward, colored more or less like cap, arising from spongy pad of brownish mycelium.
SPORE PRINT white; spores 3-4.5 microns, round or nearly round, minutely spiny.
HABITAT: Scattered to densely gregarious or clustered (several caps often fused together
but the stalks usually separate) under conifers; widely distributed and fairly common. I
have found it in Mendocino County, California, in the fall and early winter, and it is some¬
times abundant in the Pacific Northwest; just how far south it occurs is unclear.
EDIBILITY: Unknown, but too small and too tough to be of value.
COMMENTS: The small size, beautifully zoned yellow-brown to cinnamon-brown to
dark brown cap, brown flesh, slender stem, and frequently fragrant odor are good field-
marks. It bears an uncanny resemblance to polypores of the genus Coltricia (see C.
cinnamomea), but the underside of the cap features minute spines or “teeth” instead of
pores. Hydnellum zonatum (see comments under H. scrobiculatum) is also somewhat
similar, but has darker spines and brown spores. Other species: P. confluens is quite
similar, but more common under hardwoods. It is more irregular in shape, with an often
roughened or pitted, whitish cap that darkens to creamy or dark tan from the center
outward, and spines which are grayish at maturity.
PHELLODON 629

Phellodon atratus (Blue-Black Phellodon)

CAP 1-5 cm broad but often fused with others; plane to depressed or irregular; surface dry,
usually at least faintly zoned concentrically; bluish-black to purple-black or black, the
margin often slightly paler or purpler. Flesh in both cap and stalk purple-black to bluish-
black; thin, tough, fibrous, pliant, sometimes with a thin outer or upper spongy layer; odor
mild or faintly fragrant; taste mild. SPINES very short (1-2 mm), irregularly decurrent;
gray to dark purplish-gray-brown, darker where bruised. STALK 2-5 cm long, 3-5 mm
thick, usually central, sometimes compound or branched; tapering downward but usually
thickened at ground level by a felty mycelial layer; rough, often flattened, colored more or
less like cap. SPORE PRINT white; spores 4-5 x 3-5 microns, round or nearly round,
minutely spiny. Cap tissue staining blue-black in potassium hydroxide (KOH).
HABITAT: Scattered to gregarious, often forming compound or fused clusters, on
ground under conifers (particularly Sitka spruce); apparently endemic to the Pacific
Northwest and California and quite common in the fall and winter in second-growth
forests. It has been found in Big Basin State Park, but is rare south of San Francisco.
EDIBILITY: Unknown, but like myself, too tough and too small to be of value.
COMMENTS: The small size, tough texture, and bluish-black color distinguish this
conifer-lover. Hydnum fuscoindicum is somewhat similar in color but larger and fleshy-
brittle rather than pliant and tough. A closely related species, P. melaleucus, has purplish-
black to purple-gray flesh, but its cap is dark brown to purplish-gray with a pallid margin,
the spines are whitish to gray, and the stalk is very thin, dark brown to black, and sometimes
deeply rooted. It also occurs under conifers but has a wider (albeit northern) distribution.
In eastern North America a similar species, P. niger, is common. It is larger and thicker
than P. atratus, with a white to brownish, gray, or black cap and black flesh (in both the
cap and stem). Hydnellum nigellum, another eastern species, is small and slate-gray to
black, but has brown spores.

AURISCALPIUM
THIS genus contains a single odd species with a worldwide distribution. It is not closely
related to other stalked teeth fungi. In fact, microscopic characters suggest a possible
relationship to the agaric genus Lentinellus.

Auriscalpium vulgare (Ear Pick Fungus)

CAP 1-2 (4) cm broad, more or less kidney-shaped in outline, broadly convex to plane or
slightly depressed; surface dry, covered with dense fibrils or hairs, brown to dark brown,
sometimes blackish in age; margin often fringed and paler. Flesh thin, tough, pliant, white
to pale brown. SPINES whitish to flesh-colored, sometimes darkening to brown; very
fine and crowded, short (1-3 mm long). STALK 2-1 Ocm long, 0.5-3 mm thick, very slender,
equal or slightly enlarged below, usually attached to side of cap (lateral), densely hairy,
especially toward base, rusty-brown to dark brown or blackish. SPORE PRINT white;
spores 4.5-6 x 3-3.5 microns, round or nearly round, smooth or minutely spiny, amyloid.
HABITAT: Solitary or in twos and threes on rotting, often buried cones of conifers, or
sometimes on thick mats of debris made up partly of decaying cones; widely distributed,
but rare in our area (or else frequently overlooked). In my experience it favors Douglas-
fir cones, at least on the west coast.
EDIBILITY: Much too small and much too tough to be of value.
Auriscalpium vulgare. This dainty fungus grows on decaying cones (in this case, Douglas-fir). Note
the slender stalk and layer of spines on underside of cap.

COMMENTS: The small size and growth on decaying cones plus the thin, hairy, lateral
stem and fine spines that line the underside of the cap are the forthright fieldmarks of
this unique, petite fungus. The stem, though lateral, may occasionally appear to be central
when the cap is deeply indented. The stem is usually longer than the width of the cap and
far thinner than that of any other fungus with spines. The dark color and small size make
it very difficult to see unless you are specifically looking for it.

Coral and Club Fungi

CLAVARIACEAE
THIS large and lovely group of fleshy fungi includes simple, unbranched, upright clubs
and fleshy, intricately branched, coral-like forms. With the exception of Clavariadelphus
the fruiting body is not differentiated into an upper sterile surface (cap) and fertile under¬
side. Instead the spore-bearing basidia line the smooth to occasionally wrinkled surfaces of
the upright clubs or branches. A sterile base, stalk, or “trunk” is normally present, however.
Coral fungi are a conspicuous and colorful component of our woodland fungi. They
come in every imaginable color, and some of the larger branched forms (notably Sparassis)
are edible. They are difficult from a taxonomic standpoint. Nearly all the coral fungi
were originally lumped together in one unwieldy genus, Clavaria, but now more than 30
genera are recognized. These are delimited largely on microscopic and chemical charac¬
teristics (e.g., whether or not the spore-bearing surface stains green in ferrous sulfate), so
to facilitate identification the family has been divided into five groups, keyed below. The
largest and most common group, Ramaria, is microscopically similar to Gomphus of the
chanterelle family (Cantharellaceae), and some taxonomists place them together in the
family Gomphaceae in the belief that they arose from a common ancestor. Another genus,
Sparassis, is usually placed in a family of its own, but is traditionally grouped with the coral
fungi because of its branched fruiting body.

Key to the Clavariaceae

1. Fruiting body unbranched or very sparsely branched (but often tufted or clustered) ... 2
1. Fruiting body profusely branched from a stalk or common base .4

630
Left: Clavaria vermicularis has unbranched but clustered fruiting bodies (it is also shown on p. 637).
Right: Ramaria stricta, one of many species with a branched fruiting body.

2. Fruiting body entirely brownish-black to black or blackish beneath a white powdery coating or
entirely green to olive or blue-green or interior with large chambers or compartments or para¬
sitic on insects, spiders, or truffles; spores borne asexually or inasci (see Ascomycetes, p. 782)
2. Not as above (may be white, but if so then not powdery); spores borne on basidia . 3
3. Fruiting body tough, the flesh pithy, stringy, or punky; apex often enlarged; usually 7 mm thick
or more .Clavariadelphus, p. 632
3. Fruiting body typically fragile or if tough then much smaller; mostly less than 7 mm thick; apex
acute or blunt or occasionally enlarged . Clavaria & Allies, p. 634
4. Fruiting body small and tough with very thin, almost hairlike branches, brown to grayish-brown
to dark brown or purple-brown; growing on twigs, needles, etc.; rare (mostly tropical) Pterula
4. Not as above; common . 5
5. Fruiting body consisting of numerous flattened, wavy, ribbonlike, or leafy segments or lobes
arising from a common base; rather tough; overall color white to creamy, yellowish, or tan;
growing at or near the bases of trees and stumps . 6
5. Not as above . 7
6. Found on hardwood stumps or roots in tropics and along Gulf Coast; fertile surface usually
developing pores, spines, or “teeth” in age . . . (seePolyporus, Albatrellus, & Allies, p. 554)
6. Not as above; common and widespread .Sparassis, p. 657
7. Fruiting body bright yellow to orange; spore print white, or if not then branches usually viscid;
spores smooth; typically growing on wood . Clavaria & Allies, p. 634
7. Not as above . 8
8. Branch tips crownlike (in the form of small fringed cups); spore print white; growing on wood
. Clavulina& Allies, p. 640
8. Not as above .9
9. Fruiting body bright yellow to orange when fresh and small (typically 2-7 (10) cm high) . . 10
9. Not as above . 11
10. Stalk slender and not particularly fleshy; branches often hollow and/or viscid; spore print
white or yellowish; extensive mycelial mat typically absent . Clavaria & Allies, p. 634
10. N ot as above; stalk thick and fleshy or if not, then an extensive mat of mycelial threads usually
present at base and in substrate; spore print buff to tan, yellowish, or ochre Ramaria, p. 645
11. Branches tough, usually flattened, grayish-brown to dark brown to purple-brown (but tips
often pallid when growing); odor typically garliclike or fetid (see Thelephorapalmata, p. 609)
11. Not as above . 12

631
632 CLAVARIACEAE

12. Spore print creamy to yellow, tan, yellow-orange, or ochraceous (rarely white); fruiting body
medium-sized to fairly large, often brightly colored, or if dull colored then usually with a large
fleshy base (stalk); fertile surfaces staining greenish to blue in ferrous sulfate Ramaria, p. 645
12. Spore print typically white; fruiting body rather small to medium-sized, white or dull-colored
(grayish, brownish, tan, bluish-gray, or tinged purple); base typically not large and fleshy; fertile
surfaces typically not staining green or blue in ferrous sulfate . . . Clavulina & Allies, p. 640

CLAVARIADELPHUS (Club Corals)

Medium-sized, terrestrial, woodland fungi. FRUITING BODY erect, unbranched or occasionally
forked, more or less club-shaped or with a flattened top, usually at least 5 mm thick; surface smooth
to wrinkled. Flesh rather tough and fibrous or pithy. SPORE PRINT white to pale yellow, buff, or
ochre. Spores typically elliptical, smooth. Spore-bearing surface staining green in ferrous sulfate.

THESE are rather tough, club-shaped fungi with a smooth or somewhat wrinkled spore¬
bearing surface. They are larger and thicker than most fairy clubs (Clavaria & Allies)
and not nearly so fragile. In C. truncatus and its close relatives the apex of the club
is flattened and sterile—in other words, a rudimentary cap—but in the other species it is
typically rounded, pointed, or only slightly flattened.
Club corals are harmless but rather tough, stringy, and/ or bitter-tasting. They grow
only in the woods and in our area fruit mostly during cold weather. Three widespread
species are described here. If your “club coral” is small and irregularly shaped, check the
earth tongues (on p. 865) as well as Clavaria & Allies (p. 634).

Key to Clavariadelphus
1. Fruiting body with a consistently flattened (truncate) or depressed apex or even a rudimentary
cap; associated with conifers . 2
1. Not as above; apex of fruiting body rounded to obtuse or pointed, or if sometimes flattened
then associated with hardwoods .4
2. Upper portion of fruiting body red to reddish-orange C. lovejoyae (see C. truncatus, p. 634)
2. Not as above; upper portion of fruiting body orange to yellow, ochre, etc.3
3. Spore print pale ochre .C. truncatus, p. 634
3. Spore print white.C. borealis (see C. truncatus, p. 634)
4. Apex of fruiting body usually with a sharply defined point or “nipple”; associated with conifers
. C. mucronatus (see C. ligula, p. 633)
4. Not as above; apex rounded to bluntly pointed or somewhat flattened .5
5. Associated mainly with hardwoods; mature fruiting body 1-3 cm thick and 6-20 cm or more
high .C. pistillaris & others, below
5. Associated with conifers; mature fruiting body up to 1.5 cm thick and typically 2-10 cm high
.C. ligula & others, p. 633

Clavariadelphus pistillaris (Common Club Coral)

FRUITING BODY simple, erect, unbranched or sometimes forked; club-shaped or
tapering downward, the apex rounded or somewhat flattened but not normally depressed;
6-20 (30) cm high, 0.8-4 (6) cm broad; surface smooth or often longitudinally wrinkled or
grooved in age; usually pallid at first, but soon dull pinkish-brown to reddish-brown, flesh-
colored, or ochraceous-brown; staining brown to vinaceous-brown when handled or
bruised; apex often yellowish at first but soon colored like the rest of the fruiting body;
base usually pallid, with white hairs. Flesh tough, fibrous or pithy, whitish, bruising brown;
taste mild or bitter. SPORE PRINT white or tinged yellow; spores 9-16 * 5-10 microns,
elliptical, smooth.
Clavariadelphus pistillaris is a common terrestrial club-shaped species. In our area it favors oak.
Forked specimen at top is not unusual.

HABITAT: Solitary, scattered, or in groups on ground under hardwoods and in mixed

woods; widely distributed. It is common in our area from the late fall through early spring,
especially under live oak, tanoak, and madrone.

EDIBILITY: Harmless. The taste and texture are reminiscent of stale rope.
COMMENTS: The ochre-brown to flesh-colored, club-shaped fruiting body that stains
brown when handled is characteristic of this cosmopolitan club coral. It is the commonest
Clavariadelphus in our area and the only one found under hardwoods. Its apex is some¬
times quite broad, but not as flagrantly flattened as that of C. truncatus. Other species:
C. subfastigiatus is a brownish-orange species found under conifers in northern California
and elsewhere; it does not discolor as much when handled and turns bright green in
potassium hydroxide (KOH).

Clavariadelphus ligula (Strap Coral) Color Plate 171

FRUITING BODY simple, erect, unbranched or rarely forked, cylindrical to flattened-
cylindrical or club-shaped, the apex usually rounded or bluntly pointed; 2-10 cm high and
0.3-1 (1.5) cm broad at apex. Surface smooth to slightly wrinkled, dull-colored (buff to
dull yellowish, ochre-buff, pale reddish-brown, or vinaceous-buff); base whitish and hairy,
often with white mycelial threads penetrating the surrounding humus. Flesh white, pithy
but tough; taste mild or bitter. SPORE PRINT white to pale yellowish; spores 8-18 * 3-6
microns, elongated-elliptical, smooth.
HABITAT: Scattered to densely gregarious or tufted in humus under conifers; widely
distributed. It is common throughout much of the West in the summer and fall, but
absent or very rare in our area.
EDIBILITY: Worthless.
COMMENTS: This species is smaller and slimmer than C. pistillaris and typically occurs
under conifers rather than hardwoods—often in large troops (see color plate). The apex of
the club is not noticeably flattened or depressed as in C. truncatus, and is not brightly
colored. Other species: C. sachalinensis is a macroscopically identical species with larger,
buff to ochraceous spores; it also grows gregariously in humus under conifers. Another
conifer-lover, C. mucronatus, differs in its whitish, sharply nippled apex. Still another,
C. subfastigiatus, is somewhat thicker and stains green when touched with potassium
hydroxide (KOH). None of these are worth eating.
634 CLAVARIACEAE

Clavariadelphus truncatus (Truncate Club Coral) Color Plate 166

FRUITING BODY simple, erect, unbranched or occasionally forked; club-shaped or
more often with a broadly flattened or depressed apex (a rudimentary cap); 5-15 (18) cm
high and 2.5-8 cm broad at apex. Surface smooth or often wrinkled or veined (especially
near apex), more or less pinkish-brown to ochre or brownish-orange, the apex usually
brighter (yellow to golden-yellow or yellow-orange), at least when young; base often pallid,
with white hairs. Flesh rather tough or pithy, white to ochre; taste mild to sweetish or
bittersweet. SPORE PRINT pale ochre; spores 9-13 * 5-8 microns, elliptical, smooth.
HABITAT: Scattered to gregarious in duff under conifers; widely distributed, but not
nearly as common in our area as C. pistillaris. I have seen large fruitings of this species and
its look-alike, C. borealis (see comments), in the fall and winter in northern California and
in the summer in the Rocky Mountains.
EDIBILITY: Edible and delicious when sweet. It can be sauteed by itself and served for
dessert!
COMMENTS: The broad, golden, flattened top distinguishes this species from its cousins
C. pistillaris and C. ligula. The color of the fruiting body actually varies considerably from
dull pinkish-brown to bright golden-orange, and the apex can be quite inflated so as to
resemble a chanterelle (Gomphus or Cantharellus). Other species: C. borealis is a white-
spored version of C. truncatus that sometimes shows a lilac tinge to the fruiting body; it is
common and widespread under conifers. C. lovejoyae of the Rocky Mountains is also
similar, but red to reddish-orange in color.

CLAVARIA & Allies (Fairy Clubs)

Small, mostly fragile fungi found on ground, leaves, or occasionally on wood. FRUITING BODY
erect, usually unbranched or sparingly branched and finger-shaped or clublike, slender, often tufted
or clustered. Flesh usually fragile. SPORE PRINT white. Spores typically smooth. Basidia typically
4-spored. Clamp connections typically absent (Clavaria), present (Clavulinopsis). Fertile surface
often staining green in ferrous sulfate (Clavulinopsis), or not greening (Clavaria & others).

THESE are primitive fungi with an erect, relatively unspecialized fruiting body. The most
common forms are unbranched and smaller, slimmer, and frailer than the club corals
(Clavariadelphus). They are sometimes confused with earth tongues—which are tougher,
often flattened and velvety, and capitate (with a cap distinct from the stalk)—and which
belong to an entirely different group of fungi, the Ascomycetes. Fairy clubs often grow in
clumps, but the individual clubs do not usually arise from a fleshy base as in the branched
corals (Clavulina, Ramaria, etc.). The few branched species are not as fleshy as Ramarias
and usually more vividly colored than Clavulina, Ramariopsis, and Clavicorona.
Clavaria and Clavulinopsis are the two most common genera of fairy clubs, but their
defining features are esoteric, involving the presence or absence of carotenoid pigments,
clamp connections, and the behavior of nuclei in the basidia. For the sake of convenience
they are treated together here, along with several other small, miscellaneous genera (Multi-
clavula, Macrotyphula, and Typhula).
Fairy clubs are too small and fragile to have any food value, but they are an attractive
addition to our woodland decor. They are saprophytic on humus, soil, or occasionally grass
and decaying wood. In our area they fruit—as do most of the Clavariaceae—from late fall
through early spring. Seven species are described here.
CLAY ARIA & ALLIES 635

Key to Clavaria& Allies

1. Growing on algae-covered wood or soil; fruiting body minute (up to 1.5 cm high and 1 -3 mm
thick) .Multiclavula mucida & others, p. 636
1. Not as above (if growing on algae, then larger) .2
2. Fresh fruiting body yellow to orange, red, salmon, or pink . 3
2. Not as above (fruiting body white, yellow-brown, grayish, purple, etc.) . 10
3. Fresh fruiting body yellow to orange .4
3. Fresh fruiting body rose-pink to red or orange-red .9
4. Fruiting body branched . 5
4. Fruiting body unbranched or occasionally forked (but often clustered) .6
5. Usually growing on or near wood, the base often deeply rooted; branches rather tough and
usually somewhat viscid .(see Calocera viscosa, p. 674)
5. Not as above; growing on ground or wood; not viscid .... Clavulinopsis corniculata, p. 639
6. Fruiting body with a wide, often flattened head and/or fruiting body often irregular in shape;
texture rather tough; spores borne inside asci .(see Helotiales, p. 865)
6. Not as above; fruiting body typically clublike to spindle-shaped or fingerlike or rarely forked;
usually rather fragile; spores borne on basidia . 7
7. Fruiting bodies 5-15 cm tall, usually growing in bundles or large clusters, yellow .
.Clavulinopsisfusiformis(see C. laeticolor, p.638)
7. Fruiting bodies up to 6.5 (10) cm high, solitary to gregarious or tufted; yellow to orange .. 8
8. Fruiting body less than 15 mm high, usually somewhat viscid; typically growing on or near
wood . (see Calocera cornea under C. viscosa, p. 674)
8. Not as above; usually growing on ground .Clavulinopsis laeticolor & others, p. 638
9. Fruiting body pink to rose-colored when fresh; found mainly in eastern North Ameica; rare .
.Clavaria rosea (see Clavulinopsis laeticolor, p. 638)
9. Fruiting body orange to orange-red or red; widespread but rare .
.Clavulinopsis aurantio-cinnabarina & others (see C. laeticolor, p 638)
10. Fruiting body very thin, arising from a small beadlike body (sclerotium) .
.Typhula spp.(see Macrotyphula juncea, p 636)
10. Not as above; fruiting body not very thin, or if so then not attached to a beadlike body . 11
11. Fruiting body very thin (up to 2 mm), 3-10 cm tall; pallid to yellowish or brown .
.Macrotyphula juncea, p. 636
11. Not as above . 12
12. Fruiting body lavender to purple, deep purple, or grayish-purple, at least when fresh 13
12. Fruiting body differently colored . 15
13. Fruiting bodies unbranched but often clustered; brittle or fragile; found mostly under northern
and mountain conifers . Clavaria purpurea, p. 637
13. Not as above . 14
14. Fruiting body sparingly or much-branched (the branching often dichotomous); basidia 4-
spored; found in eastern North America . Clavaria zollingeri (see C. purpurea, p. 637)
14. Fruiting body much-branched; basidia 2-spored; widespread (see Clavulina & Allies, p. 640)

15. Fruiting body branched .Clavulinopsis umbrinella & others (see C. corniculata, p. 639)
15. Fruiting body unbranched or occasionally branched very sparingly (but often growing in
tufts or clusters) . 16
16. Fruiting body small (up to 2 cm tall), white, the apex broadly enlarged; growing on needles,
twigs, etc. Clavicorona taxophila(see Multiclavula mucida, p. 636)
16. Not as above . 17
17. Fruiting bodies very fragile, crumbling easily, pure white to translucent white or yellow-stained,
not prominently wrinkled; tip acute or sometimes blunt but not broadly enlarged; often
growing in tufts or clusters; very common .Clavaria vermicularis, p. 637
17. Not with above features . 18
18. Fruiting body very long (tall) and narrow (7-30 cm high; 2-8 mm thick); yellowish to brown;
not normally growing in clusters .Macrotyphula fistulosa(see M. juncea, p. 636)
18. Not as above . 19
636 CLAY ARI ACE AE

19. Growing on wood; fruiting body more or less clublike, the surface often roughened, 1-4 cm high
and 0.5-1 cm thick; spores borne inside asci .(seePodostroma alutaceum, p. 879)
19. Not as above; usually growing on ground . 20
20. Fruiting body 0.3-1.5 cm thick, texture tough; flesh white and pithy or stringy; color variable:
buff to yellowish or some shade of brown, but not white orgray; typically growinginduff under
conifers, often in troops .(see Clavariadelphus, p. 632)
20. Fruiting body usually 2-7 mm thick and/or differently colored, usually fragile.21
21. Fruiting body white and sparingly branched; branches thick, hollow, blunt, somewhat gelati¬
nous; found under hardwoods in eastern North America .. (see Tremellales& Allies, p. 669)
21. Not as above .22
22. Fruiting body fragile, usually tufted or clustered, grayish to yellowish-gray to pinkish-gray
or dingy flesh-colored; basidia typically 4-spored . Clavaria fumosa & others, p. 638
22. Not as above; fruiting body tough, or if fragile then differently colored (white or tinged gray or
buff); basidia typically 2-spored .(see Clavulina & Allies, p. 640)

Multiclavula mucida (Scum-Lover)

FRUITING BODY simple, unbranched or sometimes forked, erect; small (only 0.5-1.5
cm high and 1 -2 mm thick); cylindrical or tapered; surface smooth, white to creamy, buff,
or yellowish, but often aging salmon to brick-red. Flesh rather waxy and tough, pliant,
white. SPORE PRINT white; spores 4.5-7.5 * 2-3 microns, oblong to elliptical, smooth.
Cystidia absent.
HABITAT: In groups—but not clusters—on wet algae-covered wood or occasionally on
soil; northern in distribution. I have seen it in our area in the fall and winter, but it is in¬
conspicuous because of its small size.
EDIBILITY: Utterly inconsequential.
COMMENTS: The small size and association with green algae typify the genus Multi¬
clavula, and the whitish color of the fresh fruiting body and tendency to grow on wood are
characteristic of this species. Clavaria mucida is an older name for it. Other species:
M. vernalis(-Clavariaphycophila) is similar but pale orange, has cystidia, and grows on
algae-covered soil; Clavicorona taxophila is small and whitish but has a widened, flattened
apex and grows on twigs, needles, and other debris rather than with scum. All of these are
smaller than Clavaria vermicularis and shorter than Macrotyphula juncea.

Macrotyphula juncea (Fairy Hair)

FRUITING BODY simple, unbranched or rarely forked, erect, very thin (up to 2 mm
thick), 3-10 cm high when mature; cylindrical or tapering upward; leather-colored to
yellowish-buff or pallid, smooth; tip acute or in age sometimes blunt; base somewhat
fibrillose and often creeping horizontally, often with large whitish mycelial threads(rhizo-
morphs) attached. Flesh very thin; taste sometimes acrid. SPORE PRINT white; spores
6-12 x 3.5-5.5 microns, elliptical or almond-shaped, smooth.
HABITAT: Scattered to gregarious in humus and leaf litter, on rotting twigs, etc.; widely
distributed. It occurs in our area on oak and tanoak leaves and redwood needles, but is easy
to overlook. It is fairly common in the fall and winter, especially along streams and in other
dank places.
EDIBILITY: Utterly irrelevant—a couple hundred would be needed for a mouthful!
COMMENTS: This species is so thin that it can be mistaken for the bare stem of an herba¬
ceous plant or a small agaric that has lost its cap. Its slimness alone separates it from our
other common coral and club fungi. Its generic disposition is problematic—some authors
place it in Typhula (other Typhula species are similar in size and shape but arise from a
Left: Macrotyphula juncea, a very thin species. Right: Fragile white clusters of Clavaria vermicularis
look like bunches of bean sprouts. See p. 631 for a photo of younger specimens.

small beadlike body or sclerotium); it has also been placed in Clavaria and Clavariadelphus
and may eventually merit a genus of its own. Other species: M. fistulosa (-Clavaria¬
delphus fistulosus) has a very long (7-30 cm), slender (2-8 mm), hollow fruiting body
that is yellowish to brownish; it grows on dead sticks and debris, especially of alder.

Clavaria vermicularis (Fairy Fingers)

FRUITING BODY simple, unbranched (but usually clustered) or rarely forked at the tip,
erect or often curved, slender, soon withering; 3-12(15) cm tall, 3-5 mm thick; surface pure
white to translucent white or often stained yellow and yellowing in age from the tip down¬
ward; cylindrical or flattened somewhat, smooth or sometimes grooved, usually tapered
toward the tip, which is acute or sometimes blunt and often discolored. Flesh thin, white,
very brittle or fragile. SPORE PRINT white; spores 5-7 * 3-4 microns, elliptical to nearly
round, smooth.
HABITAT: I n tufts, clusters, or groups, often with solitary fruiting bodies interspersed, on
ground in woods or grassy places; widely distributed. Common in our area throughout the
mushroom season, but most prevalent in December and January, especially in dank
wooded areas.
EDIBILITY: Edible but insubstantial; the fragile watery flesh has little or no flavor and
dissolves when chewed.
COMMENTS: The distinctive clumps of slender white “fingers” make this species most
attractive. It is by far our most common Clavaria, growing wherever moisture is sufficient.
It is not as deeply wrinkled as C. fumosa or Clavulina rugosa (see comments under the
Clavulina cristata group), and it crumbles seemingly without provocation.

Clavaria purpurea (Purple Fairy Club) Color Plate 167

FRUITING BODY simple, unbranched but often clustered, erect, 2.5-12 cm tall, 2-6 mm
thick; grayish-purple to deep purple or purple when fresh, fading as it ages to lavender-gray,
lavender-buff, purple-brown, smoky-brown, etc.; cylindrical or tapered; tip acute or blunt;
base usually paler and/or with white hairs. Flesh white or purplish, brittle. SPORE
PRINT white; spores 5.5-9 * 3-5 microns, elliptical to oblong, smooth. Cystidia inter¬
mingled with basidia.

637
638 CLAVARIACEAE

HABITAT: Scattered to densely gregarious (often in tufts or clusters) on ground in wet

areas, usually under or near conifers; common during the summer under spruce and fir
in the Rocky Mountains, also found in the Pacific Northwest and California (but not in
our area). I have seen large fruitings in New Mexico.
EDIBILITY: Edible, but thin-fleshed and fragile.
COMMENTS: The purple color immediately distinguishes this beautiful species from
other fairy clubs. Clavaria (-Clavulina) zollingeri is a sparingly to profusely branched
purple species occasionally found in eastern North America. For other branched purple
coral fungi, see Clavulina cinerea, C. amethystina, and Ramariopsis pulchella (under
C. cinerea) and Ramaria fumigata.

Clavaria fumosa (Grayish Fairy Club)

FRUITING BODY simple, unbranched (but usually clustered), erect, 3-10 (14) cm tall,
2-7 mm thick; surface grayish to yellowish-gray (color variable), usually grooved or
wrinkled longitudinally, often somewhat flattened or twisted and/ or hollow in age; tip
usually blunt, sometimes brownish; base often paler or whitish. Flesh brittle, whitish.
SPORE PRINT white; spores 5-8 * 3-4 microns, elliptical, smooth.
HABITAT: In tufts or clusters onground in woods or grassy areas; widely distributed but
not common. I have found it several times under oak and madroneinthe winterand spring.
EDIBILITY: Harmless, fleshless, flavorless.
COMMENTS: The gray to yellowish-gray (or occasionally brownish-gray) color, fre¬
quently wrinkled surface, and clustered but unbranched fruiting bodies typify this forget¬
table fairy club. Other species: C. rubicundula is just as forgettable but more fragile, and
usually slightly pinker (dingy flesh-colored to pinkish-gray to vinaceous-buff).

Clavulinopsis laeticolor (Golden Fairy Club)

FRUITING BODY simple (unbranched or occasionally forked once) but often tufted;
erect, small, cylindrical or often somewhat flattened, grooved, and/ or twisted; 1.5-6.5(10)
cm tall but usually about 34 cm, 1-5 (10) mm thick; surface bright orange toyellow(some-
times yellower below) to yellow-ochre or in some forms orange-red, often duller(yellow-
ish-buff) as it dries or fades; extreme base whitish; tip usually acute, often brownish in age
or when dry. Flesh thin, somewhat pliant, pallid or yellowish; odor and taste mild.
SPORE PRINT white; spores 4.5-7 (9) * 3.5-5.5 (6.5) microns, broadly elliptical to nearly
round to triangular or pear-shaped, smooth, prominently apiculate.
HABITAT: Solitary, scattered, tufted, or in groups on mossy banks, wet soil, and wood¬
land humus; widespread and common, fruiting in our area in the fall, winter, and spring.
EDIBILITY: Inconsequential.
COMMENTS: Also known as Clavariapulchra, this dainty fairy club can be distinguished
from all but a few close relatives (see below) by its bright yellow to orange or sometimes
reddish-orange color. It is not viscid like the coral-like jelly fungi (Calocera), and does
not usually grow on wood. Certain earth tongues (e.g., Microglossum rufum and Neo-
lecta irregularis) are similarly colored, but have more enlarged fertile “heads” and bear
their spores in asci rather than on basidia. There are a number of closely related, brightly
colored fairy clubs, including: Clavulinopsis helvola, with anguar-warty spores; C.fusi-
formis, taller (5-15 cm) and bright yellow with round spores, usually found in bundles in
grass or humus (common in eastern North American, also found in California); C.
gracillima (-C. luteoalba), with a well-defined sterile base or “stalk”; C. appalachiensis,
creamy to creamy-yellow with a deep ochre “stalk”; C. miniata, pale pinkish-orange or
Two bright yellow to orange club fungi: Clavulina laeticolor (left) grows singly or in small tufts;
Clavulinafusiformis (right) is usually taller, slimmer, and clustered.

apricot-colored; C. aurantio-cinnabarina, a rare but widely distributed striking blood-

red to orange or pinkish-orange species with only slightly apiculate spores; and C. sub-
australis, with a delicate pink fertile portion and dark yellow to orange “stalk.” Finally,
there is Clavaria rosea, a beautiful rose-pink species that occurs occasionally in eastern
North America but hasn’t yet been found in California. None of these are worth eating.

Clavulinopsis corniculata
FRUITIN G BODY branched (sometimes sparingly) from a common base or stalk, usually
small and delicate-looking; 2-9 cm high and broad (but usually 2-5 cm). BRANCHES
smooth, mostly hollow, bright yellow to yellow-orange or egg-yellow to deep ochraceous,
sometimes fading when dry or in age to creamy-buff; tips acute. STALK 2A mm thick,
colored like branches or duller, often with a coating of downy mycelium at base. Flesh
rather tough, thin, whitish; odor mild or farinaceous; taste mild or bitter. SPORE PRINT
white; spores 4.5-7.5 microns, round or nearly round, smooth, apiculate.
HABITAT: Solitary, scattered, or in groups on ground or dead wood in forests and at
their edges, in grassy areas, etc.; widely distributed. I have found it locally under cypress
and oak in December and January, accompanied by C. laeticolor and numerous waxy
caps (Hygrocybe and Camarophyllus species).
EDIBILITY: Probably edible, but too small to be of value.

Clavulinopsis corniculata is a small brightly colored branched species. Both specimens shown here
are elaborately branched, but some specimens havea well-developed“stalk” with only a few branches.
640 CLAVARIACEAE

COMMENTS: The bright yellow to ochre color and modest size make this branched coral
fungus fairly easy to recognize. Other yellow to orange coral and club fungi are either
unbranched (see C. laeticolor) or much larger and fleshier (see Ramaria). The jelly fungus
genus Calocera is somewhat similar, but usually has a viscid fruiting body. Other small
branched species include: C. umbrinella (-Clavaria cineroides), a dingy buff to grayish
or brownish species with round spores and several primary branches that arise from a
common base or “trunk,” occasional in our area in the fall and winter(usually in forests),
but more widely distributed; C. holmskjoldii, which resembles C. umbrinella but has
purplish branch tips and an aniselike odor when fresh; C. dichotomaand C. subtilis, which
are both white to faintly yellowish; and Clavulina (-Clavaria) ornatipes, with pallid to
pinkish-gray to brownish branches and a hairy brown stalk or “trunk.”

CLAVULINA & Allies

Small to medium-sized, coral-like fungi found on ground (Clavulina, Ramariopsis, Tremelloden-
dropsis) or wood (Clavicorona). FRUITING BODY usually branched, sometimes with a fleshy
base. BRANCHES mostly erect, usually pale or dull-colored (white to gray, tan, brownish, dingy
yellowish, or sometimes tinged pinkish or purple). SPORE PRINT white (or in Clavulina some¬
times yellowish after prolonged storage). Spores smooth or spiny. Basidia typically 2-spored
(Clavulina) or 4-spored (others). Spore-bearing surface typically not greening in ferrous sulfate.

THIS is a motley, artificial grouping of mostly white or dingy colored, branched coral
fungi. Their overall aspect is intermediate between that of the fairy clubs (Clavaria &
Allies) and the Ramarias. The pale or dingy color, white spores, and moderate size dis¬
tinguish them from most Ramarias, while the branched fruiting body separates them
from most fairy clubs.
The most common of the genera treated here is Clavulina, which has smooth spores and
2-spored basidia. In Ramariopsis, also common, the spores are minutely ornamented and
the basidia are 4-spored. In Clavicorona the fruiting body grows on wood and typically has
crownlike branch tips and amyloid spores, while Tremellodendropsis has tough, usually
flattened branches and basidia which appear partially partitioned.
The species treated here are probably edible, but are not as fleshy or desirable as the
Ramarias. In addition, some are quite tough while others are exceedingly fragile. They fruit
primarily in the woods. Five species are described and several others are keyed out.

Key to Clavulina & Allies

1. Growing on wood; branch tips usually crownlike .... Clavicoronapyxidata & others, p. 642
1. Not as above; usually growing on ground .2
2. Fruiting body tough and pliant, not white, typically with a mat of copious white mycelial
threads at base or in surrounding humus .(seeRamaria, p. 645)
2. Not as above; mycelial mat absent and/ or fruiting body brittle.3
3. Fruiting body lavender to purple .... Clavulina amethystina & others (see C. cinerea, p. 641)
3. Not as above (but branches may have a purple tinge) .4
4. Texture very tough or cartilaginous to slightly gelatinous; branches usually flattened; overall
color white or pallid (or sometimes greenish from algae); found mainly under hardwoods in
eastern North America (see photo at bottom of p. 644) .
. Tremellodendronpallidum & others (see Tremellodendropsis tuberosa, p. 643)
4. Not as above . 5
5. Texture tough; “stalk” usually comprising at least one half the height of fruiting body; branches
often flattened, pallid to brownish-gray or tinged dull purplish; base often with whitish down;
basidia appearing partially septate under microscope . Tremellodendropsis tuberosa, p. 643
5. Texture brittle to fragile, or if tough then not as above.6
CLAVULINA & ALLIES 641

6. Fruiting body branched, but stalk usually at least half the total height; color of brownish; taste
often bitter and/ or stalk with brown hairs; basidia 4-spored . (see Clavaria & Allies, p. 634)
6. Not as above . 7
7. Fruiting body white or pallid (or tinged buff, gray, or pinkish) .8
7. Fruiting body darker (gray to brownish-gray, bluish-gray, purplish-gray, etc.).
. Clavulina cinerea, below
8. Fruiting body unbranched or sparingly so .. Clavulina rugosa(see C. cristata group, below)
8. Fruiting body branched .9
9. Fruiting body profusely branched, fragile; branch tips typically neither toothed nor enlarged
.Ramariopsis kunzei, p.643
9. Fruiting body branched (but at times rather sparingly so), fragile to rather tough; branch tips
often toothed and / or enlarged . Clavulina cristata group, below

Clavulina cinerea (Ashy Coral Mushroom) Color Plate 169

FRUITING BODY erect or somewhat spreading, profusely branched, 2-11 cm tall and
broad. BRANCHES often irregular in shape, often resulting in a somewhat tangled ap¬
pearance; pallid soon becoming grayish to ashy-gray, purple-gray, bluish-gray, dark gray,
or even brownish-gray; smooth or wrinkled to somewhat flattened; tips acute or blunt,
often forked. STALK present as a short, fleshy sterile base or “trunk”; colored like the
branches, or often whitish at the very base. Flesh white, brittle; taste usually mild. SPORE
PRINT white; spores 6.5-11 * 5.5-10 microns, broadly elliptical to nearly round, smooth.
Basidia 2-spored.
HABITAT: Solitary, scattered, or in groups on ground in mixed woods and under
conifers; widely distributed but mainly northern. In California I have seen it in the fall and
winter, but have yet to find it in our area.
EDIBILITY: Edible and highly rated by some authorities, but rather fragile and insipid
in my experience.
COMMENTS: The ash-gray to dark gray or purple-tinged fruiting body separates this
conifer-lover from other branched coral fungi. Microscopically it is distinct by virtue
of its 2-spored basidia with curved spore stalks(sterigmata). The C. cristata group is closely
related but paler in color. C. (-Clavaria) amethystina is a beautiful purple branched
species; it is found occasionally in eastern N orth America and I have found it( or something
very similar) near Aptos, California, under redwood and tanoak. Ramariopsispulchella
is a very small, sparingly branched, lavender species. Clavaria zo/lingeri (see comments
under C. purpurea) is also small and purple, but has 4-spored basidia.

Clavulina cristata group (Crested Coral; Wrinkled Coral)

FRUITIN G BODY erect but extremely variable in shape and form, 2-7 (12) cm high, up to
5 cm broad; “typical” form branched, but other forms sparsely or irregularly branched and
still others unbranched. BRANCHES smooth in typical form, uneven or knobby or longi¬
tudinally wrinkled or flattened in other forms; tip(s) acute and often finely toothed in
“typical” form, but blunt and often enlarged in others; color usually white, but some¬
times tinged gray, buff, yellowish, or pinkish; tip(s) often darkening in age or dry weather.
STALK present as a sterile base; slender, white or darkened by a parasite. Flesh white,
brittle to rather tough. SPORE PRINT white; spores 7-11 (14) * 6.5-10 (12) microns,
nearly round, smooth. Basidia 2-spored.
HABITAT: Solitary to scattered or densely gregarious on ground in woods and grassy
areas; widely distributed and common from sea level up to timberline. In our area this
group fruits throughout the mushroom season and is especially prevalent under pine.
EDIBILITY: Edible. Some rate it highly, others do not.
Clavulina cristata, a cosmopolitan white to grayish coral fungus. Note how the branch tips are
“crested.” In our area it is especially common under pine.

COMMENTS: This species “complex” is extremely polymorphic (variable in shape and

form), and as a result is likely to confound the beginner. The“typical” form is distinguished
from Ramariopsis kunzei (which is also white and branched) by its frequently crested
(finely toothed) branch tips, smooth spores, and tendency of the stalk to darken in age when
attacked by a fungal parasite. The unbranched to sparingly branched, wrinkled or knobby
form, which is often called C. rugosa, is also common in our area(see photo at top of next
page). It is likely to be confused with species of Clavaria, which are differently colored
and/ or smooth and more regular in appearance. It also seems to intergrade with “typical”
C. cristata. Another closely related species, C. cinerea, is usually darker (see description
on p. 641). Also see Clavulinopsis dichotoma and C. subtilis (under C. corniculata).

Clavicoronapyxidata (Crown Coral Mushroom)

FRUITING BODY profusely branched from a common base or “stalk”; 5-12.5 cm high
and 2-8 cm wide. BRANCHES usually arising in tiers from the enlarged tips of lower
branches; whitish to pale yellow when young, becoming dull ochre to tan or tinged pinkish,
but lower portion often darkening to brownish or grayish-brown in age; tips usually
enlarged and pyxidate(i.e., crownlike or terminating in fringed cups). STALK present as a
slender, short sterile base or “trunk”; colored like lower branches. Flesh white, tough and
rather pliant; taste often somewhat acrid (peppery). SPORE PRINT white; spores 3.5-6 *
2-3 microns, elliptical, smooth, amyloid.
HABIT AT: S olitary or in small groups on dead hardwoods, especially aspen, willow, and
cottonwood; very widely distributed and common in some regions. I have seen it often in
the southern Rocky Mountains. It also occurs in northern California, but I have yet to
find it in our area.
EDIBILITY: Edible, but rather stringy and tough.
COMMENTS: The crownlike branch tips, pale color, and growth on wood distinguish this
widespread coral mushroom. Several Ramaria and Lentaria species grow on wood (see
R. stricta), but do not have crownlike branch tips. Other species: C. avellanea is grayish-
brown with paler crownlike tips; it grows on rotting conifers.
Left: Clavulina rugosa (see comments under the C. cristata group) is a wrinkled clublike or sparingly
branched whitish species. Right: Ramariopsis kunzei is an elaborately branched whitish coral
fungus that is especially common under conifers.

Ramariopsis kunzei (White Coral Mushroom)

FRUITING BODY erect or somewhat spreading, profusely branched, 2.5-10 cm tall and
3-8 cm wide. BRANCHES white to creamy-white, often tinged pinkish in age; smooth,
not usually compact; tips blunt or acute. STALK absent or present only as a short, fragile,
sometimes hairy base. Flesh white, fragile; taste mild. SPORE PRINT white; spores 3-5.5
x 2.5-4.5 microns, broadly elliptical to round, minutely spiny. Basidia 4-spored.
HABITAT: Scattered to densely gregarious onground in mixed woods and under conifers,
frequently hidden in the duff; widely distributed. It is common in our redwood forests from
late fall through early spring, but is not restricted to that habitat.
EDIBILITY: Harmless, fleshless, flavorless.
COMMENTS: Formerly known as Clavaria kunzei, this ubiquitous branched coral
fungus is best recognized in the field by its white color and marked fragility. It is more
profusely branched than typical Clavulina cristata and lacks the toothed branch tips
often found in that species, and has ornamented spores. Other species: R. californica is
a rare, smooth-spored, branched, white to yellowish species.

Tremellodendropsis tuberosa
FRUITING BODY rather sparsely branched from a tough base (stalk) or sometimes
scarcely branched; 2-7 (10) cm high, 0.5-4 cm broad. BRANCHES usually somewhat
flattened (especially the lower ones), erect, tough, whitish to buff, brownish, or grayish,
sometimes with a purple or pinkish tinge (caused by a parasite?); tips often paler and
brighter (whiter) when actively growing. ST ALK well-developed, usually one-third to one-
half the height of fruiting body; colored like branches or paler, often with a coating of
white mycelial down. Flesh white, tough, not staining. SPORE PRINT white; spores
13-20 x 4.5-6.5 microns, elongated-elliptical or spindle-shaped, smooth. Many or all of
the basidia appearing partially septate (partitioned) longitudinally at their apices.
HABITAT: Solitary or in groups on ground in woods or clearings; widely distributed. Itis
fairly common in coastal California in the late fall and winter, especially with bracken fern,
redwood, or cypress, but is easily overlooked because of its humble appearance.

643
Tremellodendropsis tuberosa is best recognized by its tough texture, pale color, and upright branches.

EDIBILITY: I can find no information on it.

COMMENTS: This rather nondescript coral fungus is included here because it represents
a possible “bridge” between the coral fungi, which have“normal” club-shaped basidia, and
the jelly fungi, some of which are coral-like and have longitudinally septate (partitioned)
basidia. In the field it can be told by its rather long stalk or trunk, pale or dingy color,
tough texture, and growth on the ground. Lentaricu byssiseda (see comments under
Ramaria stricta) is a rather similar whitish to pinkish-tan species that often grows on wood,
has less flattened branches, “ normal” basidia, and white mycelial threads at the base of the
stalk. It is widely distributed, but in California is more common in the Sierra Nevada than
on the coast. Other species: Tremellodendron pallidum (-T. schweinitzii) of eastern North
America is a beautiful whitish branched species that is classified as a jelly fungus because
it has longitudinally partitioned basidia. It is common under hardwoods along with a
smaller relative, T. candidum, and can be distinguished from other whitish coral fungi by its
tough or tenacious texture and flattened branches (see photograph). Another eastern
hardwood-lover, Tremella reticulata, has very blunt, hollow, somewhat gelatinous,
white branches.
Tremellodendron pallidum of eastern North America is a coral-like jelly fungus. It may be related to
Tremellodenropsis tuber osa (see comments under that species for more details).
 645

RAMARIA (Coral Fungi)

Medium-sized to large, coral-like fungi found on wood or ground. FRUITING BODY profusely
branched from a common, often fleshy base or stalk. BRANCHES mostly erect, smooth, never
ribbonlike; often brightly colored. SPORE PRINT typically yellowish to tan or ochraceous. Spores
usually ornamented (warted or spiny), produced on the surfaces of the branches. Spore-bearing
surface staining green or bluish in ferrous sulfate.

THESE are brittle or pliant fungi with elaborately branched fruiting bodies. They repre¬
sent an evolutionary advancement over the simpler coral fungi or fairy clubs (Clavaria
& Allies) insofar as branching greatly increases the available surface area on which to
produce spores. (The same can be said for the teeth fungi (Hydnaceae), except that their
“branches” hang downward and are called spines.)
Ramaria can be distinguished from other branched coral fungi (see Clavulina & Allies)
by its tan to ochraceous or orange-yellow spores (which are usually ornamented with
minute warts, spines, or ridges) and frequently colorful fruiting body. Virtually every hue
is represented, with yellow, orange, red, pink, and tan predominating. It is easily the most
attractive and prominent group of coral fungi, and never fails to attract the attention of
collectors. However, it is also the largest and most complex group, with over 35 species
in California, many more in the Pacific Northwest, and at least 100 in North America.
To facilitate identification, Ramaria can be crudely divided into two groups: medium¬
sized to large, terrestrial species with a fleshy, gelatinous, or brittle fruiting body (e.g.,
R. botrytis)\ and wood- or duff-inhabiting species with a fairly small, slender, pliant-tough
fruiting body (such as R. stricta, and the small, smooth-spored “satellite” genus Lenlaria,
which intergrades with Ramaria). However, pinpointing the exact identity of a species
within either group is a difficult task, even for a specialist. In part this is because of
the nature of the fruiting body—aside from color and texture, there are few criteria by
which to separate species in the field. As a result, certain names have been applied indis¬
criminately to a slew of similar—but autonomous—species. Any attempt to correct this
trend short of an exhaustive study of Ramaria would only contribute to the confusion.
Therefore, the descriptions offered here are rather broad in scope. This may not satisfy
/fa ma n'tf-researchers, but it will enable collectors to refer most of the Ramarias they find to
a species group or “complex” without resorting to detailed microscopic study and special
chemical tests. Besides, it is not necessary to know the exact identities of these coral fungi
to appreciate their beauty. The manner in which they arise from the murky depths of the
forest floor is indeed reminiscent of corals.
Ramarias are a popular group for the table, probably because they are so distinctive
and none are known to be dangerously poisonous. Only the large, fleshy species are worth
collecting, but some are bitter or prone to attack by fungal parasites such as Hypomyces
transformans, and all are apt to be riddled with maggots and are difficult to clean. A few
(notably R. gelatinosa and the R. formosa group) are mildly poisonous, and even the
so-called “edible” species have a laxative effect on some individuals. Therefore it is best
to sample each type cautiously (if at all) and not to overindulge.
Ramarias are woodland fungi. The slender, pliant forms are saprophytic on humus and
wood and are especially common under conifers. The large fleshy types are usually terres¬
trial and may or may not be mycorrhizal. In the Pacific Northwest they are partial to
hemlock and other conifers, in our area they favor tanoak, and in the Rocky Mountains
they gravitate toward spruce, but are by no means restricted to these habitats. Only eleven
species are described here but a number of others are keyed out. Most of them occur in the
Pacific Northwest because that region is particularly rich in Ramarias. (The key is based
on my own field experience plus information in The Ramarias of Western Washington by
Currie Marr and Daniel Stuntz, and Trial Key to the Species of Ramaria in the Pacific
Northwest, by Kit Scates.)
646 CLAVARIACEAE

Key to Ramaria
1. Growing on wood . 2
1. Growing on ground (or occasionally on very rotten wood) . 5
2. Branches pale yellow to tawny-buff, orangish, or pinkish-tan, the tips yellow at first; spores
warty.R. stricta, p. 648
2. Not as above; fruiting body pallid to buffy-tan, pinkish-tan, reddish-brown, brown, or darker,
(occasionally yellowish but then spores smooth); branch tips not usually yellow when fresh 3
3. Taste acrid (peppery); branches usually with a pinkish tinge, becoming darker (reddish-brown)
in age .R. acris (see R. stricta, p. 648)
3. Not as above; branches pallid to pinkish-tan, buffy-tan, brown, greenish-tinged, etc.4
4. Fruiting body creamy to yellowish or pinkish-tan (branch tips sometimes greenish); spore print
whitish and spores smooth . . . Lentaria byssiseda & L. pinicola (see Ramaria stricta, p. 648)
4. Fruiting body buffy-tan to pinkish-tan to brown or darker (sometimes greenish-tinged); spore
print yellowish to ochraceous and spores ornamented . . R. apiculata (see R. stricta, p. 648)
5. Fruiting body pliant and rather tough, small or medium-sized (rarely taller than 10 cm); stalk
or “trunk” slender to practically absent, with a mat of conspicuous white mycelial threads
attached to the base and/ or permeating the substrate (see photo on p. 650) . 6
5. Fruiting body medium-sized to large; mycelial mat absent, or if present then not as above
(sometimes tough and pliant but usually fleshier) . 8
6. Spore print whitish; fruiting body creamy to pinkish-tan or yellowish, sometimes with greenish
tips; often found near wood or in lignin-rich humus .
.Lentaria byssiseda & L. pinicola (see Ramaria stricta, p. 648)
6. Spore print yellowish to ochraceous; fruiting body pallid to yellowish, ochraceous, cinnamon-
tan, etc., sometimes with greenish stains; found in duff . 7
7. Fruiting body (or at least the lower branches) bruising or aging blue-green to olive-green,
especially in cold weather .R. abietina, p. 650
7. Not as above . R. myceliosa & others, p. 649
8. Flesh in base of stalk rusty to rusty-brown when cut open lengthwise (i.e., vertically) .9
8. Not as above . 11
9. Branches orange to pinkish or pinkish-red .R. amyloidea
9. Branches creamy to yellow or yellow-orange . 10
10. Branches whitish to pale yellow with whitish tips . R. velocimutans
10. Branches yellowish to yellow-orange with yellow tips. R. celerivirescens
11. Branches white to creamy when fresh', tips similarly colored or pinkish to purplish, brick-red,
orangish, brownish, or yellowish-buff (but not yellow); fruiting body often (but not always)
compact when young; flesh in base not gelatinous or semi-gelatinous .40
11. N ot as above; branches darker or more brightly colored when young and fresh or tips yellow or
flesh gelatinous to semi-gelatinous . 12
12. Odor fragrant, like cocoa butter; branches pinkish-lavender to vinaceous or duller; found in
eastern North America .R. cacao
12. Not as above . 13
13. Branches dingy-colored (yellow-brown to olive-brown or olive-gray, etc.) with a distinct violet
tinge to lowermost ones (when fresh) and/or upper stalk; base (stalk) usually well-developed
and fleshy (see photo on p. 651) . R.fennica, p. 650
13. Not as above (but branches may be entirely violet or have violet to vinaceous stains) .... 14
14. Branches entirely violet to lavender when fresh (but may fade or discolor in age) . 15
14. Branches differently colored (but may have some vinaceous or purplish stains) . 16
15. Spore print white; base not particularly fleshy.(see ClavulinaSt Allies, p. 640)
15. Spore print yellowish to ochre; base usually fleshy .R. fumigata & others, p. 651
16. Branches red to coral-red, rose-pink, scarlet, or magenta when fresh; branch tips not yellow,
of if yellow then branches red ... 17
16. Branches differently colored (pink to orange, salmon, yellow, brown, etc.); tips yellow or not
yellow . 18
RAMARIA 647

17. Branches usually red to coral-pink when fresh; flesh in base of stalk not amyloid .
.R. araiospora & others, p. 655
17. Not as above .R. stuntzii& others (see R. araiospora, p. 655)
18. Branch tips blunt and conspicuously swollen (like the fruiting body of a Clavariadelphus) 19
18. Not as above . 20
19. Branches white to yellowish or becoming tan; widespread but not common . . R. obtusissima
19. Branches tan to dull grayish-orange; taste often bitter; found in the Pacific Northwest .
.R. claviramulata
20. Base or lower branches covered with a cottony white tomentum (fuzz), or if not then branches
or stalk often hollow . 21
20. Not as above . 23
21. Fruiting body consisting of a dense bundle of upright, slender, sparsely or densely branched
“stems” arising from a common base; branches (“stems”) salmon to yellow-orange to peachy
with yellow tips when fresh (sometimes with purplish stains), pliant to somewhat waxy and
often hollow; widely distributed under both hardwoods and conifers.R. conjunctipes
21. Not as above . 22
22. Branches yellow or pale yellow; odor often fragrant or pungent-sweet .
.R. cystidiophora & R. synaptopoda (see R. sanguinea group, p. 653)
22. Not as above; differently colored or soon becoming so. 23
23. Flesh in base (stalk) gelatinous or showing gelatinous streaks or pockets when sliced open or
if not gelatinous then cartilaginous (tough and brittle) . 46
23. Flesh in base firm or fibrous or with watery areas, but not gelatinous or cartilaginous ... 24
24. Branches brown to dark brown with white or pallid tips when fresh; found in eastern North
America . R. grandis
24. Not as above . 25
25. Base or stalk (and sometimes the branches) staining brown to reddish-brown to wine-red,
violet, or blackish when bruised (often already with vinaceous to reddish-brown stains) . 26
25. Not as above; fruiting body not staining appreciably when bruised . 30
26. Branches becoming yellow-brown to reddish-brown in age; branches and base staining reddish-
brown to brown when bruised; found under conifers in the Pacific Northwest .
. R. testaceoflava (see R.formosa group, p. 654)
26. Not as above; branches usually lighter or brighter in color, at least when fresh; common . 27
27. Branches and tips yellow to creamy . R. sanguinea group, p. 653
27. Branches pinkish to orangish or yellow-orange, the tips sometimes yellow . 28
28. Fruiting body with a yellow zone or band near ground level (on lower branches or just below),
at least when young and fresh .R. sandaracina (see R. gelatinosa, p. 655)
28. Not as above . 29
29. Fruiting body staining brown to blackish, at least at base .R.formosa group, p. 654
29. Fruiting body staining wine-red to burgundy, at least at base .
. R. rubribrunnescens & R. maculatipes (see R.formosa group, p. 654)
30. Fresh branches and branch tips bright orange to yellow-orange with a distinct yellow zone or
band just below them (near ground level) . . R. aurantiisiccescens (see R. rasilispora, p. 652)
30. Not with above features (but may have some of them) . 31
31. Branches and branch tips bright orange to deep orange to yellow-orange or pinkish-orange 32
31. Not as above (but branches may be pinkish or orangish with yellow tips) . 34
32. Branches yellow-orange when young and fresh, becoming pale or dull orange in age. 37
32. Branches bright orange to deep orange or pinkish-orange when fresh but not yellow-orange 33
33. Branches bright orange to deep orange in all stages R. largentii (see R.formosa group, p. 654)
33. Branches usually scarlet to pinkish or pinkish-orange, at least when young and fresh .
.R. stuntzii, R. subbotrytis, & R. cyaneigranosa (see R. araiospora, p. 655)
34. Odor usually fragrant; branches yellow becoming yellow-brown in age; fruiting mainly in the
fall under conifers . R. flavobrunnescens (see R. rasilispora, p. 652)
34. Not as above . 35
648 CLAVARIACEAE

35. Branches orange to pinkish or pinkish-tan, the tips usually yellow or yellowish when fresh
.R. formosa group & others, p. 654
35. Not as above . 36
36. Branches yellow to yellow-orange or whitish with yellow tips (when fresh) . 37
36. Branches differently colored (usually dull) . 39
37. Found under hardwoods in eastern North America .... R. aurea (see R. rasilispora, p. 652)
37. Found under both hardwoods and conifers; common in western North America . 38
38. Fruiting body with a large, broad, conical or steeply tapered, rooting base; upper branches
never orange when fresh; found under northern and mountain conifers in the spring or
summer.R. magnipes (see R. rasilispora, p. 652)
38. Not as above; fruiting body with a large or small base and with yellow to orangish branches;
found in above habitat and season as well as many others .R. rasilispora, p. 652
39. Branches yellow-brown to orangish-brown or duller even when fresh (sometimes with a purplish
tinge); branches typically erect and slender; taste usually bitter when cooked; common under
conifers in the Pacific Northwest .R. acrisiccescens (see R.fennica, p. 650)
39. Not as above . 40
40. Fruiting body compact, creamy to tan, dingy yellow-brown, or cinnamon-buff; taste mild;
spores not striate; found in eastern North America . R. caulifloriformis
40. Not with above features; widespread and common . 41
41. Branch tips typically pinkish when young but soon fading so that the mature fruiting body is
entirely white or creamy; spores striate; found under conifers in the Pacific Northwest .
.R. rubrievanescens (see R. botrytis, p. 656)
41. Tips never pinkish, or if pinkish when young then retaining their color longer; spores striate
or not striate . 42
42. Base with reddish-brown or vinaceous stains or staining these colors when bruised; branch
tips whitish to yellow; spores not striate R. vinosimaculans (see R. sanguinea group, p. 653)
42. Not as above . 43
43. Odor usually pungently sweet; branch tips dull orange to brownish, yellowish-buff, or whitish
when fresh; spores striate .R. strasseri & others (see R. botrytis, p. 656)
43. Odor mild, or if sweetish then branch tips differently colored; spores striate or not . 44
44. Fruiting body whitish with yellow to creamy-yellow tips, at least while under the duff (often
yellower or brighter in color when exposed); spores not striate R. rasilispora & others, p. 652
44. Not as above; branch tips usually pinkish to reddish or purplish, at least when young .... 45
45. Taste typically mild (occasionally bitter or acrid); spores striate; very common .
.R. botrytis & others, p. 656
45. Taste typically bitter to very bitter; spores not striate or only obscurely so; widespread but not
particularly common .R. botrytoides (see R. botrytis, p. 656)
46. Fruiting body brownish-yellow to tan or yellowish-tan; flesh cartilaginous.
.R. cartilaginea (see R. gelatinosa, p. 655)
46. Not as above; differently colored and/ or flesh in base gelatinous or semi-gelatinous .47
47. Branches usually yellow or yellow-orange .R. flavigelatinosa (see R. gelatinosa, p. 655)
47. Branches creamy to pinkish, orangish, brownish, etc., but not yellow (but fruiting body may
have a yellow zone just below branches) . 48
48. Fruiting body usually with a yellow zone or band near ground level (near bases of branches),
at least when young.R. gelatiniaurantia & R. sandaracina (see R. gelatinosa, p. 655)
48. Fruiting body lacking marked yellow zone .R. gelatinosa, p. 655

Ramaria stricta (Strict Coral Mushroom)

FRUITING BODY profusely branched from a poorly developed base or stalk; 4-12 cm
high and 3-10 cm broad. BRANCHES erect, slender, mostly parallel, usually compact
but sometimes openly branched, sometimes grooved or flattened slightly; pale yellow
becoming pinkish-tan to tawny-buff, orangish, or sometimes light brown, usually
RAMARIA 649

staining slowly brown or vinaceous-brown when bruised; branch tips fine, pale yellow to
pale greenish-yellow when fresh. STALK rudimentary or practically absent, not well-
developed. Flesh tough, pliant; odor often faintly sweetish; taste often somewhat metallic
or bitter. SPORE PRINT cinnamon-buff to yellowish; spores 7-10 * 3.5-5.5 microns,
elliptical, minutely roughened.
HABITAT: Solitary or in groups or tufts on rotting logs and branches (mostly of hard¬
woods but also conifers); widely distributed. It is common in our area in the late fall and
winter, but seldom fruits in large numbers.
EDIBILITY: Inedible—there is very little flesh and the flavor is not pleasing.
COMMENTS: This is one of several Ramarias that grow on wood, though it may form
“log lines” (i.e., grow in rows originating from buried or extremely decomposed logs). The
upright, parallel orientation of the branches (see photo on p. 631) plus the yellowish branch
tips are distinctive. The form that grows on conifers is often rather bushy and yellowish
when young, while the one that grows on hardwoods is usually oranger. Other wood-
inhabiting species: /?. apiculata favors conifers but is dull buffy-tan to dull orange-brown
or vinaceous-cinnamon (often with whitish tips when young). It is widespread and
common, also bruises brown, and in one form exhibits green to blue-green tints on the
branch tips and/or stalk base. R. acris(-R. rubella) is usually acrid-tasting and pinkish-
tinged when young with paler tips but becomes reddish-brown or darker throughout
in age. Lentaria pinicola and L. byssiseda (both placed in Ramaria by some authorities)
grow on logs (mostly coniferous) or lignin-rich humus and have smooth whitish spores.
The former is tan to yellowish with spores shorter than 10 microns, while the latter is
pallid to pinkish-tan, sometimes has greenish branch tips, and has longer spores. Both
species have white mycelial threads or cottony material at the base; L. byssiseda is espe¬
cially common in the Sierra Nevada (but is widespread). None of these are worth eating.

Ramaria myceliosa
FRUITING BODY abruptly and profusely branched from a slender base (stalk); 2-6 (10)
cm high and broad. BRANCHES slender, spreading, pliant, yellowish to tan, ochraceous,
olive-ochre, cinnamon-buff, or dull orange, etc.; tips same color. STALK slender, pliant,
not very fleshy; same color as branches or paler, with abundant white mycelial threads
(rhizomorphs) attached to base and/or permeating the surrounding humus. Flesh thin,
whitish, pliant; taste usually bitter. SPORE PRINT yellowish or pale ochraceous; spores
3.5-6 * 2-4 microns, elliptical to nearly round, minutely spiny.
HABITAT: Scattered to densely gregarious in duff under conifers; known only from the
west coast, but similar species (see comments) are more widely distributed. In our area it
is often abundant under redwood in the fall, winter, and early spring.
EDIBILITY: Unknown, but hardly worth experimenting with because of its small size
and bitterish taste.

COMMENTS: This is one of several small (usually less than 10 cm high), dull, pliant,
terrestrial Ramarias with a conspicuous white mycelial mat and minutely spiny spores.
Other members of the club include: R. pusilla, an eastern version of R. myceliosa; R.
invalii, with golden to dull yellowish-orange branches and spores 6-8.5 * 4-6 microns;/?.
flaccida, pale creamy-ochre with paler tips and spores 5-8 microns long;/?, suecica, pallid
to pinkish-tan with spores 8-10 microns long;/?, gracdis, with minutely roughened rather
than spiny spores and a frequent licorice-like odor; and/?, murrillii, with cinnamon-brown
to dark reddish-brown branches and a whitish base plus spiny spores, found in south¬
eastern North America. None of these grow on wood like R. stricta and relatives, nor
do they develop the greenish or blue-green stains typical of R. abietina.
Ramaria abietina (-R. ochraceovirens) is an ochraceous species that develops greenish stains in age
or cold weather. Note the extensive mat of white mycelial threads at base of fruiting body.

Ramaria abietina (Green-Staining Coral Mushroom)

FRUITING BODY profusely branched from a slender base (stalk); 2-10 cm high and
broad. BRANCHES slender, pliant, ochraceous to pale cinnamon becoming olive-brown
to dingy brown in age, slowly staining greenish or blue-green when bruised or as it matures,
especially the lower branches. STALK up to 2.5 cm long, slender, white to ochraceous or
colored like branches, with white mycelial threads (rhizomorphs) at base and/or in
surrounding humus. Flesh pliant, often discoloring slightly vinaceous-brown; taste usually
bitter. SPORE PRINT pale yellowish-tan; spores 5.5-8 x 3-4.5 microns, elliptical, minutely
spiny.
HABITAT: Scattered to densely gregarious in duff under conifers (less commonly hard¬
woods); widely distributed. It is common in our area from the fall through early spring,
especially under redwood.
EDIBILITY: Too small, tough, and/or bitter to bother with.
COMMENTS: The tendency of the lower branches to develop greenish stains distinguishes
this species, which is also known as R. ochraceovirens, from the throngs of other small,
pliant, buff to dingy ochraceous, terrestrial Ramarias (see R. myceliosa). The green-
staining is especially evident in cold weather, and can be enhanced—if there is any doubt—
by placing the fruiting body in a freezer. R. apiculata (see comments under R. stricta)
may stain greenish, but normally grows on wood.

Ramaria fennica (Bitter Coral Mushroom)

FRUITING BODY typically with two to four large primary branches arising from a
common base (stalk), and many smaller secondary branches arising from the primary ones;
6-18 cm high and 5-12 cm broad. BRANCHES smooth, mostly erect, olive-gray to olive-
umber to smoky-yellow, grayish-tan, or yellow-brown, the basal (primary) branches
dark olive-brown with a violet tinge when fresh; tips olive-yellow to dingy buff. STALK
large, fleshy, white below, upper portion colored like the branches and usually tinged
violet when fresh. Flesh white, firm but brittle; taste often bitterish. SPORE PRINT pale
ochraceous; spores 8.5-12 * 3.5-5 microns, elliptical, roughened.
HABITAT: Solitary, scattered, oringroupsinhumus under both hardwoodsand conifers;
widely distributed. In our area it is usually associated, like other fleshy Ramarias, with
tanoak. It is not normally a common species, but occasionally fruits in great abundance
in the fall and early winter.
EDIBILITY: Unknown; the bitterish taste (when present) is a deterrent.

650
Ramaria fennica is a handsome upright coral fungus. Note the well-developed fleshy base or “trunk.”
Fresh specimens are usually tinged purple at base of branches.

COMMENTS: One of our most lovely coral fungi, this species is easily told by its overall
shape (see photograph) and dull color, plus the subtle violet tinge to the lower branches and
top of the trunk (when fresh). In contrast to many of the larger Ramarias, it is typically taller
than it is wide. R.fennnica var. violaceibrunnea is a more precise name for the west coast
variant. Other species: R. fumosiavellanea is a rare similarly-colored species with a much
smaller base. R. acrisiccescens, called the “Blah Ramaria” by Kit Scates in her extensive
key to Ramarias of the Pacific Northwest, is a common dull-colored species with slender,
erect branches and a slim, tapered base plus an acrid or bitter taste, at least when cooked.
It is common under conifers in the Pacific Northwest, but I have not seen it in our area.

Ramaria fumigata (Violet Coral Mushroom)

FRUITING BODY typically with two to four large primary branches arising from a
common base (stalk), and many smaller secondary branches arising from the primary ones;
5-12 cm high and 8-12 cm broad. BRANCHES smooth, violet or lilac, slowly discoloring
dingy yellowish-brown or smoky-yellow in old age; tips also violet. STALK distinct as
a white fleshy base or “trunk” below the main branches. Flesh white, firm; taste slightly
bitter or acrid. SPORE PRINT pale ochraceous; spores 8.5-11 x 3-4 microns, elliptical,
roughened.
HABITAT: Solitary or in small groups on ground in woods; widely distributed, but
apparently very rare in California. I have found it only once in our area, near Boulder
Creek, under tanoak and Douglas-fir, in December.
EDIBILITY: Unknown
COMMENTS: The violet branches and branch tips make this beautiful Ramaria un¬
mistakable. It is closely allied to R. fennica, but is somewhat smaller in my experience,
and more vividly colored. Other distinctly purple coral fungi are more sparsely branched
and/ or have white spores (see comments under Clavariapurpurea and Clavulina cinerea).
Other species: R. cedretorum, reported by Kit Scates from northern Idaho, is another rare
amethyst-colored species. It does not discolor as much in age and has slightly larger spores.

651
Ramaria rasilispora is the most common of several yellow to pale orange coral fungi. Note that the
flesh in sliced specimen is solid and white, not gelatinous.

Ramaria rasilispora (Yellow Coral Mushroom)

FRUITING BODY abundantly branched from a fleshy base or stalk, 5-40 cm high and
broad. BRANCHES smooth, mostly erect, creamy to pale yellow (var. scatesiana) or
yellow to yellow-orange or pale orange (var. rasilispora), but the lower branches often
whitish when very young and branches in both varieties often pale or dull orange to ochra-
ceous in old age; tips yellow or same color as branches, sometimes brownish in age. STALK
fleshy, tapering to a point or rooting; white or whitish. Flesh brittle or fibrous, often with
watery areas, white; taste mild. SPORE PRINT pale orange-yellow or ochraceous; spores
8-12 x 3-4.5 microns, cylindrical, smooth or slightly roughened.
HABITAT: Solitary to scattered or in groups or rings on ground in woods; common in
the West and probably more widely distributed. In our area it fruits in the fall and winter
under tanoak or less commonly live oak, but in other regions it is common under conifers,
especially fir. In mountain ranges such as the Sierra Nevada it usually fruits in the spring
and early summer, as does R. magnipes(see comments), which is common from the Siski¬
you Mountains northward under hemlock, fir, and other conifers.
EDIBILITY: Edible and quite popular, but difficult to clean and sometimes lacking in
flavor. Coral fungus connoisseur Herb Saylor says it’s good raw in salads or candied like
grapefruit rinds. Some people, however, are adversely affected by it, and R. magnipes
(see comments) is sometimes bitter when cooked.
COMMENTS: The fleshy, terrestrial, yellowish Ramarias do not lend themselves to glib
categorization. As a group, however, they are easily recognized by their shape and color,
and none appear to be dangerously poisonous. The names R.flava and R. aurea have
traditionally been applied indiscriminately to practically all yellow coral fungi, but are
inadequate for the many variants in the group. R. rasilispora is probably the most com¬
mon yellow species in California. R. magnipes is a very similar, characteristic feature
of the springtime mushroom flora of western mountains. It differs in having a larger
(7-14 x 4-6 cm), rooting, steeply tapered or broadly conical stalk, yellow (never orange)
branches, and slightly longer spores. Other fleshy, terrestrial, yellow Ramarias include:
R. flavobrunnescens, which usually has a sweet odor, yellow branches that become
brownish-yellow (or brownish-tipped) in age, and is very common at times in the Pacific
Northwest under western hemlock; R. flavigelatinosa(see comments under R. gelatinosa),
with watery-gelatinous or sometimes cartilaginous flesh; R. aurantiisiccescens, with
yellow-orange to orange branches and a yellow zone near ground level; and R. aurea,
a pale orange to yellow species that is said to be common under hardwoods in eastern
North America. None of these bruise wine-red at the base as in the R. sanguinea group
(or the “true” R.flava), and none are choice edibles.

652
Ramaria sanguinea group. Branches are yellow to nearly white and the base stains reddish or bur¬
gundy when bruised.

Ramaria sanguinea group (Bleeding Coral Mushroom)

FRUITING BODY profusely branched fromafleshybaseor“stalk”;4-25cmormorehigh
and broad. BRANCHES smooth, mostly erect; pale or clear yellow, the tips usually slightly
brighter yellow; lower (primary) branches turning dark red or vinaceous to reddish- or
vinaceous-brown when bruised. STALK fleshy, tapered below, often very short; white or
whitish, typically staining dark red or wine-red to vinaceous-brown when bruised or
already with such stains. Flesh white or yellowish, rather brittle; taste mild. SPORE
PRINT pale ochraceous; spores 8.5-12 * 3.5-4 microns, oblong or elongated-elliptical,
roughened or smooth.
HABITAT: Solitary, scattered, or in groups on ground under both hardwoods and coni¬
fers; widely distributed. In our area this species “complex” fruits in the late fall and winter
under tanoak, but is not as numerous as R. rasilispora. In the Pacific Northwest and
northern California several similar species (see comments) are common under conifers in
the fall, and I have also seen large fruitings near Lake Tahoe in October.
EDIBILITY: Presumably consumable; I haven’t tried it.
COMMENTS: The yellow branches and tendency to stain vinaceous or reddish-brown
plus the mild taste and non-gelatinous flesh are the telltale traits of this lovely coral mush¬
room and its close relatives. R.flava is a very similar, longer-spored species that has not
yet been reported from the west coast. Actually, there is considerable doubt as to whether
the “true” R. sanguinea occurs in the West, but there are a number of similar vinaceous-
staining species, including: R. vinosimaculans, with creamy to light yellow branch tips
and vinaceous stains at the base (and sometimes on the branches), and R. rubiginosa, a
fleshy species with darker yellow tips and prominent vinaceous or purplish-brown stains at
the base. Both of these species are common under conifers in California and the Pacific
Northwest, particularly in the fall. Two other yellow, vinaceous-staining species, R.
cystidiophora and R. synaptopoda, differ in possessing a coating of cottony white fuzz
at the base. The former is often fragrant while the latter resembles a bundle of strings with
“ox-blood red” spots at the base and usually has a pungent-sweet odor. For vinaceous-
staining species with oranger or pinker branches, see comments under the R. formosa
group, and also check R. conjunctipes (couplet #21 of the key to Ramaria) and the species
discussed under R. gelatinosa.

653
Ramaria formosa group. Branches are pinkish to orangish and tips are usually yellow when fresh.

Ramaria formosa group (Pinkish Coral Mushroom)

FRUITING BODY profusely branched from a fleshy base (stalk); 5-20 cm high and wide.
BRANCHES smooth or grooved, mostly erect; pinkish to pinkish-orange, salmon,
pinkish-tan, or even reddish, fading to pinkish-buff, tan, or dingy ochraceous in age; tips
yellow or colored like branches when fresh, also fading. STALK fleshy, usually tapering
downward; white at base, otherwise colored like branches (or paler). Flesh brittle to
fibrous, whitish or becoming pinkish to orange in age, not gelatinous; taste usually mild
but often astringent or bitter when cooked. SPORE PRINT pale ochraceous; spores
9-12.5 x 3.5-6 microns, elliptical, minutely roughened.
HABITAT: Solitary, widely scattered, or in groups or rings on ground in woods, widely
distributed. In the Pacific Northwest and Rocky Mountains it is one of the most common
Ramarias, fruiting mainly under conifers, but also hardwoods. In our area it favors tanoak
(like just about every other fleshy Ramaria) and is fairly common in the fall and winter.
EDIBILITY: Said to have cathartic effects on some (or many) individuals. Until the
identity of our variety (or varieties) is definitely established, it is best left alone.
COMMENTS: The above description encompasses several forms that pose great dif¬
ficulties to the avid coral-categorizer but have collectively passed under the name R.
formosa in North America. The pinkish to pinkish-orange to pinkish-tan branches and
non-gelatinous flesh are the main fieldmarks, but older specimens are easily confused with
other species. The European R. formosa is said to stain cinnamon to vinaceous-brown or
blackish when bruised (at least at the base). Several forms in the Pacific Northwest do,
whereas the one in our area does not. Which is the “true” R. formosa is for the Ramaria-
researchers to decide! Similar species which stain include: R. rubribrunnescens and R.
maculatipes, both bruising vinaceous or burgundy (the former often fragrant); the R
formosa of the Pacific Northwest already alluded to, which bruises brown; and./?, testa-
ceoflava var. brunnea (-R. brunnea), a handsome, bitter-tasting conifer-lover of the
Pacific Northwest with “apricot-buff’ to golden-yellow branches that soon become
brownish and quickly stain reddish-brown, plus a brown-staining base. Similarly-colored
western species that do not stain include: R. leptoformosa and R. rubricarnata, common
in the Pacific Northwest under conifers, the latter with a stout, chunky fruiting body and
salmon-tinged flesh inside the branches; R. longispora, with a yellow zone or band near
or at ground-level when young; R. cyaneigranosa, whose fruiting body is often flattened
(somewhat fanlike); and R. largentii, a beautiful species that is orange to deep orange
throughout (never pinkish and not yellow-tipped).

654
RAMARIA 655

Ramaria gelatinosa (Gelatinous Coral Mushroom)

FRUITING BODY profusely branched from a fleshy base or “stalk”; 5-15 cm high and
wide. BRANCHES mostly smooth and erect, orange to pinkish-orange to orange-buff,
pinkish-brown, or occasionally yellow-orange, sometimes developing a faint purple-gray
tinge in age. STALK or main trunk whitish, rather tough. Flesh colored like branches or
paler and showing translucent, often gelatinous pockets or streaks when stalk is sectioned
lengthwise; taste often bitter. SPORE PRINT pale ochraceous; spores 7-10 * 4.5-6
microns, elliptical, roughened (warted).
HABITAT: Widely scattered to gregarious in duff under conifers (hemlock, fir, etc.);
common in the fall in the Pacific Northwest and northern California, but more widely
distributed. I have yet to find it south of San Francisco.
EDIBILITY: Poisonous to some people (causing diarrhea and digestive upset); apparently
edible for others, but hardly worth the risk.
COMMENTS: The gelatinous interior of the base and overalldingy pinkishtodull orange
color are the forthright features of this fungus. It is most likely to be mistaken for the
R. formosa group, which does not have gelatinous flesh. The above description is for
R. gelatinosa var. oregonensis. The typical variety, which was originally collected under
hardwoods in the southeastern states, is similar but paler (creamy) when young. Other
western species with gelatinous or semi-gelatinous flesh include: R.flavigelatinosa, which
is yellower in color and quite common on the west coast;R. gelatiniaurantia and/?, san-
daracina, both with orange to deep orange branches and orange to yellow tips, and a band
of yellow just above ground level (near the bases of the branches); and R. cartilaginea,
a yellowish-tan to tan or brownish-yellow species with cartilaginous to very slightly
gelatinous flesh. None of these are worth eating.

Ramaria araiospora (Red Coral Mushroom) Color Plate 168

FRUITING BODY profusely branched from a fleshy base (stalk); 4-13 cm high, 3-10 cm
broad or more. BRANCHES smooth, brilliant coral-red to red, crimson, oreven magenta,
but often fading to coral-pink, light red, orange, or even paler in old age; tips red to
yellowish, fading slightly in age. STALK usually rather short, white or whitish (or dis¬
colored) at base, colored like branches (or paler) above. Flesh brittle; taste mild or slightly
krautlike. SPORE PRINT pale ochraceous; spores 8-13 * 3-5 microns, elliptical to
cylindrical, roughened. Flesh in base of stalk not amyloid.
HABITAT: Solitary, widely scattered, or in small groups on ground in woods; known
only from the west coast. I find it every year in the fall and winter under tanoak, but it is
not as common in our area as some of the other fleshy Ramarias. Farther north it fre¬
quently occurs with hemlock and other conifers.
EDIBILITY: Presumably edible, but not as fleshyas/?. botrytisand R. rasilispora. Besure
not to confuse faded specimens with the R. formosa group!
COMMENTS: The brilliant red color of fresh specimens makes this gorgeous coral
mushroom unmistakable. It used to pass under the name R. subbotrytis, but that species
is apparently eastern in distribution. There are at least two distinct varieties of R. araio¬
spora: var. araiospora, with yellowish branch tips in age, and var. rubella, with red tips.
As in other Ramarias, the color tends to fade with age, and old specimens barely hint at
their former splendor. R. cyaneigranosa is a somewhat similar species that has broader
spores and is usually more peach-colored when fresh. R. stuntzii, common in northern
California and the Pacific Northwest, is similar to R. araiospora in color or can show
more yellow below, deep red or magenta near the tips, and orange in between, but it has
amyloid flesh in the stalk. R. subbotrytis is an eastern species with salmon-orange to pink
branches and branch tips when fresh, but it becomes duller or slightly yellower in age.
Ramaria botrytis, young specimens. Note how compact the fruiting body is—it looks more like a
cauliflower than the cauliflower mushroom (Sparassis). Older specimens are not necessarily com¬
pact, however. Branch tips are purple-red to pinkish when young, but the color may fade with age.

Ramaria botrytis (Pink-Tipped Coral Mushroom)

FRUITING BODY profusely branched from a fleshy base or stalk; sometimes massive,
7-20 cm high and 6-30 cm broad or more when mature; usually very compact when young,
but less so in age. BRANCHES often crowded, white or pallid when young, gradually
becoming buff to dull ochraceous or tan in age; tips typically short, clefted, blunt, pink to
purple, vinaceous, or red when fresh (but orange to brownish-orange in var. aurantii-
ramosa), fading in age. STALK thick (1.5-6 cm), fleshy, the base often rounded and
swollen at or above ground level; white or whitish when fresh, tan to ochraceous in old age.
Flesh firm, solid, white; taste mild in some forms, somewhat bitter in others. SPORE
PRINT pale ochraceous to ochre-buff; spores 11-16 (20)x 4-6 microns, oblong-elliptical to
nearly cylindrical, longitudinally lined (striate).
HABITAT: Solitary, scattered, or in groups or rings on ground in woods, often partially
buried in the humus; very widely distributed. Along with R. strasseri (see comments) it is
common in the Pacific Northwest and Sierra Nevada under conifers in the summer, fall,
and spring (but is less common in the spring than R. rubripermanens—see comments). In
our area it favors tanoak and other hardwoods and fruits in the fall and winter.
EDIBILITY: Edible and choice, according to some. However, it has laxative effects on
some individuals and bitter look-alikes (see comments) occur. It is one of the largest and
fleshiest of all the Ramarias, so it is definitely worth trying.
COMMENTS: This handsome, compact coral mushroom is reminiscent of a cauliflower
—more so, in fact, than the cauliflower mushroom (Sparassis). Young specimens with
stubby pink- or purple-tipped branches are the most readily recognized of all the Ramarias
(see photo). In age, however, the fruiting body discolors and often loses its compact form,
and it is then easily confused with other Ramarias. The longitudinally striate spores,
however, are distinctive. There are several closely related species with longitudinally striate
spores. One, R. strasseri, is quite common (at least in western North America), but typically
has a tapered, rooting stalk and a frequently fragrant or spicy-sweet odor; it is whitish to
ochraceous or tan with whitish to ochre, tan, dull orange, or brownish tips. A second,/?.
secunda, closely resembles R. strasseri, but has shorter spores. A third,/?, rubripermanens,
is colored like typical R. botrytis, but has shorter spores (less than 13 microns long). A
fourth, /?. rubrievanescens, also has short spores, but shows pink on the branch tips only
when very young and is often entirely white or creamy at maturity. All of these occur in
western North America, mainly (but not exclusively) with conifers. Finally, there is /?.
botrytoides, a widespread species that mimics R. botrytis in color (whitish with pink
to purplish branch tips) but has an exceedingly acrid taste that can linger in one’s mouth
for up to twelve hours after tasting it! This species, which occurs occasionally in our area,
also differs in having non-striate (or only very faintly striate) spores, suggesting a closer
kinship to R. araiospora than to R. botrytis.
656
 657

SPARASSIS (Cauliflower Mushrooms)

Medium-sized to large fungi parasitic on the roots of trees (especially conifers). FRUITING BODY
typically a compact mass of flattened, wavy, ribbonlike branches or leafy lobes, often with a tough
rooting base; white to buff or tan. SPORE PRINT white. Spores smooth, elliptical.

THE branched fruiting body with wavy, flattened, leafy or ribbonlike lobes is unique to
Sparassis, and most mycologists now sequester it in a family of its own. Despite the
common name, our species looks more like a giant brain than a cauliflower. It fruits at the
bases of trees (another distinctive feature), appearing year after year in the same spots.
From a size and edibility standpoint, Sparassis is indisputably the king (or queen) of
the coral fungi. It is highly esteemed for its fragrance, flavor, and keeping quality, and I
found it to be a staple item in the monsoon diet of many Himalayan villagers. Only two
species occur in North America, one western and one eastern; a key hardly seems necessary.

Sparassis crispa (Cauliflower Mushroom) Color Plate 172

FRUITING BODY a compactly branched mass of flattened, leafy lobes arising from a
tough, rooting base or “stalk”; 12-60 cm or more high and broad. BRANCHES (lobes)
flattened, thin, usually leafy in appearance; wavy, crisped, or crimped, pliant; surfaces of
lobes smooth, whitish to creamy or yellowish, often becoming cinnamon-buff or tan in age
or dry weather, sometimes with darker brown stains on the edges. ST ALK 5-13 cm long, 2-5
cm thick, usually tapered downward, fleshy but very tough, buried deep in ground (or
wood). Flesh firm, white, fairly tough or elastic; odor spicy-fragrant. SPORE PRINT
white; spores 5-7 x 3-5 microns, broadly elliptical, smooth.
HABIT AT: Parasitic on the roots of conifers, usually growing solitary at or near the bases
of trunks or stumps; found in the coniferous forests of western North America. It is not
uncommon in our coastal pine forests, fruiting in the same spot year after year (or some¬
times twice a year), usually in the late fall or winter. It also grows on Douglas-fir and other
members of the pine family. It produces a brown or yellow carbonizing rot in its host, and
is economically significant in some regions. The best place to look for it is in older forests
where there are plenty of mature trees. S. spathulata{see below) can grow on hardwoods.
EDIBILITY: Edible and exceptional. The perfect mushroom for a special occasion, it is as
elegant as it looks intelligent. Thorough cooking is necessary, however, to render it tender,
and it is a royal pain-in-the-posterior to clean! Gentle sauteeing or parboiling followed by
baking or stewing is best. Fresh specimens can be stored for a week or two in a cool dry
place, but be sure to check first for maggots!

COMMENTS: Better known as S. radicata, this fantastic fungus looks more like a
sea sponge or “bouquet of egg noodles” (Alexander Smith) than a cauliflower. The
flattened, ribbonlike lobes or “branches”, plus the overall white to yellowish color and
tough, rooting base distinguish it from other coral fungi. The spicy odor is also distinctive,
but difficult to characterize. In the Pacific Northwest, 20-30(oreven50!) pound specimens
are not unheard of, but in our area the size range is generally 1 -5 pounds—or about the size
of a human head (a cross-section, coincidentally, reveals brainlike convolutions or
“canals”). Certain cup fungi(e.g., Pezizaproteana form sparassoides and Wynneasparas-
soides) sometimes mimic it, but are brown or lilac-tinted and/ or smaller and bear their
spores in asci rather than on basidia (a microscopic distinction). The name S. crispa has
also been applied to the equally edible cauliflower mushroom of eastern North America,
which favors oak and pine, has thicker, more erect, rigid-looking branches or lobes, and
lacks a rooting base. However, recent studies indicate that its “correct” name isS. spathu-
lata or S. herbstii, and that the western species, which has been called S. radicata, is the
“true” S. crispa. (I’m tempted to say that mushroom taxonomy is as intricately twisted
and convoluted as a cauliflower mushroom, but I won’t!)
658

Chanterelles spores

CANTHARELLACEAE
Fairly small to medium-large, mostly terrestrial woodland fungi. FRUITING BODY with a cap
and stalk. CAP typically depressed to vase-shaped or trumpet-shaped at maturity, surface not
typically viscid. UNDERSIDE smooth, wrinkled, veined, or with primitive foldlike, forking,
shallow, blunt, decurrent gills. STALK fleshy or hollow, continuous with cap. VEIL and VOLV A
absent. SPORE PRINT white to buff, yellowish, pinkish, salmon-tinged, or tan. Spores smooth
or roughened, not amyloid. Basidia long and narrow.

THESE are colorful and conspicuous mushrooms with a more or less vase-shaped to
funnel-shaped fruiting body differentiated into an upper or inner sterile surface (the cap)
and lower or outer spore-bearing surface (the hymenium). The latter may be smooth,
wrinkled, veined, or gilled, but the gills when present are foldlike (thick, shallow, blunt,
and usually joined by connecting veins) rather than bladelike and/ or thin-edged as in the
true agarics. The long, narrow basidia and relatively unspecialized hymenium suggest a
kinship between the chanterelles and coral fungi. In fact, if the flattened apex of a club coral
(Clavariadelphus) were broadened into a cap and the wrinkles on its sides amplified, you
would essentially have a chanterelle!
The three principal genera in the Cantharellaceae are treated together here. In
Craterellus the fruiting body is more or less trumpet-shaped and dark brown to gray or
black, and the spores are smooth; in Cantharellus it is broadly convex to plane or vase¬
shaped and often brightly colored, gills are usually present, and the spores are smooth; and
in Gomphus the fruiting body is cylindrical to vase-shaped with a veined or wrinkled
hymenium, and the spores are warted or wrinkled. A fourth genus, Polyozellus, is also
included here though it is usually relegated to a family of its own.
The chanterelles are great favorites with fungophiles. With the exception of the G.
floccosus group, all are apparently edible and many are excellent. The best (and most
plentiful!) of the local species are the renowned chanterelle (Cantharellus cibarius)a.nd the
unheralded but far tastier horn of plenty (Craterellus cornucopioid.es). Both refrigerate
well and are usually maggot-free, and since large crops are commonplace, they can be
preserved for later use. The horn of plenty is excellent dried. Chanterelles, on the other
hand, dry miserably (they become as tough as leather), but can be pickled, canned, or
sauteed and frozen. (Enormous quantities of chanterelles are gathered in the Pacific
Northwest and shipped in brine to Germany—five million pounds annually according to
one estimate!)
The chanterelles are strictly woodland fungi and are saprophytic or mycorrhizal with a
broad range of tree hosts. They are not a particularly large group, but are quite conspi¬
cuous, easy to identify, and have an interesting distribution pattern. Many species are
characteristic components of cold northern and montane coniferous forests—e.g., Gom¬
phus clavatus, G. floccosus, and Cantharellus infundibuliformis. A slew of others favor
the warm hardwood forests of eastern North America—e.g., Cantharellus cinnabarinus,
C. ignicolor, and C. lateritius. But there is a puzzling paucity of species in our area, with
only a few of the truly cosmopolitan ones (like the chanterelle) represented. Nine species
are depicted here (most of them with color plates) and several others are keyed out.

Key to the Cantharellaceae

1. Fruiting body gelatinous or rubbery and pliant .(see Tremellales & Allies, p. 669)
1. Fruiting body fleshy to rather tough and pliant, but not rubbery-pliant or gelatinous .2
The chanterelle, Cantharellus cibarius, is a prized edible mushroom. Note shallow, blunt, veined gills.
Cap is depressed or funnel-shaped in these mature specimens, but can be convex when very young (see
description on p. 662).

2. Fruiting body small or minute (usually less than 2 cm broad), typically white to grayish- or
brownish-tinged; stalk absent, or if present often short and off-center to lateral; fertile surface
(underside of cap) with meandering veins or shallow gills; growing on sticks, moss, etc.
(not often terrestrial); especially common in northern regions (see Pleurotus &. Allies, p. 132)
2. Fruiting body medium-sized to large, or if small then not as above .3
3. Fruiting body a gilled mushroom (usually Laetarius or Russula) covered by a layer of minutely
pimpled tissue that completely engulfs the gills or makes them look blunt and chanterelle-like;
flesh crisp, brittle, usually white; spores borne inside asci .(see Pyrenomycetes, p. 878)
3. Not as above; surface not finely pimpled; spores borne on basidia . 4
4. Cap dark when fresh (dark gray to dark brown, black, violet-black, or bluish-black) .5
4. Cap brighter or lighter than above (including olive-buff, tan, light brown, etc.) . 11
5. Entire fruiting body deep blue to blue-black or violet-black, usually compound (a cluster of
spoon- or fan-shaped caps); found under northern and montane conifers (especially spruce
and fir) .Polyozellus multiplex, p. 668
5. Not as above (but fruiting body may be black or bluish-gray or more or less funnel- to trumpet¬
shaped, and may occur in clusters) . 6
6. Underside of cap (fertile surface) with distinct gills, the gills often forked .7
6. Underside of cap smooth to wrinkled or with sinuous (squiggly) veins, but lacking gills ... 8
7. Entire fruiting body dark when fresh (colored more or less like cap or the fertile surface some¬
times paler) . Craterellus cinereus & others, p. 665
7. Not as above; either the stalk or gills (or both) yellowish to orange (or stalk with a yellowish to
orange undersurface and darker fibrils) . 17
8. Underside of cap (fertile surface) with prominent veins (at least at maturity); odor sickeningly
sweet in age, or if not then fertile surface usually developing a strong yellowish tinge as spores
mature; not common .Craterellusfoetidus& C. sirtuosus (see C. cinereus, p. 665)
8. Not as above; underside of cap smooth to slightly wrinkled; common .9
9. Fruiting body typically 4 cm or more high when mature, trumpet-shaped to petunia-shaped
or tubular (i.e., hollow nearly to base) . Craterellus cornucopioides & others, p. 666
9. Fruiting body smaller and/ or differently shaped (not hollow); cap margin sometimes white or
pale when actively growing . 10
10. Cap typically less than 2 cm broad and stalk 1 -3 mm thick; fruiting body grayish to brown or
blackish but not chocolate-brown or purple-brown; spores smooth; found in eastern North
America (especially the Southeast) . . . Craterellus calycuius (see C. cornucopioides, p. 666)
10. Not as above; cap up to 5 cm broad (or larger if compound), usually purple-brown to chocolate-
brown (but sometimes grayish-brown) and/ or compound (with more than one cap); cap
margin often fringed; spores often angular or warty . (see Stereaceae & Allies, p. 604)

659
660 CANTHARELLACEAE

11. Fruiting body more or less vaselike and coarsely scaly at the center (or throughout), at least
in age (see Color Plate 174) . 12
11. Not as above; cap smooth or breaking up to form small scales, but not coarsely scaly .... 13
12. Cap reddish to bright orange to orange-buff, at least when fresh (may fade to tan or paler) . .
.Gomphusfloccosus group, p. 661
12. Cap with tan to brown scales, never brightly colored .
.Gomphus kauffmanii(see G. floccosus group, p. 661)
13. Flesh and gills (or underside of cap) white or whitish, but staining purplish to purple-brown
when bruised; cap usually salmon-colored to orange; known only from the Southeast, not
common .Cantharellus (-Gloeocantharellus)purpurascens
13. Not as above (but gills or fertile surface may be purple or purple-tinged). 14
14. Fertile surface and/ or upper stalk purple or with a purplish tinge when fresh\ cap tan to olive-
buff, olive-brown, yellow-brown, or purplish; stalk thick and solid, firm, continuous with cap
(i.e., fruiting body resembling a chopped-off, often lopsided club, at least when young);
found under conifers, often (but not always) in fused clusters . . . Gomphus clavatus, p. 661
14. Not with above features (differently colored or stalk slender and hollow in age, etc.) . 15
15. Fertile surface (underside of cap) with gills (or lacking them only in the button stage), the gills
often forked or with connecting veins . 16
15. Fertile surface (underside of cap) smooth to slightly wrinkled or at times developing shallow
veins at maturity, but lacking gills . 22
16. Fresh fruiting body bright red to pink to orange-red .Cantharellus cinnabarinus, p. 664
16. Fruiting body differently colored (white, yellow, orange, salmon-orange, brown, etc.) ... 17
17. Either growing on wood (or lignin-rich humus) and stalk absent or rudimentary or gills orange
to deep orange and usually dichotomously forked and fairly thin inage and cap often brownish,
at least at the center.(see Paxillaceae, p. 476)
17. Not as above; gills usually blunt and thick (at least when young) and often forked irregularly
(rather than dichotomously), orange to yellow-orange, white, or differently colored .... 18
18. Cap grayish to brown or yellow-brown when young (but may break up into scales or fibrils,
revealing the yellowish to orangish undersurface); stalk often hollow, at least in age .... 19
18. Not as above; cap white, yellow, or orange when fresh (unless faded); stalk hollow or solid 20
19. Stalk brown or with brownish fibrils when young; gills yellowish to orangish; found in eastern
North America . Cantharellus appalachiensis
19. Stalk usually yellowish to orange when young and fresh; gills often with a grayish, brownish, or
purplish tinge; widespread .Cantharellus infundibuliformis group, p. 665
20. Cap white or whitish, sometimes with orange or yellow stains Cantharellus subalbidus, p. 662
20. Cap orange-buff to salmon-buff to ochre-tawny to bright yellow or orange, at least when fresh
(may fade to whitish in sunlight) . 21
21. Stalk slender (usually less than 1 cm thick) and usually hollow in age; flesh very thin; cap smallish;
common in eastern North America .
.Cantharellus ignicolor & C. minor (see C. inf undibuliformis group, p. 665)
21. Stalk slender to very thick (0.5-2 cm or more), solid (unless maggot-eaten); cap medium-sized to
very large or sometimes small; flesh thick; common, widespread Cantharellus cibarius, p. 662
22. Mature fruiting body with a flattened top, but without a true cap (i.e., cap margin blunt or non¬
existent—see Color Plate 166); flesh often punky or stringy . . (see Clavariadelphus, p. 632)
22. Not as above; fruiting body normally with a cap that has a well-defined, acute edge (margin) 23
23. Fruiting body whitish or dull-colored; stalk thin (1-3 mm); cap usually small, often rosette-like
(with petal-like lobes) in age; flesh very thin and tough . . . (see Stereaceae & Allies, p. 604)
23. Not as above; fruiting body fleshy though sometimes thin-fleshed, usually brightly colored
at least in part . 24
24. Cap brown to ochre-brown or yellow-brown, or sometimes more brightly colored; flesh very
thin; odor typically mild . . Cantharellus xanthopus (see C. infundibuliformis group, p. 665)
24. Cap brightly colored (usually yellow to orange or pinkish-orange); flesh fairly thick (at least
at center of cap); odor often fruity or fragrant . 25
25. Fruiting body more or less trumpet-shaped, usually clustered and fragrant.
.Cantharellus odoratus (see C. cibarius, p. 662)
25. Not as above .Cantharellus lateritius & C. confluens (see C. cibarius, p. 662)
GOMPHUS 661

Gomphus clavatus (Pig’s Ears) Color Plate 176

FRUITING BODY 6-20 cm or more high, nearly cylindrical to more or less club-shaped
but with a flattened top; often growing in fused or compound clusters. CAP(s) 2-10 (15)
cm broad, plane or depressed with an uplifted, often wavy or lobed margin that is often
more highly developed on one side than the other; surface moist or dry but not viscid,
smooth or breaking up into minute scales, light purplish to purplish-tan to olive-brown,
olive-buff, tan, or yellowish-buff. Flesh thick, firm, white or buff; odor mild. UNDER¬
SIDE (fertile surface) with numerous shallow, blunt, deeply decurrent, forking veins
or wrinkles which occasionally give it a poroid appearance; color variable but usually dull
purple to purplish-tan or showing at least some purple tints below (near the stalk), slowly
fading to dull ochre, tan, orbuff. STALK(s)3-5(10)cmlong, 1-3 cm thick, continuous with
cap, central or off-center, often fused together below, equal or narrowed at base (if soli¬
tary), often curved; solid, firm, buff to pale purple. SPORE PRINT pale tan to pale ochre;
spores 10-13 * 4-6.5 microns, elliptical, slightly wrinkled or warted.
HABITAT: Scattered to gregarious, often in fused pairs or clusters, on ground under
conifers; widely distributed in northern North America, fruiting mainly in the late summer
and fall. It is common in the Pacific Northwest and northern California, but I have not
seen it south of San Francisco.
EDIBILITY: Edible and considered choice by some, but I am not particularly fond of it.
COMMENTS: Also known as Cantharellus clavatus, this northern conifer-lover is easily
distinguished by its overall shape (like a chopped-off, frequently lopsided club), dull
purplish veined spore-producing surface, and growth habit in fused clusters. It is aptly
characterized in one book as “that funny looking thing which is purplish underneath.’’The
common name, “pig’s ears,” is also applied to an Ascomycete, Discina perlata. Other
species: G. (-Pseudocraterellus) pseudoclavatus is said to be similar, but has smooth
spores and favors hardwoods. It is apparently quite rare; I have not seen it.

Gomphus floccosus group Color Plate 174

(Scaly Chanterelle; Woolly Chanterelle)
FRUITING BODY nearly cylindrical, soon becoming more or less trumpet- or vase¬
shaped, 5-20 cm high. CAP3-15cm broad, depressed and soon hollow in the center; surface
reddish to bright orange, buffy-, oryellow-orange(butoftenfadingl), with large cottony or
woolly scales, the ones at the center often recurved in age; margin often wavy or lobed.
Flesh rather fibrous, whitish; taste mild to sour. UNDERSIDE (fertile surface) milky
white to creamy, buff, dingy yellowish, or pale ochre; with blunt, shallow, frequently
forking, deeply decurrent veins or ridges, sometimes with a somewhat poroid appearance
in age. STALK 3-10 cm long, 1-3 cm thick, continuous with cap, central or off-center,
tapering downward, solid becoming hollow in age, white to orange or yellowish. SPORE
PRINT ochraceous; spores 10-16 * 5-8 microns, elliptical, slightly wrinkled or warted.
HABITAT: Scattered to gregarious or sometimes clustered on ground under conifers;
widely distributed in northern and montane North America. Along with G. bonari (see
comments), it is common in northern California and the Sierra Nevada in the summer,
fall, and even spring, but like G. clavatus, it does not seem to occur on the coast south of
San Francisco. It is also common in the Southwest, Pacific Northwest, and Rockies.
EDIBILITY: Not recommended. Some people pronounce it delicious, but others are
adversely affected by it (they suffer gastric upsets) and the specimens I’ve sampled had
an unpleasant sour taste. Its only asset is its distinctiveness—it would be very difficult
to confuse it with anything other than its close relatives (see comments).
662 CANTHARELLACEAE

COMMENTS: Also known as Cantharellus floccosus, this striking mushroom and its
close relatives are easily told by their vase-shaped fruiting body with a reddish-orange to
orange-buff scaly cap and pallid veined exterior. The name G. bonari has been given to the
western variety with milky white exterior, smaller spores, tendency to grow in clusters, and
duller, paler, or more cinnamon-colored cap; it is oftencommon under mountainconifers.
Another western conifer-lover, G. kauffmanii, is similar but has a tan to brown or ochre-
tawny cap up to 35 cm broad with very prominent, brittle scales. It is sometimes common
in northern California and the Sierra Nevada as well as in the Pacific Northwest and
Southwest. Like G. floccosus and G. bonari, it should be eaten very cautiously if at all.

Cantharellus subalbidus (White Chanterelle) Color Plate 179

CAP 4-15 cm broad, plane to broadly depressed with an often wavy or irregularly lobed
margin; surface smooth or breaking up into small scales in age, moist to dry but not viscid;
dull white or whitish, bruising yellowish-orange to orange-brown. Flesh thick, firm, white;
odor mild or faintly fragrant. UNDERSIDE (fertile surface) with thick, well-spaced,
shallow, blunt, deeply decurrent, foldlike gills which are usually forked or veined between;
dull white or pinkish-tinged, often staining yellowish to orange in age or where bruised.
STALK 2-7 cm long, 1 -5 cm thick, equal or tapered downward, central or off-center, solid,
firm, smooth, dull whitish, discoloring yellowish-orange to orange-brown in age or where
bruised. SPORE PRINT white; spores 7-9 * 5-5.5 microns, elliptical, smooth.
HABITAT: Solitary, scattered, or gregarious on ground in woods; known only from
the Pacific Northwest and California. It is quite abundant under second-growth conifers
in the late summer, fall, and winter in Washington, Oregon, and northern California, and
I have also seen large fruitings in Idaho. In our area it fruits in the fall and early winter, but
is only occasionally common. The best places to look for it are on the high, exposed ridges
of our coastal mountains under tanoak and madrone or in manzanita thickets mixed with
knobcone pine. It often grows with Armillaria ponderosa, another choice edible.
EDIBILITY: Edible and choice—as good as C. cibarius or even better, and like that
species, rarely attacked by maggots. In the Pacific Northwest the buttons are firm, meaty,
and very compact—ideal for marinating or prolonged refrigeration. Unfortunately, it is
not nearly so common in our area and the fruiting bodies, when found, are often water¬
logged and overripe.
COMMENTS: The white chanterellecanbemistakenfora Clitocybeor Hygrophorus, but
the gills are distinctly “chanterellesque”—thick, shallow, blunt, and usually veined. It
differs from the common chanterelle (C. cibarius) in its dull whitish color and white spores,
but in size and shape the two are quite similar and can even be found growing together.

Cantharellus cibarius (Chanterelle) Color Plates 175,177,178

CAP (2) 3-15 (25) cm broad, broadly convex when young, becoming plane to depressed or
vase-shaped in age; surface smooth or occasionally cracked, not viscid; orange to bright
golden-orange, egg-yellow, or yellow (but in dry weather or direct sunlight often bleached
out and cracked or even greenish from algae); margin usually lobed or wavy, at first in¬
curved. Flesh thick, firm, whitish or tinged yellow to orange underthecuticle;odormild or
faintly fruity (like pumpkins or apricots); taste mild to somewhat peppery or occasionally
bitter. UNDERSIDE (fertile surface) with thick, well-spaced to close, shallow (narrow),
blunt, foldlike, deeply decurrent gills which are often forked or cross-veined; colored like
cap or more often paler (but brighter if cap has faded); not fading, but sometimes
developing dingy orange-brown stains in old age. STALK 2-10 cm long, 0.5-3 (5) cm thick,
Cantharellus cibarius. Note the white flesh in sliced specimen (upper left) and shallow, decurrent,
veined gills. It varies greatly in stature and color (see color plates) and favors oak as well as conifers.

equal or tapered downward or sometimes enlarged at base, solid, dry, firm, colored like cap
or paler, often staining ochraceous to orange-brown. SPORE PRINT creamy or yellowin
most forms, but pinkish in one variety; spores 7-11 * 4-6 microns, elliptical, smooth.
HABITAT: Widely scattered to gregarious on ground in woods, occurring throughout the
north temperate zone and very common on the west coast in the fall, winter, and spring.
In northern California and the Pacific Northwest it grows mainly with conifers, but in our
area it favors live oaks, especially those at the edges of pastures. It has a long growing season
and is a joy to find—brilliant splashes of gold against the subdued backdrop of decaying
leaves. If you see one, probe around and you’ll probably uncover more—they often hide
under the humus. If rainfall is sufficient, successive crops are produced over a long period
of time, so check your patches regularly (but don’t check mine!).
EDIBILITY: Edible and choice—the best known wild mushroom in California, if not in
North America. Itissoplentifulandpopularonthe west coast that it is now being harvested
commercially and shipped abroad or sold to restaurants and markets. A few tips for the
uninitiated: Chanterelles should always be cooked as they are somewhat fibrous raw. Pick
selectively—firm specimens will keep in the refrigerator for a week, whereas waterlogged
individuals will rot rapidly and cook up slimy. Even firm ones have a high water content
and should be sauteed slowly and thoroughly in an open pan. You waste them if you can’t
taste them, so don’t just mix them in with a bunch of other vegetables. They are best in
simple dishes that highlight their delicate flavor and fruity fragrance—cream of chanterelle
soup is a traditional favorite. On the west coast chanterelles are virtually free of maggots,
an asset keenly appreciated by those who hunt them—but they’re often full of dirt, a quality
deeply detested by those who have to clean them! (If your “chanterelles” are pristine and
wormy, they may not be chanterelles!). Curiously, it’s just the opposite in the eastern
United States, where they are usually quite spotless, but often lunched on by maggots!
Imported chanterelles can be purchased in small tins at delicatessens—for an outrageous
price, of course. They are low in protein but high in vitamin A. Attempts to cultivate
them commercially have not been successful.

COMMENTS: Cantharellus cibarius is the proud possessor of a plethora of popular

names—more than any other mushroom with the possible exception of Boletusedulis. The
best known are chanterelle and girolle (both French) and Pfifferling (German). From
region to region the chanterelle varies considerably in size, stature, color, and fruiting
habits (several forms have been described), but is easily identified by its yellow-orange cap

663
664 CANTHARELLACEAE

and “primitive” gills (blunt, thick, shallow, veined or forking, and foldlike rather than
bladelike). The firm whitish flesh and wavy cap margin are also characteristic. On the west
coast it fruits in cool weather and is often very large and thick-stemmed (one pound
specimens are not uncommon), with an orange cap, faintly fruity odor, and pale, copiously
veined gills (although a smaller, slimmer, cleaner form grows under Sitka spruce). In
eastern North America it is commonest in the summer and is usually much smaller (caps
average 3-6 cm broad) and often yellower, with a slender, well-developed stalk and little
or no odor (see Color Plate 175). Also, the gills may be less “primitive” (deeper, with
thinner edges and few if any cross-veins, as shown in Color Plate 177), and are sometimes
brightly colored. In the southern Rocky Mountains, on the other hand, the fruiting bodies
are often packed closely together and are small and stubby, with bleached-out caps
(if growing in sunlight) and brilliant yellow to yellow-orange gills.
Mushrooms most likely to be mistaken for the chanterelle include the false chanterelle
(Hygrophoropsis aurantiaca), various waxy caps (Camarophylluspratensis, Hygrocybe
flavescens, etc.), Lactarius alnicola, Leucopaxillus albissimus, the Gymnopilus
spectabilis group, Hypomyces lactifluorum, and the jack-o-lantern mushrooms
(Omphalotus speciesj. The latter are poisonous, but have brightly colored flesh and thin,
crowded, unforked, bladelike (“true”) gills, and usually grow in clusters on or near wood.
Except for the Hypomyces, which is a parasitized gilled mushroom, the others also have
bladelike gills. (It is sometimes said that true chanterelles have “false” gills, while false
chanterelles have “true” gills.) Aside from the many variations within C. cibarius, there is
C.formosus, a northern conifer-lover with a convex, yellow to brownish cap and pinkish-
tinged gills, and three similar edible species with a smooth to only slightly wrinkled
fertile surface. These are common in eastern North American and are often mistaken for
C. cibarius. The most common and widespread of the three, C. lateritius (-Craterellus
cantharellus) has a yellow-orange to pinkish-orange fruiting body; C. confluens has a
yellower, often compound fruiting body and a tropical-subtropical distribution; C.
odoratus (-Craterellus odoratus) is a gorgeous trumpet-shaped, yellow-orange-capped
southern species that is usually strongly fragrant and cespitose (clustered). C. lateritius
has also been reported (but not verified) from the Pacific Northwest.

Cantharellus cinnabarinus (Red Chanterelle) Color Plate 181

CAP 1-4 (7) cm broad, convex to nearly plane at first, plane to shallowly funnel-shaped
in age, the margin often lobed or wavy and often remaining incurved for a long time;
surface more or less smooth, dry, bright red to red-orange to flamingo-pink when fresh,
often paler (or even whitish) in age or sunlight. Flesh thin, whitish below, usually tinged
cap color above; odor typically mild, taste mild to peppery. UNDERSIDE(fertile surface)
with thick, shallow, blunt, decurrent gills which are fairly close to well-spaced, usually
forked or veined, and colored like the fresh cap or slightly paler (or pinker). STALK
1-4 (6) cm long, 0.3-1 cm thick, equal or tapered downward, often curved, solid, firm, dull
red or colored like fresh cap (or pinker), the base sometimes whitish. SPORE PRINT
usually pinkish-tinged (pinkish-cream); spores 6-9(11) * 4-6 microns, elliptical, smooth.
HABITAT: Scattered to gregarious orclusteredongroundin woodsorattheiredges, open
areas, in beds of moss, etc.; fairly common in eastern North America (especially south¬
ward) in the summer and early fall. I have seen large fruitings under oak and pine in Michi¬
gan, Pennsylvania, and North Carolina. It apparently does not occur in the West, but is
to be looked for in Arizona (one unverified report places it in the Pacific Northwest).
EDIBILITY: Edible and quite good, though delicate in flavor. It sometimes grows with
C. cibarius, and the two species make colorful companions in the same dish.
COMMENTS: The modest size and bright pinkish-red to orange-red color make this
chanterelle practically unmistakable. It is vaguely reminiscent of a waxy cap (Hygrocybe),
but the blunt, veined or forked gills distinguish it.
CANTHARELLUS 665

Cantharellus infundibuliformis group Color Plate 180

(Funnel Chanterelle; Winter Chanterelle)
CAP 1-6(11)) cm broad, convex to plane with a depressed center or becoming broadly
depressed to funnel-shaped (often with a perforated or hollow center) in age; surface
smooth to slightly wrinkled or finely fibrillose-scaly, not viscid, dark brown to brown to
dingy yellow-brown, sometimes fading to paler brown or the yellowish background color
becoming dominant as it dries or ages; margin usually wavy or irregularly lobed. Flesh
thin, rather tough and pliant, colored like cap or paler; odor mild or slightly fragrant; taste
mild or bitterish. UNDERSIDE (fertile surface) with shallow (narrow), thick, blunt, well¬
spaced, decurrent, forked or veined gills; pale yellow or pale orange-yellow becoming
grayish, brownish, or violet-tinted in age. STALK 2-8 cm long, 0.3-1 (2) cm thick, equal or
tapered at either end, smooth, often grooved and/ or flattened, becoming hollow; orange-
yellow to yellow, duller in age (dingy yellow or yellow-brown to grayish-orange). SPORE
PRINT creamy-buff to yellowish (or white in one form); spores 9-12 * 6.5-8 microns,
elliptical, smooth.
HABITAT: Scattered to gregarious in moss and humus and on rotting wood in cold, damp
coniferous forests and bogs throughout northern North America; fruiting from late
summer through the winter, common. The farthest south I’ve seen it is in Mendocino
County, California, in December, January, and February.
EDIBILITY: Edible, but small and thin-fleshed. Some people relish it nevertheless,
and in Finland it is harvested commercially.
COMMENTS: This characteristic feature of northern bogs and cold coniferous forests
can be recognized by its modest size, dark brown to dingy yellow-brown cap, yellowish to
gray or purple-tinged gills, and slender, hollow, yellow to yellow-orange stalk. The gills
have a somewhat waxy look (leading to possible confusion with Hygrocybe and Camaro-
phyllus), but are characteristically “chanterellesque”: thick, blunt, shallow, and con¬
spicuously forked or veined and/or wavy. The name C. tubaeformis has been used for
the white-spored variety, but many cantharellologists treat it as a synonym for C. infun¬
dibuliformis. In eastern North America there are numerous other small, edible, hollow¬
stemmed chanterelles, including: C. ignicolor, very similar but with a brighter cap when
fresh(orange to yellow-orange to ochraceous-tawny or salmon-buff), growing underboth
hardwoods and conifers; C. minor, small and entirely yellow to orange (including the gills),
favoring hardwoods; and C. xanthopus (also called C. lutescens), widely distributed
(also found in the West), similar to C. infundibuliformis in cap color or brighter, but with
a smooth to slightly veined (rather than gilled) orangish to yellowish underside.

Craterellus cinereus (Black Chanterelle)

FRUITING BODY more or less funnel- or vase-shaped but not tubular; 3-12 cm high.
CAP 1.5 -5 cm broad, shallowly to deeply depressed, surface smooth or minutely scaly-
scurfy, not viscid; black when moist, becoming dark grayish-brown inage orasitdries out;
margin usually wavy, lobed, or torn. Flesh thin, tough, colored more or less like cap.
UNDERSIDE (fertile surface) with thick, well-spaced, shallow, blunt, foldlike gills which
fork frequently near the cap margin and/or have cross-veins; bluish-black becoming
bluish-gray to gray, or paler from spore dust. ST ALK 2-8 cm long,0.4-1.3 cm thick, nearly
equal or tapering downward, central or off-center, tough, hollow except at base, colored
like cap or underside. SPORE PRINT whitish; spores 8-11 * 5-6 microns, elliptical,
smooth.
HABITAT: Solitary to scattered or in groups or clusters in mixed woods and under oaks;
widely distributed but not common. In ourarea itfruits in the winterand early spring, often
in the company of C. cornucopioides, or a few days before it.
Craterellus cinereus, a bluish-gray to black species with veins or shallow gills on underside of cap.

EDIBILITY: Edible and delectable—as good or even better than C. cornucopioides. It is

delicious simply sauteed in butter.
COMMENTS: This species, which has also passed under the name Cantharelluscinereus,
closely mimics its commoner cousin, Craterellus cornucopioides, but has primitive gills
on the underside of the cap and is not as brown in dry weather. Other species: C. caeruleo-
fuscus of eastern North America is similar, but has smaller spores; C. foetidus, also
eastern, has a sickeningly sweet odor, veined fertile surface, and thicker stalk (1-3
cm thick at apex); C. (-Pseudocraterellus) sinuosus has a gray to dark grayish-brown cap,
slightly larger spores, and a veined or copiously wrinkled (but not gilled) underside that
is grayish at first but acquires a distinct yellowish or ochre tinge as the spores mature; it
is edible and widely distributed, but rather rare (Craig Mitchell has found it in mixed
woods near Felton, California, in February).

Craterellus cornucopioides (Horn of Plenty) Color Plate 182

FRUITING BODY 3-14 cm high, tubular becoming trumpet- or funnel-shaped. CAP
2-8 (10) cm broad, hollowat the center with the marginatfirst decurved (folded down), then
spreading outand becoming wavy, split, or lacerated (sometimes witha lacy appearance or
convoluted to form small “suction cups”); surface not viscid, usually minutely scaly or
scurfy, grayish-black to very dark brown or black when moist, paler (brown to grayish-
brown) when dry (but one form sometimes developing a yellowish margin or
yellowish blotches). Flesh thin, brittle but tough, colored like cap or paler; odor pleasant.
UNDERSIDE (fertile surface) smooth to uneven or with slight, deeply decurrent wrinkles
(but not gills); colored like cap but usually paler or grayer, in age or dry weather with a
whitish to yellowish or buff coating of spore dust. STALK 1-5 cm long, 0.5-1 (1.5) cm thick,
continuous with cap and tapering downward, central or off-center, hollow except at very
base, tough, often twisted; colored like cap or underside. SPORE PRINT whitish to buff
or pale yellow; spores 8-11 * 5-7 microns, elliptical, smooth.
HABITAT: Scattered or in groups, clumps, or lacy clusters in woods; widely distributed.
It is common in our area in the winter and spring under hard woods(live oak, tanoak, man-
zanita, madrone, huckleberry), but farther north it fruits in the late fall under conifers.
South of San Francisco it rarely appears before Christmas, usually peaks at the same time
that the fetid adder’s tongue blooms, and may fruit on into April. In eastern North America

666
Craterellus cornucopioides. This delicious mushroom is easily recognized by its dark trumpetlike
fruiting body. The undersides of these mature specimens are dusted white or buff by spores. Note
complete absence of gills.

it is largely replaced by C.fallax{see comments), which also occurs in our area. Though
terrestrial, it often fruits near fallen branches or at the bases of trees or manzanita burls.

EDIBILITY: One of my fifteen “five favorite flavorful fleshy fungal fructifications.” Like
myself, it is thin, tough, and dark, and like myself, it goes largely unappreciated. It is
forever being passed up in favor of the larger, fleshier, more colorful and impressive types,
yet its flavor is superb and its potential unlimited. It is partly to blame for this sad state of
affairs, for its appearance is admittedly drab and rather somber, and it shuns attention,
blending unobtrusively into the dark, secretive situations where it thrives.
But though it takes an accomplished eye to detect its presence in the woods(from the top
it looks like a hole in the ground), anyone can detect its presence in a dish—for it cooks up
black, announcing itself with unmistakable earthy authenticity. It can play any culinary
position with equal finesse, from first base to second fiddle, enhancing practically any dish,
be it soup, souffle, or sauce. Like its more popular cousin the chanterelle, it is not often

Craterellus cornucopioides (right) often grows in clusters, blending into its surroundings so
uncannily that you have to search for black holes in the ground, as shown on left (actually a photo of
Craterellus fallax, essentially the same but for its salmon-tinged spores).
668 CANTHARELLACEAE

inhabited by maggots. Dried and powdered it has a cheesy odor and is known, quite
appropriately, as “Poor People’s Truffle.”
COMMENTS: This cryptically-colored fungus usually occurs in large groups. It is hard to
pick out in the forest gloom, but once you locate one, you’re bound tofind more. In France
it is sometimes called trumpet de mort (trumpet of death)—a tribute to its somber ap¬
pearance, not its edibility. Actually, it is quite lovely when fresh—fully-grown specimens
look more like black or brown petunias than anything else, and clusters often have a lacy
look. The horn of plenty that is so common in the summer and fall in eastern North Amer¬
ica, C. fallax, looks and tastes identical but has a salmon- or yellow-tinted underside in
age and longer, salmon or ochre-yellow spores; it also occurs in California. Otherwise,
there is little that C. cornucopioides can be confused with—the dark trumpet-shaped
fruiting body with smooth or slightly wrinkled exterior and hollow center is unique. C.
cinereus (the black chanterelle) is somewhat similar but has primitive gills; Polyozellus
multiplex is also similar but is dark blue or violet-tinted, while C. sinuosus (see comments
under C. cinereus) has yellowish-ochre spores and a more copiously wrinkled or veined
hymenium (underside). Several Ascomycetes are black, but not trumpet-shaped
(Plectania species are cup-shaped; Urnula is urn-shaped). Thelephora terrestris is
sometimes funnel-shaped but is thinner, smaller, and purple-brown rather than gray or
black. Occasionally hollow, deformed, clublike specimens of C. cornucopioides occur
alongside normal ones. These are identical in color and edible. Other species: C. calyculus
of eastern North America is one of several similar but much smaller and thinner(cap less
than 2 cm broad, stalk 1-3 mm thick), solid-stemmed, mostly southern species.

Polyozellus multiplex (Blue Chanterelle) Color Plate 183

FRUITING BODY usually compound—in compact, clustered masses 5-15 cm high and
up to 1 meter broad (but usually smaller). CAP(s) 2-10 cm broad, spoon-shaped to fan¬
shaped, often centrally depressed and lop-sided; surface dull purple to purple-gray to
black, deep blue, or frosted blue; margin usually irregularly lobed or wavy. Flesh deep
violet to bluish-black, soft and rather brittle; odor mild or fragrant. UNDERSIDE(fertile
surface) nearly smooth or more often with shallow, crowded wrinkles or veins or at times
nearly poroid (but lacking gills); colored more or less like cap or slightly paler. STALK(s)
2-5 cm long, 0.5-2 cm thick, central to off-center or lateral, usually short and oftenfusedat
base; colored more or less like cap or underside, often grooved; solid or becoming hollow.
SPORE PRINT white; spores(4) 6-8.5 x 5.5-8 microns, round to broadly elliptical, warted
or appearing angular. Cap tissue staining olive-black in KOH.
HABITAT: Clustered on ground under conifers (usually spruce and fir) or aspen in late
summer and fall; known only from northern and montane North America. It is usually
listed as rare, but is locally common in the Rocky Mountains. I have seen it in the Oregon
Cascades and it occurs very rarely in northern California. It was originally described
from Maine, and the color photograph was taken near Santa Fe, New Mexico, in an
aspen-spruce forest, in August.
EDIBILITY: Edible, and according to some sources, delicious. I have tasted it only once—
the texture was reminiscent of Craterellus, but the flavor was far inferior.
COMMENTS: This strikingly beautiful mushroom is easily told by its unusual deep
violet to bluish-black color and clustered growth habit. It might be mistaken at first glance
fora Craterellus, but the caps are spoon-to fan-shaped rather than tubular or trumpetlike.
The warty-angular spores suggest a closer relationship to the genus Thelephora (or per¬
haps to the teeth fungi) than to other chanterelles, but it has traditionally been treated
as a chanterelle because of its superficial resemblance.
 669

Jelly Fungi

TREMELLALES & Allies spores

Small to medium-sized fungi found mostly on wood. FRUITING BODY typically gelatinous or
rubbery when fresh; variously shaped, but most often lobed, convoluted, or bloblike. SPORE
BEARING SURFACE smooth, warty, lobed, or with small spines, covering the outer or lower
surfaces of the fruiting body. SPORE PRINT white to yellowish, but difficult to obtain. Spores
smooth, sometimes septate. Basidia septate or forked.

JELLY fungi differ fundamentally from other Basidiomycetes in the structure of their
basidia, which are partitioned (septate) or forked rather than simple and clublike. The
rust and smut fungi also have partitioned basidia, butarenotconsidered to be mushrooms.
Although the basidia are microscopic, jelly fungi can usually be told in the field by the
gelatinous (jellylike) or rubbery texture that gives them their name. The most familiar
types are collectively called “witch’s butter.” They have lobed to convoluted or rather
amorphous (shapeless) fruiting bodies that look—at least to the hungry mushroom hunter
staggering home under a basketful of boletes—like lumps of melting butter. Other jelly
fungi have a cap and stalk, still others are cuplike, and a few are tough, branched, and coral¬
like. The spore-bearing surface ranges from smooth to veined, lobed, or in one case,
toothed, and is always exposed (hence the jelly fungi are classed here as Hymenomycetes).
The design of the basidium (see illustration) forms the basis for dividing the jelly fungi
into three separate groups. The Tremellales, which are the most common, have obliquely
or longitudinally septate basidia that look like hot cross buns when viewed from above, the
Auriculariales have transversely septate basidia (i.e., with cross walls), and the Dacrymy-
cetales have Y-shaped basidia that look like tuning forks. As these distinctions are micro¬
scopic, the three groups are treated as one unit in this book.

U Different types of basidia in the jelly fungi.

Left to right: longitudinally septate
(Tremellales); Y-shaped (Dacrymy-
cetales); and transversely septate (Auri¬
culariales).

Most of the jelly fungi grow on rotten wood. They thrive in cool wet weather, shrinking
down to almost nothing when it dries out, then swelling up again as soon as it rains. None
are known to be poisonous, but most are very watery and flavorless. Some species,
however, can be marinated, candied, or even eaten raw, and in the Orient two types are
very popular: a translucent white species called Tremella fuciformis, and the brown to
black “tree ears” or “cloud ears” (Auricularia). The latter can be purchased in dried form
in many specialty stores. Eight jelly fungi are described here and several others are keyed
out. Tremella mesenterica is the most common of the lot, at least in our area.

Key to the Tremellales & Allies

1. Fruiting body brightly colored (yellow, orange, pink, red, or greenish) when fresh, but sometimes
losing its color in rainy weather or old age . 2
1. Not as above; fruiting body white, grayish, black, reddish-purple, brown, yellow-brown, etc. 9
2. Fruiting body spatula-shaped to funnel-shaped (but with one side usually incised or split),
pink to orange or red; stalk usually present .Phlogiotis helvelloides, p. 672
2. Not as above . 3
670 TREMELLALES & ALLIES

3. Fruiting body erect, either simple and clublike (unbranched) or antlerlike or branched, or with a
cap and stalk . 4
3. Not as above; fruiting body cup-shaped to cone-shaped, cushion-shaped, irregularly lobed and
contorted, or amorphous (bloblike) . 6
4. Fruiting body with a yellow-orange to greenish “head” or cap .(see Helotiales, p. 865)
4. Not as above; fruiting body simple and unbranched or antlerlike to coral-like . 5
5. Fruiting body branched, usually more than 15 mm high .Calocera viscosa, p. 674
5. Fruiting body unbranched (but may be clustered), or if sparingly branched then typically less
than 15 mm high .Calocera cornea (see C. viscosa, p. 674)
6. Fruiting body cup-shaped to cone-shaped (with a narrowed base or point of attachment); usually
found in the spring (often near melting snow) . Guepiniopsis alpinus & others, p. 674
6. Not as above; fruiting body bloblike to cushion-shaped to irregularly lobed or brainlike . . 7
7. Fruiting body 1-5 mm broad, drop-like, usually growing in large masses or rows .
. Dacrymyces deliquescens (see Tremella mesenterica, p. 673)
7. Not as above; fruiting body usually larger . 8
8. Typically growing on hardwoods; bone-hard when dry; basidia shaped like hot-cross buns in
top view .Tremella mesenterica, p. 673
8. Usually growing on conifers; collapsing when dry, but with a tough whitish basal point of
attachment; basidia Y-shaped . . . Dacrymyces palmatus (see Tremella mesenterica, p. 673)
9. Fruiting body translucent to whitish, grayish, or brownish, with a cap (and usually a stalk),
the underside of the cap lined with tiny spines or “teeth” Pseudohydnum gelatinosum, p. 671
9. Not as above; underside of cap lacking minute spines or “teeth” . 10
10. Fruiting body tough, erect, and usually branched (coral-like), white or pallid; found mainly on
ground under hardwoods in eastern North America . 11
10. Not as above; usually found on wood or plants . 12
11. Branches few and very thick, blunt, hollow, and somewhat gelatinous . . . Tremella reticulata
11. Not as above; texture very tough, branches often flattened .
.Tremellodendronpallidum & T. candidum (see Tremellodendropsis tuberosa, p. 643)
12. Fruiting body black (or nearly black) when fresh . 13
12. Not as above (but fruiting body may be dark brown and/ or may blacken as it dries out) . 14
13. Fruiting body small and cushion-shaped, warty, or lobed, but often fusing to form large con¬
tinuous patches or sheets; growing on dead hardwoods .Exidia glandulosa, p. 672
13. Not as above; fruiting body broadly top-shaped to cuplike and not fusing to form sheets . . . .
. (see Pezizacaceae & Allies, p. 817)
14. Fruiting body a gelatinous mass of wavy or leafy lobes up to 20 cm broad (or sometimes larger),
some shade of brown; spores borne on basidia . Tremella foliacea, p. 673
14. Not as above (if brown and leafy, then fruiting body thin and rubbery instead of gelatinous) 15
15. Fruiting body reddish to brown or dark brown and bracketlike to cuplike, ear-shaped, broadly
top-shaped, or occasionally forming a cluster of earlike lobes . 16
15. Not as above . 17
16. Fruiting body cup-shaped to broadly top-shaped, reddish to purplish to brown or dark brown;
flesh thick or thin; usually on hardwoods; spores borne in asci .(see Helotiales, p. 865)
16. Fruiting body bracketlike to earlike or cuplike (or with several earlike lobes); flesh thin; on
hardwoods or conifers; spores borne on basidia .Auricularia auricula & others, p. 675
17. Fruiting body containing whitish granules; overall color whitish becoming pinkish- to reddish-
brown or vinaceous-brown . Exidia nucleata (see E. glandulosa, p. 672)
17. Not as above . 18
18. Fruiting body white or pallid when fresh (may be tinged tan in age) . 19
18. Fruiting body not white or pallid . 21
19. Fruiting body gelatinous or tough and usually hollow; growing on herbaceous stems and other
vegetable matter in eastern North America .Tremella concrescens
19. Not as above; usually growing on wood; widespread . 20
TREMELLALES & ALLIES 671

20. Fruiting body lobed or convoluted, 1.5-7 cm broad (or merging to form larger patches up to
12 cm or more); spores elliptical; found in the southern U nited States and warmer parts of the
world (including the Orient) . Tremellafuciformis
20. Not as above; smaller and/ or spores sausage-shaped . Exidia alba (see E. glandulosa, p. 672)
21. Fruiting body more or less brownish-yellow and top-shaped or cone-shaped .
. Exidia recisa (see Guepiniopsis alpinus, p. 674)
21. Fruiting body bloblike to lobed, convoluted, or brainlike and flesh-colored to brownish,
yellowish, or buff. Tremella encephala & T. frondosa(see T. foliacea, p. 673)

Pseudohydnum gelatinosum (Toothed Jelly Fungus)

FRUITING BODY flexible and flabby, rubbery-gelatinous, more or less tongue-shaped
to spoon-shaped or fan-shaped (with a cap and usually a stalk). CAP 1-6 (7.5) cm broad;
surface minutely roughened or downy to nearly smooth, not viscid; translucent white to
watery gray or bluish-gray, or sometimes dingy brownish (or spotted brown). Flesh
rubbery-gelatinous. UNDERSIDE lined with small pallid spines or “teeth” that are 0.5-3
(5) mm long. STALK up to 6 cm long, usually lateral, continuous with cap and similar in
color and texture. SPORE PRINT white; spores borne on the “teeth,” 5-8.5 microns,
round or nearly round, smooth. Basidia longitudinally septate, elliptical to nearly round.
HABITAT: Solitary, scattered, or gregarious on rotting logs, twigs, and humus under
conifers; widely distributed. It is sporadically common in our area in the late fall and winter,
especially in dank, dark, damp situations under Douglas-fir.
EDIBILITY: Edible. It is said to be fairly good with honey and cream—but what isn't! It
can also be marinated for use in salads. The texture is interesting, the flavor nonexistent.
COMMENTS: Also known as Tremellodon gelatinosum, this denizen of dank places is
one of my fifty “five favorite fleshy fungal fructifications.” The rubbery or flabby tongue¬
shaped fruiting bodies with small “teeth” on the underside are as attractive as they are
unique, and look funnier than they do fungal—in fact, it is hard to take them seriously!
(They remind me of the “Creepy Crawlers” I used to buy at the dime store with my lunch
money.) The small teeth resemble those of the teeth fungi (Hydnaceae), hence the name
Pseudohydnum, which means “false tooth fungus.” Specimens on the west coast tend to
have a well-developed, often vertical stalk whereas those in eastern North America tend
to have little or no stalk, especially when growing from the sides of logs.

Pseudohydnum gelatinosum. Underside (left) is lined with tiny “teeth.” Cap (right) is whitish to
grayish or brown. Entire fruiting body is rubbery.
Left: A mature, flabby specimen of Phlogiotis helvelloides. Note how it is split down one side. Right:
Exidia glandulosa, whose small fruiting bodies often fuse to form black rubbery or gelatinous sheets.

Phlogiotis helvelloides (Apricot Jelly Mushroom)

FRUITING BODY flabby or rubbery, with a cap and stalk; spatula-shaped to somewhat
funnel-shaped, but usually indented or split down one side; 2-8 (18) cm high. CAP 1-7(10)
cm broad, pale to deep rosy-pink to reddish-orange, apricot , or salmon-colored; surface
more or less smooth, margin often wavy. Flesh rubbery to somewhat gelatinous.
UNDERSIDE smooth to faintly veined or coarsely wrinkled, colored like cap or paler.
STALK 1 -6 cm long, off-center or lateral but usually vertical (upright); continuous with
cap, with the same color and texture, or the base whitish. SPORE PRINT white; spores
borne on underside of cap, 9-12 (16) x 4-6.5 microns, oblong-elliptical, smooth. Basidia
longitudinally septate.
HABITAT: Scattered to gregarious on ground, debris, and rotten wood under conifers;
widely distributed, but uncommon in most regions. In California I have seen impressive
fruitings in Yosemite National Park and the Trinity Alps, but have found it only once in
our area, near Big Sur. The fruiting bodies do not decay readily and seem to like cool
weather (either late in the season or in the spring).
EDIBILITY: Highly prized by some people—presumably for its color and texture—for it
has no flavor that I can detect. It can be eaten raw in salads, pickled, or candied, but should
not be cooked because of its high water content. Larry Stickney says if it is sucked on a
hot day it “cools you right down.”
COMMENTS: The pink to orange color, rubbery texture (similar to that of Pseudo-
hydnum gelatinosum), and overall shape—which is sometimes reminiscent of a miniature
calla lily—identify this fetching little fungus. Actually, the word “little” is not always
appropriate, as specimens 7 inches (18 cm) high have been reported by Sam Ristich of New
York. These apparently developed over a period of two months and had smaller ones
riding “piggy-back” on top of them! The normal size range, however, is 1-3 inches high.

Exidia glandulosa (Black Witch’s Butter; Black Jelly Roll)

FRUITING BODY flabby or gelatinous, beginning as a pallid or translucent blister but
soon becoming cushion-shaped to irregularly lobed; reddish-black to olive-black soon
becoming jet-black (or black from beginning); 1-2 cm broad but often fusing with others
to form rows or masses up to 50 cm long; upper surface smooth to minutely roughened or
warty. Flesh gelatinous, black. STALK absent. SPORE PRINT whitish; spores borne on
the lobes, warts, or wrinkles, 10-16 * 3-5 microns, sausage-shaped, smooth. Basidia longi¬
tudinally septate.
EXIDIA 673

HABITAT: Scattered to densely gregarious or in fused rows and masses on rotting hard¬
wood logs and branches; widely distributed. It is fairly common in our area throughout the
mushroom season, but is usually overlooked because of its dark color.
EDIBILITY: Unknown, and like most of us, likely to remain so.
COMMENTS: This sinister-looking black jelly fungus can be confused with little else.
Individual fruiting bodies are quite small but usually fuse to form rows or sheets that look
like black gelatin or slime. The frequently roughened spore-bearing surface and sausage¬
shaped spores help distinguish the genus Exidia from Tremella. Other species: E. alba is
small and whitish; E. nucleata varies from white or colorless to pinkish- or vinaceous-
brown, but is sprinkled with small white granules or “nuclei.” Both are widely distributed.

Tremella foliacea (Brown Witch’s Butter) Color Plate 173

FRUITING BODY flabby or gelatinous when moist, bone-hard when dry; 2.5-20 cm
broad or more, typically consisting of a complicated mass of wavy or leaflike folds, lobes,
and convolutions; reddish-cinnamon to brown, vinaceous-brown, or tinged purple; often
paler when waterlogged. Flesh gelatinous. STALK absent. SPORE PRINT white to
yellowish; spores produced on the upright lobes, 7-9 (13) * 6-9 microns, round to broadly
elliptical, smooth. Basidia longitudinally septate.
HABITAT: S olitary or several on hardwood stumps, logs, and fallen branches (occasion¬
ally on conifers as well); widely distributed. In our area it is not uncommon on oak in the late
fall, winter, and early spring, but is not nearly as numerous as T. mesenterica.
EDIBILITY: Harmless, but mostly water. As for the flavor—well, I once encountered an
Asian-American gathering this species and T. mesenterica in a live oak woodland. “How
do you prepare them?” I said. “With garlic and soy sauce,” she said. “And how do they
taste?” I said. Slight hesitation. “Like garlic and soy sauce,” she said.
COMMENTS: This brown version of T. mesenterica is our largest Tremella and is quite
striking and seaweed-like when fresh. The fruiting body is never cuplike or earlike as in
Auricularia auricula, and is much more gelatinous. T. frondosa is a somewhat similar
yellowish to buff species. T. encephala is also similar but smaller (1-6 cm) and flesh-colored
to brownish. It is thought to be parasitic on Stereum, a wood-inhabiting bracket or parch¬
ment fungus. M ost Tremellas, in fact, are frequently found with Stereum, suggesting some
kind of relationship between the two. The lobed, brainlike, or convoluted fruiting bodies of
Tremellas are reflected in their names: foliacea means “leaflike”; mesenterica means
“middle intestine”; and encephala means “brainlike.”

Tremella mesenterica (Witch’s Butter) Color Plate 170

FRUITING BODY flabby or gelatinous when fresh, bone-hard when dry; 1-10 cm broad,
typically consisting of several to many convoluted or brainlike lobes or folds, but in wet
weather or old age often bloblike or amorphous; clear yellow to golden-yellow to bright
orange, paler (to nearly colorless) when old or waterlogged. Flesh gelatinous. STALK
absent. SPORE PRINT pallid to yellow; spores produced on the upright lobes or folds,
7-18 x 6-14 microns, elliptical to nearly round, smooth. Basidia longitudinally septate.

HABITAT: Solitary or in groups on hardwood sticks, logs, etc.; very common and wide¬
spread. In our area it favors dead oak and fruits throughout the mushroom season, often
on the same branches as Stereum species. Shrivelled up specimens are inconspicuous, but
usually revive in wet weather. In colder climates it may appear during winter thaws.
EDIBILITY: Harmless but flavorless (see comments on edibility of T. foliacea)—it is
mostly water. My one attempt at cooking it was a failure: most of it evaporated!
 674 TREMELLALES & ALLIES

COMMENTS: Also known as T. lutescens, this fungus and its look-alikes (see below) are
a familiar sight in rainy weather across the continent. At first the fruiting body is lobed or
brainlike and relatively firm, but as it swells with water it often loses its original shape and
looks like a dollop of melting butter on a log. Also common is Dacrymyces palmatus,
which closely mimics T. mesenterica but is slightly smaller (1-6 cm broad), tends to be
oranger in color, favors conifers, and has a small, tough, whitish, basal point of attachment.
The resemblance is only superficial, however, for Dacrymyces species have Y-shaped
basidia and long narrow spores that develop cross walls, while Tremella species have
longitudinally septate basidia and simple (non-septate) spores. Other species: T.frondosa
is buff to yellowish but larger and leafier (like T. foliacea); D. deliquescens is a small (1-5
mm broad), yellow-orange to reddish species that grows on decaying wood, usually in
large masses or rows. Like most jelly fungi, the above species are not worth eating.

Guepiniopsis alpinus (Alpine Jelly Cone; Poor Man’s Gumdrop)

FRUITING BODY flabby or gelatinous; top- to cone-shaped or sometimes cup-shaped,
0.3-2.5 cm broad; yellow to orange or occasionally reddish-orange; surface smooth. Flesh
gelatinous. STALK present only as a small, narrowed point of attachment. SPORE
PRINT yellowish; spores borne on the concave surface of the cone, 11-18 * 4-6 microns,
sausage-shaped, developing 3-4 cross walls. Basidia Y-shaped.
HABITAT: Scattered to gregarious on coniferous logs, stumps, branches, twigs, etc.;
fruiting in the spring (usually as or just after the snow melts) at higher elevations from the
Rocky Mountains westward (rarely in the summer or fall); sometimes abundant.
EDIBILITY: Presumably consumable, but of negligible value.
COMMENTS: The golden cone-shaped fruiting body and fondness for dead conifers
typify this omnipresent “snowbank” mushroom of western mountains. The gelatinous
texture distinguishes it from the cup fungi, and the gumdrop-like fruiting body separates
it from Tremella and Dacrymyces. The Y-shaped basidia place it in the Dacrymycetales,
alongside Dacrymyces and Calocera. G. chrysocomus is a closely related species with a
yellow cup-shaped fruiting body, larger spores, and a preference for hardwoods. Exidia
recisa resembles G. alpinus in shape, but has longitudinally septate basidia and is brownish-
yellow in color; it grows on hardwoods and is exceptionally abundant in some regions
(especially the southern United States).

Calocera viscosa (Staghorn Jelly Fungus)

FRUITING BODY erect, branched (antler- or coral-like); 2-7 (10) cm high, 1-3 (5) cm
wide. BRANCHES tough, pliant, usually somewhat viscid, the tips usually forked (with

Left: Guepiniopsis alpinus looks like a gumdrop. Right: Calocera viscosa. (Herb Saylor)
CALOCERA 675

2-3 tips); bright yellow to deep yellow to orange. ST ALK present as a yellow to whitish base
or “trunk”, often rooting. Flesh tough but often somewhat gelatinous. SPORE PRINT
dingy yellowish to ochraceous; spores produced on the branches, 9-14 * 3-5 microns,
elongated-elliptical to sausage-shaped, smooth, developing one cross wall at maturity.
Basidia Y-shaped.
HABITAT: Solitary, scattered, or in small groups on or near coniferous logs, stumps,
roots, and debris; widely distributed. I have found it several times in our coastal pine
forests in the winter and spring, but never in quantity.
EDIBILITY: Inconsequential.
COMMENTS: This dainty little jelly fungus looks like a coral mushroom (especially
Clavulinopsis corniculata) and is keyed out as one. However, it is usually viscid and has
Y-shaped basidia, plus it typically grows on or near wood. Another widespread species,
C. cornea, is smaller (up to 15 mm high), yellow, and clublike or very sparingly branched;
it grows in groups or clusters on twigs and branches of both hardwoods and conifers. The
forked basidia place Calocera in the Dacrymycetales.

A uricularia auricula (Wood Ear; Tree Ear)

FRUITING BODY rubbery to pliant or flabby to somewhat gelatinous when fresh, hard
when dry; 2-15 cm broad; cup-shaped to ear-shaped or sometimes with several earlike
lobes originating from a central point of attachment. Outer surface sterile, often
veined or ribbed, minutely silky or with fine downy hairs, pale brown to brown to liver-
brown, drying blackish. Inner (fertile) surface smooth to slightly wrinkled, somewhat
gelatinous when wet, tan to yellow-brown, grayish-brown, brown, liver-brown, or tinged
purple; blackish when dried. Flesh thin, rubbery. STALK absent or rudimentary. SPORE
PRINT white; spores borne on inner (usually the lower) surface, 12-18 * 4-8 microns,
sausage-shaped, smooth. Basidia transversely septate (with 3 cross walls).

HABITAT: Solitary or in groups or clusters on logs, dead branches, stumps, etc. (attached
centrally or laterally); very widely distributed on both hardwoods and conifers, and often
common in cool weather. I have seen it in the springtime in the Cascades, but it seems to
be rare in our area.
EDIBILITY: Edible, but I haven’t tried it. A similar but hairier species, A. polytricha, is
prized in the Orient and can be purchased dried in many stores. Along with the shiitake
(Lentinus edodes) it is the mushroom most often served in Chinese restaurants, usually
under the name Y ung Ngo or Muk Ngo. Dried specimens of A. auricula and A. poly tricha
are quite dark, hard, and unappetizing, but billow up like clouds when soaked in water,
showing off their delicate curves and convolutions to great effect.
COMMENTS: This species can be mistaken for a cup fungus, but has a more rubbery
texture, bears its spores on basidia (a microscopic feature), and usually—but not always—
grows with its fertile (concave) surface facing downward. Cup fungi, on the other hand,
bear their spores in asci with the concave (fertile) surface facing upward, and usually have
a brittle or fragile texture. Sometimes Auricularia is irregularly lobed rather than cuplike,
and I have found old specimens which looked more like pieces of soggy seaweed than
anything else. Hirneola auricula-judae is a sinister-sounding synonym. It also used to be
called “Judas’ Ear” because it was believed that when Judas’ hanged himself on an elder
tree, these ear-shaped “excrescences” were condemned to appear on elders thereafter.
Auricularia is the most prominent genus in the Auriculariales. It embraces several similar
brownish species, including: A. mesenterica, which is more or less bracketlike with a
hairy, concentrically zoned sterile surface and veined fertile surface; and A. delicata,
a striking gelatinous tropical species with large honeycomb-like pits or “pores.” For a
photo of Auricularia, see p. 958.
676

GASTEROMYCETES
THIS large division of the Basidiomycetes includes those fungi better known as puffballs,
earthstars, stinkhorns, bird’s nest fungi, false truffles, and gastroid agarics. Gastero-
(meaning “stomach”) describes the manner in which the spores are produced—internally,
rather than externally as in the Hymenomycetes (agarics, boletes, coral fungi, etc.). The
Gasteromycetes are also unique in that their spores are not forcibly discharged. Instead
the basidia disintegrate and the leftover spore mass is dispersed by wind, rain, and animals.
The most familiar fungi in this group are the puffballs and earthstars. They bear spores
in a round to oval “stomach” or spore case. The mature spore mass is powdery and easily
dispersed. The false truffles are similar, but their spore mass remains intact and does
not become powdery, while the gastroid agarics resemble gilled mushrooms that haven’t
opened. The stinkhorns strike a decidedly different pose—their slimy spore mass is
initially enclosed by a membrane, but later it is elevated on a stalk, arms, or latticed ball,
with the membrane forming a volva at the base. Last and least, there are the bird’s nest
fungi, which look like tiny nests with spore-containing eggs or peridioles.
Gasteromycetes occur in a wide range of habitats. They are especially prominent in
arid regions where there is a selective advantage to producing spores internally (it affords
protection from moisture loss and heat). They are notable more for their size range (giant
puffballs weigh up to 50 pounds each, bird’s nest fungi are only a few millimeters broad)
and different strategies for spore dispersal than for their colors. Only the puffballs are
of importance to the mushroom-eater. Six major groups are keyed below.*

Key to the Gasteromycetes

1. Fruiting body minute (typically less than 15 mm high), consisting of a “nest” (cup, vase, or bowl)
containing one or more “eggs” (peridioles); (older specimens, however, may lack peridioles
and young ones often have a covering or “lid” over the top of the “nest”) Nidulariales, p. 778
1. Fruiting body differently constructed and usually larger than above .2
2. Fruiting body at first enclosed in a membrane, then emerging as a cylindrical, phallic, branched,
tentacled, or latticed structure with the membrane forming a volva or sack at the base; mature
spore mass slimy or mucilaginous (never powdery) and usually foul-smelling, coating the
head, branches, or latticework of the fruiting body . Phallales, p. 764
2. Not as above (but fruiting body may be slimy or malodorous at some stage).3
3. Fruiting body with a stalk below the spore case or “cap” .4
3. Stalk absent or rudimentary . 7
4. Stalk penetrating the spore case or “cap” and usually percurrent (i.e., extending to top of
fruiting body); stalk not normally branched inside spore case . . . Podaxales & Allies, p. 724
4. Not as above; stalk not percurrent, sometimes branched . 5
5. Spore mass slimy and greenish or olive, divided into several large chambers; fruiting body
club-shaped to pear-shaped; usually found on rotten wood Hymenogastrales & Allies, p. 739
5. Not with above features . 6
6. Stalk well-developed and clearly defined, almost always longer than diameter of spore case;
stalk not composed of tough rootlike fibers, or if so then fruiting body usually brightly colored
and/or gelatinous .Tulostomatales, p. 715
6. Not as above; stalk a tough mass of “roots” or fibers (and sand or dirt) or stalk an elongated,
narrowed sterile base which often shows minute chambers when sliced open lengthwise .. 7
7. Spore case rupturing or disintegrating at maturity; spore mass firm and solid when young
(chambers if present hardly discernible), powdery or cottony when mature and usually dis¬
persing fairly soon; columella (internal stalk) typically absent; mature fruiting body usually
(but not always) above the ground; found in many habitats .. Lycoperdales& Allies, p. 677
7. Spore case and spore mass remaining intact fora long time; spore mass often chambered (cham¬
bers large to minute), firm, rubbery, spongy, or slimy but not cottony or powdery at maturity;
columella present or absent; usually growing underground in association with trees or shrubs,
but sometimes surfacing at maturity .Hymenogastrales & Allies, p. 739
*Some agarics, boletes, or other mushrooms superficially resemble puffballs, especially when young. Examples are
Entoloma abortivum, Amanita“eggs,” parasitized Boletus spp., and Cryptoporus volvatus (a common polvpore).
 677

Puffballs and Earthstars

spores

LYCOPERDALES & Allies

MANY people don’t think of puffballs and earthstars as mushrooms, and indeed they
have little in common with the cap-gill-and-stem commodity you buy at the grocery store.
The fruiting body simply consists of a roundish to oval spore case, sometimes with a sterile
region beneath it called, quite appropriately, the sterile base. The skin (peridium) of
the spore case is usually composed of an inner and outer layer (endo- and exoperidium).
In the earthstars, the outer skin separates completely from the spore case and splits into
several starlike rays; in the puffballs it does not. The sterile base, which is composed of
minute compartments that give it a spongelike appearance, is best seen by slicing open the
fruiting body lengthwise. In some species the sterile base is absent or inconspicuous;
in others it is as large as the spore case and narrowed below so that it looks like a stem.
However, only the stalked puffballs (discussed on p. 715) have a true stem.
The interior of the spore case is called the spore mass (or gleba). It is usually white and
firm when young, but turns yellow, greenish, brown, or purplish as the spores mature,
first becoming mushy as moisture is released, then powdery or cottony as the moisture
evaporates. The spore color corresponds to that of the mature spore mass, and is usually
some shade of brown or purple. Once the spores have matured, the spore case either splits
open or ruptures irregularly or disintegrates—thereby exposing the spore mass to the
elements— or a mouth or slit (apical pore) forms at the top, so that the spore case looks and
acts like a miniature volcano. It has been suggested that the word “puffball” is a derivative
of “puckfist,” which in turn was derived from “pixie fart.” All of these names testify to the
puffballs’ distinctive method of spore dispersal. If a mature(ruptured) specimen is poked,
tapped, squeezed, or kicked (thereby duplicating the action of a raindrop or gust of wind),
a cloud or “puff’ of spore dust will emerge.
The true puffballs (Lycoperdales) also have several distinctive microscopic features.
The spores are typically borne in a hymenium(palisade of basidia) and are usually round,
sometimes with a pedicel (“tail”) attached. They are often warted or spiny like sea urchins,
and bounce around like furry bon-bons under the microscope. Usually intermingled with
the mature spores are microscopic threadlike cells called capillitium. These are the
remains of the hyphae on which the basidia form. Their shape and branching pattern are
of great significance to the puffball taxonomist, but are not emphasized here.

spore mass (gleba)

Vertical sections of two puffballs and one stalked puffball. Left to right: Calvatia{stalk and sterile
base absent), Tulostoma(stalk present), and Lycoperdon{sterile base present).
Make sure every puffball you eat is firm, white, and solid( homogeneous) inside. These are rather scaly
specimens of coastal California’s giant puffball (see Calvatia gigantea group, p. 682). (Joel Leivick)

Besides the Lycoperdales, which includes the true puffballs such as Calvatia and
Lycoperdon and the earthstar genus Geastrum, a superficially similar order, the Sclero-
dermatales, is traditionally included under the title “puffballs and earthstars.” This
group differs microscopically in lacking capillitium and/ or a hymenium. It includes
the earthballs (Scleroderma), which can usually be separated from puffballs by their
hard, one-layered peridium and purple-black spore mass, certain earthstars (e.g., As-
traeus), and two dusty monstrosities that bear their spores in small chambers within the
spore case (Pisolithus and Dictyocephalos). A third group, the Tulostomatales (stalked
puffballs), differs in possessing a true stalk. It is treated separately here, although micro¬
scopically it shares features of both the Lycoperdales and Sclerodermatales.
It is often said that all puffballs are safe to eat so long as they’re firm and white inside.
This is not necessarily the case, however. Several species of Scleroderma are poisonous,
some of the so-called edible puffballs have purgative effects on certain individuals, and
others don’t taste good. Therefore, it behooves you puffball-pickers to identify each
puffball before you eat it and to sample it cautiously. Once picked, puffballs must be
refrigerated or they will ripen rapidly. Any showing the slightest traces of color (usually
yellow or green) should be discarded, as they become bitter and indigestible. Also be
sure that you don’t inadvertently mistake a deadly poisonous Amanita ^ egg" fora puff¬
ball. When sliced open lengthwise (i.e., perpendicular to the ground), puffballs are solid
and homogeneous within (see above photo), whereas Amanita “eggs” or other agaric but¬
tons reveal the outline of cap, gills, and stalk (see photo at top of p. 679). False truffles are
sometimes mistaken for puffballs, but they usually grow underground and do not have a
powdery spore mass at maturity, while stinkhorn “eggs” are gelatinous within.
Puffballs and earthstars can be found almost anywhere at any time, but are especially
prominent in prairies, deserts, and high mountains, where other fungi are not so plentiful.
Though they form a distinctive group, identifying the various species can be difficult be¬
cause you often have to know the characteristics of both mature and immature fruiting
bodies, as well as microscopic features such as the size and shape of the spores and capil¬
litium. It is tempting to eat puffballs without bothering to identify them, but indiscrimi¬
nate sampling of any mushrooms—even puffballs—is poor practice, so at least make an
attempt to key out each kind you find, even if you have only young (or old) specimens at
hand. In the following key, the various genera in the Lycoperdales and Sclerodermatales
have been grouped into several categories.

678
Amanita''1’ eggs” can look like puffballs, but reveal a mushroom“embryo” of cap, gills, and stalk when
sliced open lengthwise (perpendicular to the ground). Although this Amanita calyptrata “egg” is
edible, the “eggs” of several Amanitas are deadly poisonous!

Key to the Lycoperdales & Allies

1. Outer layer(s) of fruiting body splitting into several starlike rays which unfold or bend under
(at least in wet weather) to expose the inner skin or spore case .
. Geastrum, Astraeus, & Myriostoma, p. 699
1. Not as above (fruiting body may rupture in starlike fashion, but if so then there is no separate
spore case within) . 2
2. Spore mass containing numerous minute peridioles (spore-bearing chambers) which look like
particles of sand; fruiting body small to medium-sized . 3
2. Spore mass not containing peridioles, or if so then the peridioles considerably larger than grains
of sand (usually appearing more like seeds) .4
3. Spore mass penetrated by a columella (internal stalk); fruiting body with an external stalk also,
but the stalk often falling off; found in deserts (often mingling with Endoptychum arizonicum)
.A raneosa columellata
3. Fruiting body puffball-like, i.e., lacking a columella and stalk; found mostly in sandy soil or
grassy or open places; widespread but not very common .Arachnion album & others
4. Mature fruiting body sticking out of the ground like a dusty root or half-rotted stump (but
may be roundish to pear- or club-shaped when young); spores produced inside numerous small
chambers or seedlike bodies (peridioles) imbedded in the fruiting body (the peridioles are best
seen by slicing open a young fruiting body because they soon disintegrate); usually found in
poor soil, along roads, in deserts, etc.Pisolithus& Dictyocephalos, p. 711
4. Not as above; peridioles absent; spores produced in a single large chamber (the spore case) 5
5. Spore case typically hard or tough with a thick rindlike skin, at least when young; spore mass
white when very young b ut soon darkening! usually purple-gray to black) while remaining firm,
eventually becoming dark brown to blackish and powdery; basidia not borne in a hymenium;
capillitium absent .Scleroderma, p. 707
5. Not as above; skin (peridium) thick or thin; spore mass white when young and normally soften¬
ing or becoming mushy as it darkens, then becoming powdery; basidia usually borne in a
hymenium; capillitium usually present .6
6. Fruiting body thick-skinned, not rupturing, usually underground; spore mass revealing a thick
short columella (internal stalk) when sectioned lengthwise through the center {but columella
sometimes disintegrating in old age)—see photos on pp. 761-762 . . . (sttRadiigera, p. 760)
6. Not as above . 7
7. Spore mass with prominent veins or cords running through it, the veins seeming to originate
from the base or peridium; rare .Scleroderma, p. 707
7. Not as above . 8
8. Sterile base present, often as a narrowed stemlike base beneath the spore case (section fruiting
body lengthwise if unsure) .9
8. Sterile base absent or rudimentary . 10

679
680 LYCOPERDALES & ALLIES

9. Fruiting body medium-sized to quite large, rupturing (in old age) irregularly or through radial
tears or general disintegration; peridium (skin) thick or thin .Calvatia & Allies, below
9. Fruiting body small to medium-sized (usually smaller than a baseball), typically rupturing
through an apical pore, slit, or large mouth; usually thin-skinned Lycoperdon& Allies, p. 690
10. Fruiting body golfball-sized to very large, rupturing (in old age) irregularly or through radial
tears or general disintegration; peridium (skin) thick or thin; found in many habitats .
.Calvatia & Allies, below
10. Fruiting body usually marble- to golfball-sized or occasionally as large as a baseball, usually
rupturing through a pore or large mouth (and often blowing about in the wind when old); peri¬
dium usually rather thin; found mostly in grassy or open places Bovista& Disciseda, p. 696

C ALV ATI A & Allies (Giant Puffballs)

Medium-sized to very large terrestrial puffballs. FRUITING BODY round to top- or pear-shaped
(broader above) to somewhat flattened. Peridium (skin) two-layered, disintegrating or rupturing
irregularly at maturity, smooth or warty. STERILE BASE present or absent. SPORE MASS firm
and white when immature, then slowly darkening to olive-brown, dark brown, or purple and
becoming powdery or cottony. Spores typically more or less round, smooth to warted or spiny.
Capillitium present.

THESE are baseball- to basketball-sized puffballs that disintegrate in old age, i.e., a dis¬
tinct apical pore is not formed as in Lycoperdon. In some species the outer layer of the
peridium takes the form of large warts which break up into plates and then flake off, but
in other species it is smooth and in many it adheres to the inner layer so that the two are
indistinguishable. Calvatias with a thick peridium are sometimes mistaken for earth-
balls (Scleroderma species), but are usually whiter (both inside and out) when immature
and not as hard-fleshed. The texture (whether powdery or cottony) and color of the
mature spore mass are important features in the identification of Calvatias, as is the
presence or absence of a sterile base.
Calvatias are among the most prolific of living organisms. It has been calculated that
an average-sized (30 cm) specimen of the giant puffball (Calvatia gigantea) contains
7,000,000,000,000 (7 trillion) spores! In these inflationary times that may not sound like
much, but consider this: if all 7 trillion spores (each one measuring 1/200 of a millimeter)
were lined up in a row, they would circle the earth’s equator! If each spore produced a
30 cm offspring, the resulting puffballs would stretch from the earth to the sun and back,
and if their spores were equally successful, the formidable puffball mass would weigh 800
times as much as the earth! Each spore is theoretically capable of germinating, yet very
few (obviously!) do. It would be interesting to know why so many don’t, or conversely,
why such a surplus of spores is (needlessly?) produced.
The truly giant puffballs (the C. gigantea group and C. booniana) are among the best
known and most popular of all edible mushrooms. In fact, they are eaten by people who
don’t know a gill from a gall. Some of the smaller species (e.g., C. sculpta and C. cyathi-
formis) are also excellent, but a few (e.g., C.fumosa) are bitter. None are known to be
poisonous, but the edible species have purgative or laxative effects on some people. Each
kind should be tested cautiously and eaten in moderation, and specimens that have begun
to ripen should be discarded.
Calvatia species are partial to arid climates, which is ironic considering the gigantic
size of some. It is a large and complex genus, especially in the prairies of the Midwest and
the sagebrush deserts and mountains of the West, where many endemic species still await
classification. (In our area there are several odd species which I haven’t been able to
identify to my satisfaction.) Only a handful of the more common or easily recognized
Calvatias are described here, and three small, superficially similar genera, Mycenastrum,
Calbovista, and Abstoma, are also treated.
CALVATIA& ALLIES 681

Key to Calvatia & Allies

1. Sterile base absent or rudimentary (make a perpendicular section of the fruiting body if unsure)
. 2
1. Sterile base present (but sometimes small), often but not always chambered . 14
2. Fruiting body large (10-40 cm broad or more unless very young); peridium (skin) thick in im¬
mature specimens but thinner in age and disintegrating soon after the spore mass matures
or turns powdery . 3
2. N ot as above; fruiting body small to medium-sized (typically less than 12 cm broad), or if larger
then peridium persistent (remaining intact), tough or hard, and thick even at maturity ... 4
3. Fruiting body with large warts or plaques when young, usually depressed-globose (broader
than it is tall); found in sagebrush flats and mountains of the West.C. booniana, p. 684
3. Fruiting body smooth or areolate (cracked) when young (or western form often breaking up into
plaques in dry weather), round to somewhat flattened; found mostly in grassy or open areas,
roadsides, etc.; widely distributed .C. gigantea group, p. 682
4. Peridium (skin) thick and quite tough or hard, even after spore mass has matured (i.e., skin not
readily disintegrating); spore mass not purple at maturity . 5
4. Peridium thin, fragile, and/ or disintegrating soon after spores mature (may be thick when
young); spore mass variously colored at maturity, including purple . 11
5. Fruiting body typically less than 10 cm broad; found under conifers, especially in mountains;
spores not reticulate .6
5. Fruiting body consistently larger than above, or if not then usually found elsewhere (in open
areas, livestock corrals, etc.) and/ or spores reticulate. 7
6. Peridium (skin) smooth when young or sometimes finely cracked (areolate); fruiting body often
with a small cord at base. C.fumosa, p.688
6. Not as above; peridium with whitish to grayish or smoky-brown warts that eventually break up
into plaques and flake off .C. subcretacea, p. 688
7. Outer layer of peridium (skin) a thick, white felty coat which turns grayish and fibrillose and
wears away in patches, exposing the hard thick (about 2 mm) brown persistent inner layer;
capillitium thorny; often (but not always) found in areas where livestock loiter .
.Mycenastrum corium, p. 689
7. Not with above features . 8
8. Peridium (skin) with distinct warts when young; found in arctic and tundra regions .
. C. cretacea (see C. subcretacea, p. 688)
8. Not as above . 9
9. Fruiting body averaging 2-8 cm in diameter. 10
9. Fruiting body averaging 5-20 cm or more in diameter .
. C. p achy derma & C. lepidophora (see C.fumosa, p. 688)
10. Peridium (skin) rupturing through radial tears and/ or spores reticulate .
.Abstoma townei& A. reticulatum (see Mycenastrum corium, p. 689)
10. Not as above; fruiting body usually breaking up irregularly at maturity; spores not reticulate
.C. hesperia(see C.fumosa, p. 688)
11. Mature spore mass purple or dull purple; fairly common in grass or other open areas .
.C.fragilis{sQe C. cyathiformis, p.687)
11. Not as above; mature spore mass typically some shade of brown or ochre. 12
12. Mature spore mass typically persistent (i.e., rather cottony in texture).
.C. lycoperdoides & others, p. 687
12. Mature spore mass typically powdery and easily dispersed . 13
13. Peridium (skin) often rupturing through radial tears and/ or spores reticulate .
.Abstoma townei& A. reticulatum (see Mycenastrum corium, p. 689)
13. Not as above; spores not reticulate .C. paradoxa, C. owyheensis, & others
14. Fruiting body with prominent warts, at least when young; warts large (4-20 mm), polygonal
or pyramidal and often lined (i.e., adorned by lines); found in western mountains . 15
14. Not as above (but fruiting body may have small warts), or if large warts present then habitat
different . 16
682 LYCOPERDALES & ALLIES

15. Warts usually pyramidal or pointed when young and sometimes very exaggerated, their tips
sometimes joined; usually found in forests; capillitium not thorny .C. sculpta, p. 684
15. Warts often flattened or truncated but sometimes pointed, not joined at their tips; usually found
in open places or edges of woods, roadsides, etc.; capillitium thorny .
.Calbovista subsculpta (see Calvatia sculpta, p. 684)
16. Outer surface of fruiting body staining yellow when bruised or rubbed, at least when young;
found mainly in cultivated soil. C. rubroflava (see C. lycoperdoides, p. 687)
16. Not as above . 17
17. Outer surface of fruiting body with red or reddish spots when fresh; mature spore mass pur¬
plish; found under pine and perhaps other trees; not common .
.C. rubrotincta (see C. cyathiformis, p. 687)
17. Not with above features . 18
18. Spore mass distinctly purple when mature (powdery); common in grass or other open areas
.C. cyathiformis, p. 687
18. Mature spore mass ochre to brown, etc., but not purple. 19
19. Spore mass cottony at maturity and persisting for a long time (remaining intact); peridium (skin)
smooth or granular to very wrinkled but lacking distinct warts, plaques, or spines; fairly com¬
mon in eastern and southern North America . C. craniiformis (see C. lycoperdoides, p. 687)
19. Not as above . 20
20. Sterile base very prominent and elongated (up to 12 cm long) to form a stalklike base; spore
case typically less than 8 cm broad and usually smaller in height than sterile base .
.C. elata& C. excipuliformis (see C. bovista, p. 686)
20. Sterile base sometimes prominent but not as above, and/ or spore case larger . 21
21. Typically found in pastures, lawns, roadsides, and other grassy or open places; common at low
elevations . 22
21. Typically found in arid regions (deserts, sagebrush flats, etc.) or under conifers, sometimes also
in mountain meadows. 23
22. Fruiting body rather small, typically less than 5 (rarely 7) cm broad.
.(see Lycoperdon & Allies, p. 690)
22. Fruiting body medium-sized to large (5-25 cm broad) unless very young .. C. bovista, p. 686
23. Fruiting body typically developing warts or blunt spines (at least on top) when young or as it
matures; found under western conifers, mainly in mountains C. lloydii(see C. bovista, p. 686)
23. Not as above; fruiting body lacking distinct warts and/or habitat different . 24
24. Fruiting body averaging 5-10 cm broad; base often wrinkled or furrowed and found in arid
habitats, or if not wrinkled, then found under conifers .. C. tatrensis(see C. bovista, p. 686)
24. Not with above features; usually smaller; if growing in arid habitats, then base not usually
wrinkled, and if growing in woods then base usually wrinkled .
.C. pallida & C. Candida (see C. bovista, p. 686)

Calvatia gigantea group (Giant Puffball) Color Plate 184

FRUITING BODY softball- to basketball-sized (8-60 cm or more in diameter), round
or sometimes lobed or occasionally somewhat flattened on top in age. Outer layer of
peridium (skin) thick when young (2 A mm), at first smooth (with texture of kid glove) and
pure white or brownish-stained (but in the California version often breaking up into
brownish scales even when young), then cracking up into flat scales or plates which
eventually flake off to expose the thin olive-brown inner layer—which soon disintegrates
—or both layers falling off together. STERILE BASE absent or rudimentary, but a cord¬
like “root” often present. SPORE MASS at first white and firm or cheesy, becoming
greenish-yellow and mushy with an unpleasant odor (like old urine), finally deep olive-
brown to brown and powdery. Spores 3.5-6 microns, round or nearly round, smooth or
minutely spiny, apiculate.
HABITAT: Solitary, scattered, or in groups or large circles in fields, pastures, open
Immature giant puffballs, Calvatia gigantea group. In the fall or spring, a casual jaunt through a
“puffball pasture” can yield quite a haul—the skillet in the foreground is one foot in diameter! For
scalier specimens, see photo on p. 678. (Bill Everson)

woods, cemeteries, on exposed hillsides, along roads, in drainage ditches, etc.; fairly
common in eastern North America and the Midwest. The west coast form, which is slightly
different (see comments), is sometimes abundant in our area after the first fall rains and
again in the spring. In fact, when conditions are favorable it is not unusual to find 30-40
pounds on a casual jaunt through a “puffball pasture.” Because of its preference for open
hillsides, it can often be spotted from the road. Large specimens, in fact, have been mis¬
taken by passersby for herds of grazing sheep! (Mushroom hunters, on the other hand,
are more likely to mistake grazing sheep for giant puffballs.) Dried specimens found
under houses have been mistaken for bleached skulls, while a sinister-looking individual
found in England during the war was labelled “Hitler’s Secret Weapon” and used for
propaganda purposes at an exhibition to raise war funds!
EDIBILITY: Edible and choice when the flesh is firm and white, but with laxative
effects on certain individuals. It can be sliced and fried like pancakes, or better yet, cubed
like tofu and dropped into clear soups or eaten raw in salads. The tough outer skin should
be peeled and those which have begun to ripen should be discarded. Size it not necessarily
an indication of maturity, so slice them open in the field. This will enable you to check for
maggots, which are fanatically fond of them. Infested areas can be trimmed away and the
solid portions carried home. Dried giant puffballs have been used as sponges, tinder
(before matches were invented), toys, and dyes. They were burned under beehives to
stupefy bees and used to squelch bleeding.
COMMENTS: The giant puffball is one of the best known and most familiar of all the
fleshy fungi—and it is fleshy—5 foot, 50 pound specimens have been recorded! (Alas,
the largest one I’ve found weighed “only” 7 pounds.) The exact identity of our local giant
puffball, strangely enough, is a minor mystery. It does not seem to be either the “true”
C. gigantea of Europe and eastern North America, or C. booniana of arid regions. The
former is smoother and whiter than our giant puffball, while the latter is wartier and
broader. To most people, however, its “true” identity is an academic problem best left
to the puffball pundits, who are paid to pore over such matters—after all, any large puff¬
ball is a giant puffball, and any giant puffball is a giant meal! Langermannia gigantea
and Calvatia maxima are synonyms for C. gigantea. C. bovista can also be quite large, but
has a prominent sterile base, while C. cyathiformis has purple spores at maturity. Also see
C. pachyderma and C. lepidophora (under C. fumosa).

683
Immature western giant puffball, Calvatia booniana. Note the shape (longer than it is high) and
large warts on the surface. See color plate for mature specimens. (Chuck Barrows)

Calvatia booniana (Western Giant Puffball) Color Plate 186

FRUITING BODY 15-60 cm or more broad and 7-30 cm or more high; sometimes round
or lobed, but more often somewhat flattened or depressed on top. Outer layer of peridium
(skin) thick, white to buff or tan and finally brown; at first sculptured with large warts which
soon separate to form flattened scales, plaques, or plates and eventually disintegrate along
with the thin inner layer. STERILE BASE absent or rudimentary. SPORE MASS firm
and white at first, then turning yellow or greenish and mushy and stinky, finallly becoming
powdery and olive-brown to brown. Spores4-6.5 * 3-5.5 microns, round or nearly round,
smooth or minutely spiny.
HABITAT: Solitary, widely scattered, or in groups or “flocks” infields, under sagebrush
or juniper and in other open areas; confined to the arid and semi-arid regions of western
North America, and sometimes common, especially in the late spring and summer. I
have seen it in New Mexico and Idaho, and in the mountains of southern California. A
similar but less warty species is common along the west coast (see the C. gigantea group).
EDIBILITY: Edible and choice when firm and white inside, but with laxative effects on
certain individuals. Like C. gigantea, it was apparently eaten by pioneers as well as by
some Native Americans. Be sure to check for maggots when gathering them, and re¬
member: any specimens whose flesh shows the slightest traces of color(yellow, greeen, or
brown) are no longer good to eat (unless you’re a maggot!).
COMMENTS: This giant puffball is approximately the same size as C. gigantea (50 lb.
specimens have been reported), but is not normally as round and has larger plaques or
warts—even when young. Calbovista sub sculpta (see comments under Calvatia sculpta)
is somewhat similar but much smaller (softball- or grapefruit-sized).

Calvatia sculpta (Sierran Puffball; Sculptured Puffball)

FRUITING BODY more or less egg-shaped, pear-shaped, top-shaped, or somewhat
irregular, (4) 7-18 cm high and/or broad. Outer layer of peridium composed of long,
pointed, white pyramidal warts (up to 2.5 cm long) which are erect or bent and joined at
their tips; warts often lined or grooved and arising from angular plaques which soon crack
and fall away, exposing a fragile inner layer which soon disintegrates. STERILE BASE
present, often prominent but sometimes inconspicuous, white to yellowish, often with

684
The Sierran puffball, Calvatia sculpta, is easily recognized by its enormous pyramidal or polygonal
warts. Note sterile base beneath spore case (right); specimen on left is being viewed from the top.

a purplish interior (especially in age). SPORE MASS white and firm at first, turning
yellow and then deep olive-brown as it ripens, eventually powdery. Spores 3.5-6.5 microns,
round or nearly round, minutely spiny.
HABITAT: Solitary or in small groups under conifers or sometimes in the open; known
only from the mountains of the West. It is fairly common in the Sierra Nevada in the late
spring, summer, and fall; I have not seen it on the coast.
EDIBILITY: Edible when immature, and better than most puffballs.
COMMENTS: This is easily the most spectacular of all the puffballs, and is well known
to hikers in the Sierra Nevada. The white pyramidal “peaks” make fresh specimens look
like a cross between a geodesic dome and a giant glob of meringue. Amanita magni-
verrucata can sometimes resemble it superficially (especially in the egg stage), but has
gills and a stalk. Other species; Calbovista subsculpta is a common edible western moun¬
tain puffball that is similar in size but has flatter (but sometimes pointed), less flagrant
warts. It used to be placed in Calvatia, but its capillitium have thornlike branches—a mo¬
mentous enough difference in the eyes of puffball specialists to merit a genus of its own.
It is quite common in the spring, summer, and early fall in open and grassy places or at the
edges of woods in the Sierra Nevada, Cascades, and other western mountains.

Left Calvatia sculpta in the bush(Bob Winter). Right Calvatia sculpta in the hand (Nancy Jarvis).
Calvatia bovista, immature specimens. N ote large size and prominent sterile base. S pecimen in center
is being viewed from above, hence the sterile base is not visible.

Calvatia bovista
FRUITING BODY 10-25 cm high and 5-25 cm broad, top-shaped or pear-shaped with a
broad, often flattened top in age and a large, prominent stemlike sterile base. Outer layer
of peridium (skin) white to grayish or occasionally yellowish-brown, covered with pointed
scurfy warts or soft particles, slowly breaking up into flat scales that slough off, exposing
the thin inner layer which soon disintegrates. STERILE BASE very large, constituting
up to one half the total height of the fruiting body; chambered; exterior white when young,
brown in age; smooth, persisting long after the spore case has decomposed. SPORE
MASS white and cheesy, then yellow or olive and finally olive-gold to olive-brown or
dark brown and powdery. Spores 4-6.5 microns, round, minutely warted or spiny.
HABITAT: Solitary, scattered, or in groups in pastures, exposed soil, open woods, etc.;
widely distributed. It is fairly common in our coastal pastures from fall through spring,
sometimes mingling with the giant puffball (Calvatia gigantea groups
EDIBILITY: Edible when immature, but rather soft and in my experience insipid or even
bitter. According to puffball scholar William Burk, it has been used to stop up holes in
drafty dwellings as well as nosebleeds.
COMMENTS: Also known as C. utriformis and C. caelata, this is the second largest of
our local puffballs and is easily recognized by its flattened top and large sterile base (see
photograph). It looks something like a gigantic Lycoperdonperlatum, but does not form a
pore at the top. The bowl-shaped base persists long after the rest of the spore case has
disintegrated, and old bowls are often found filled with rainwater, mosquito larvae, and
spores, or they may be windblown and completely empty. In the latter condition you may
not recognize them as puffballs, but don’t let this faze you—a special genus, Hippoperdon,
was once erected based on these empty bowls!
C. excipuliformis (-C. saccata) and C. data are two similar species which have a nar¬
rower spore case (3-10 cm broad) that is granular or coated with small spines and pale buff
to brownish, plus a narrower and greatly elongated (stemlike) sterile base. Both are wide¬
spread in wooded and open areas; the latter has practically smooth spores. C. tatrensis
is a western species with a purplish to purple-brown sterile base; it occurs under sagebrush
and in other arid waste places, or occasionally under conifers, and usually has a wrinkled
or furrowed base. C. pallida and C. Candida are two small species (up to 5 cm broad) with
a sterile base and powdery spore mass; the former has a wrinkled base and grows in woods
and mountain meadows, while the latter prefers more arid habitats. Finally, there is C.
lloydii, a fairly common species in the dry coniferous forests of the Sierra Nevada and
other western mountains. It is also rather small (less than 10 cm broad) and has a sterile
base, but features warts or blunt spines on the top, at least in age. It sometimes grows
with C.fumosa and C. subcretacea, but is easily separated from those species by its thinner
skin, and from C. sculpta and Calbovista subsculpta by its smaller warts.

686
Calvatia cyathiformis. Left: Immature specimen (turned sideways) with a well-developed sterile base.
Right: Mature specimen in which the outer layer of the peridium (spore case) is disintegrating. The
purplish color of the mature spores is the most distinctive characteristic of this species.

Calvatia cyathiformis (Purple-Spored Puffball)

FRUITING BODY 5-20 cm high and/or broad; nearly round whenyoung, becomingtop¬
shaped or pear-shaped, or round with a flattened top and narrowed base. Outer layer of
peridium (skin) smooth at first, but soon cracking into small, flat scales or patches, at
least on top; white to tan or pinkish-tan becoming purplish or purple-brown in age; inner
layer dark purple or purple-brown, smooth, thin and delicate; both layers flaking off
in old age. STERILE BASE usually present (but see comments), chambered, white to
dingy yellow or darker, persisting as a deep purplish to purple-brown cuplike structure
after the spores have dispersed. SPORE MASS firm and white when young, becoming
yellowish, then brownish and finally dull purple and powdery. Spores 3.5-7.5 microns,
round, spiny or warty to nearly smooth.
HABITAT: Widely scattered to gregarious or in rings in pastures and other grassy places,
widely distributed. It is common locally in the fall, occasional at other times.
EDIBILITY: Edible and quite good when firm and white inside. It is not quite as tasty
as the giant puffball, but is not as rich either.
COMMENTS: The striking purple color of the mature spore mass sets apart this Calvatia
from its brethren. It frequents the same habitats as C. bovista and our version of C. gigan-
tea, but there is normally only one major fruiting (at least locally)—in the fall. It is smaller
and firmer than C. bovista, and usually darker in color than the C. gigantea group. Fairy
rings of this species in the prairies of Colorado are estimated to be 420 years old—or older
than most trees! C. cyathiformisform fragilis (also called C.fragilis) is very similar, but has
only a rudimentary sterile base; it is also common in grassy areas on the west coast, and is
widely distributed. Other species: C. rubrotincta has purplish spores, but is pallid with red
spots when young; it has been found in Oregon under ponderosa pine.

Calvatia lycoperdoides (Cotton-Spored Puffball)

FRUITING BODY 1.5 -5 cm high and broad, more or less round to somewhat cushion¬
shaped. Outer layer of peridium (skin) white or pallid becoming brownish in age, at first
with soft granules, flakes, or spines (or often with small warts on top); adhering to the inner
layer and breaking up into large flakes at maturity. STERILE BASE absent or
rudimentary. SPORE MASS firm and white when young, olive-brown to brown at
maturity, with a cottony texture that causes it to remain intact for a long time. Spores
4-6.5 microns, round, with small warts.

687
688 LYCOPERDALES & ALLIES

HABIT AT: S olitary or in small groups on ground in woods and under trees; known only
from western N orth America, but C. craniiformis{see comments) is common in the eastern
states. It can be found in our area in the late fall, winter, and spring, but is not common.
EDIBILITY: Presumably edible when immature, but not big or plentiful enough to
warrant collecting. I haven’t tried it.
COMMENTS: This is one of several Calvatias with a cottony(instead of powdery) mature
spore mass. Others include: C. umbrina, a dark brown to black species; C. diguetii, a
smooth-spored westerner with an ochre gleba; and C. rubroflava, with a greenish-orange
mature spore mass and yellow-staining exterior, found mostly in cultivated soil(especially
in southern latitudes). In these species an apical pore is not formed as in Lycoperdon, and
there is no sterile base (except in C. rubroflava). Another species with a cottony spore mass,
C. craniiformis, is often wrinkled, softball-sized, and white to tan when young, with a
well-developed sterile base and yellow-green to yellow-brown spores. It is quite common in
southern and eastern North America in open areas and under hardwoods, but to my
knowledge does not occur on the west coast. It is edible, like most Calvatias, when firm
and white inside. There are several other species with a sterile base that may or may not
have a cottony spore mass when mature. These include C. elata, C. excipuliformis, and
C. lloydii(sQQ comments under C. bovista for more details).

Calvatia subcretacea (Small Warted Mountain Puffball)

FRUITING BODY golfball- to baseball-sized, 1.5-5 (7) cm in diameter, round or more
often cushion-shaped to somewhat flattened. Peridium(skin) thick and white when fresh,
the outer layer composed of white to smoky-gray or grayish-tipped warts which break up
into plates or polygons and then flake off, exposing the yellowish to brownish inner layer,
which breaks up irregularly. STERILE BASE absent or rudimentary. SPORE MASS
white and firm becoming yellowish, then olive-buff to olive-brown or brown and powdery.
Spores 3.5-6.5 microns, round or nearly round, smooth or minutely ornamented.
HABITAT: Solitary to widely scattered or in small groups in duff under mountain coni¬
fers, especially spruce and fir; fairly common in the mountains of western N orth America
from late spring through early fall. I have seen it several times in the Sierra Nevada (often
with C.fumosa), but never on the coast.
EDIBILITY: Edible when young, but difficult to find in sufficient quantity for a meal.
COMMENTS: The modest size, warty peridium, and growth under mountain conifers
help to distinguish this commonplace Calvatia (see photo at top of p. 689). It is not as
smooth-skinned as C. fumosa, and is usually much smaller than Calbovista subsculpta
(see comments under Calvatia sculpta). Mature specimens lack the apical pore charac¬
teristic of Lycoperdon, and their habitat is also distinctive; Sclerodermas are different in
color and texture. C. cretacea is a similar but larger species of arctic and tundra regions.

Calvatia fumosa
FRUITING BODY golfball- to baseball-sized, round to oval, 3-8 (10) cm broad.
Outer and inner layers of peridium (skin) adhering to each other, thick (1-5 mm) and
persistent (not disintegrating); at first smooth and white, but soon becoming grayish to
brownish and often areolate (cracking to form small scales), the undersurface and cracks
white. STERILE BASE absent or rudimentary, but a mycelialcord often present. SPORE
MASS firm and white at first, then yellowish or olive, finally dark brown and powdery (or
yellow-brown in one variety); odor often unpleasant (“a combination of sour milk, diesel
oil, and pit toilet”—Robert Ramsey). Spores4.5-8 microns, round or nearly round, spiny.
Calvatia fumosa (two specimens in center) and Calvatia subcretacea (specimens at far left and far
right) are common under mountain conifers and sometimes grow together. Both have a thick, tough
skin. C. subcretacea has a warty exterior, while C. fumosa is smoother and has a cord at the base.

HABITAT: Solitary to gregarious in duff (sometimes buried) under spruce, fir, and other
mountain conifers; common in the western United States, spring through early fall. A
similar species occurs in our area under cypress, and others occur in cultivated or hard-
packed soil (see comments).
EDIBILITY: Bitter-tasting according to Orson K. Miller; I haven’t tried it.
COMMENTS: The thick, tough, persistent skin (peridium) and modest size plus the
absence of warts distinguish this Calvatia from most others. Old specimens might be
mistaken for Sclerodermas, but possess capilliitum and are differently colored. The
peridium seldom disintegrates of its own accord, but rodents are apparently fond of
chewing on it, thus enabling the spores to escape. There are several similar Calvatias
with a thick, persistent peridium, including: C. hesperia, similar in size but white to gray¬
ish, with smooth spores and growing mostly in open places (farmland, deserts, etc.); C.
pachyderma, larger (5-15 cm in diameter) and whitish when young, with smooth to
minutely warted spores, found in open, cultivated, and arid places; and C. lepidophora of
the Midwest prairies, which is even larger(15-20 cm) and has densely warted spores. See
also Mycenastrum corium.

Mycenastrum corium
FRUITING BODY 4-20 cm broad or more, round to somewhat pear-shaped when young,
eventually rupturing in irregular fissures to form rays or plates which may bend back
somewhat in a star-shaped pattern. Outer layer of peridium (skin) a thick, white, felty
coat which becomes areolate (separates into blocklike areas), forming thin, grayish, fibril-
lose patches which eventually wear away to expose the tough, hard, persistent, smooth
inner layer, which is brown to purple-brown and about 2 mm thick. STERILE BASE
rudimentary or absent, but mycelial fibers often present. SPORE MASS firm and white
becoming olive-yellow to olive-brown and finally dark brown to purple-brown and
powdery. Spores 8-12 microns, round, warted-reticulate. Capillitium branched, thorny.
HABITAT: Scattered to gregarious on ground (sometimes partially buried) in horse
corrals, composted areas, and fields where livestock have been grazing; widely distributed,
but especially common in the West. In our area it occurs year-round. The tough spore
cases persist for months, sometimes breaking loose to blow about in the wind.
EDIBILITY: Presumably edible when firm and white inside; I haven’t tried it.

Mycenastrum corium, immature specimens. These were found in a horse corral, a favorite haunt of
this species. Note the thick skin and white felty material on exterior.
690 LYCOPERDALES & ALLIES

COMMENTS: This peculiar puffball is easy to recognize but difficult to describe. The
thick, tough inner peridium (skin) distinguishes it from Bovista and the thin-skinned
Calvatias, while the white, felty outer layer separates it from Scleroderma and the thick-
skinned Calvatias. The presence of capillitium in the spore mass indicates a much closer
kinship to the true puffballs (Calvatia, etc.) than to the earthballs (Scleroderma). Its
tendency to fruit in localities where livestock loiter is another helpful (but fallible) field-
mark. A bstoma townei and A. reticulalum are smaller and dirtier puffballs (2-6 cm broad)
with reticulate spores and unbranched capillitium. The first is said to occur in old pastures
and other waste places; the latter has been found in coastal California under cypress.

LYCOPERDON & Allies (Common Puffballs)

Small to medium-sized puffballs found mostly on rotten wood or on ground in woods (Lyco-
perdon), or in grass (Vascellum). FRUITING BODY round to pear-shaped or top-shaped; peri¬
dium two-layered, the outer layer usually with spines, warts, granules, or particles; usually rup¬
turing through an apical pore. STERILE BASE usually present and often conspicuous or stem¬
like. SPORE MASS firm and white when young, becoming powdery and olive-brown to brown or
purplish in old age. Spores more or less round, smooth to warted or spiny, sometimes pedicellate.
Capillitium present (Lycoperdon) or replaced by paracapillitium (septate, colorless hyphae) in
Vascellum.

THESE are small to medium-sized puffballs that rarely exceed 10 cm (4 inches) in dia¬
meter. In contrast to Calvatia and Scleroderma, the spores are usually released through
an apical pore (a hole, slit, or mouth that forms at the top of the mature spore case). The
frequent presence of a well-developed, often stemlike sterile base distinguishes Lyco¬
perdon from Bovista, but the two genera intergrade through a series of forms with a slight
sterile base. In most species the spore case is initially coated with a layer of spines, warts,
or fine particles, but these eventually fall away to expose the inner layer of the peridium,
in which the pore forms.
Lycoperdons are our most common woodland puffballs, but also grow in open areas,
waste places, and sawdust piles. In our area the major fruiting is in the fall and winter, but
old weathered specimens can be found at any time. All Lycoperdons are thought to be
edible when firm and white inside, but some taste better than others and it is imperative
to discard any specimens that have begun to ripen. (In Charles Mcllvaine’s words, “one
ageing L. pyriforme will embitter a hundred.”) The various species are rather difficult
to distinguish—particularly when immature—but it is always a good practice to identify
each type before eating it. Five representative species are described here, plus one species
of Vascellum, a small “satellite” genus that differs microscopically. Two other small
genera, Bovistella and Morganella, are included in the key.

Key to Lycoperdon & Allies

1. Exterior of fruiting body covered with dark brown to black spines when young; yellow tones
often developing in age .L.foetidum, 692
1. Not as above; fruiting body not dark brown when young (but may be pale to medium brown
when young and become dark brown in old age) . 2
2. Growing on wood, sawdust, or lignin-rich humus (if in humus, then base with white mycelial
threads or rhizomorphs and spore case with inconspicuous spines if any) . 3
2. Not as above; usually terrestrial . 4
3. Fruiting body with cinnamon-buff to brown spines when young, conspicuously pitted at
maturity (after spines have worn off); sterile base well-developed to practically absent; found
mainly in eastern North America Morganella subincarnata (-Lycoperdon subincarnatum)
3. Not as above; fruiting body never pitted, usually with white mycelial threads (rhizomorphs)
at base or in surrounding substrate; sterile base well-developed; common and widespread
. L. pyriforme & others, p. 691
LYCOPERDON & ALLIES 691

4. Fruiting body often broader than it is tall, white or pinkish-tinged when young; outer layer of
skin peeling away in sheets at maturity (see photo on p. 695) L. marginatum & others, p. 694
4. Not as above (if peeling in sheets, then much taller than it is broad) .5
5. Fruiting body 3-12 (14) cm broad, with a narrowed rooting base; outer layer of peridium(skin)
composed of spines and granules, the spines often tufted or joined at their tips; usually in open,
sandy, or cultivated ground . . . Bovistella radicata (see Lycoperdon puleherrimum, p. 694)
5. Not with above features; rooting base absent and/ or fruiting body considerably smaller . . 6
6. Fruiting body golden-orange to bright yellow when young . (set Bovista& Disciseda, p. 696)
6. Not as above; differently colored .7
7. Fruiting body lavender-tinged or with lavender-tinged spines when young; found mainly in
eastern North America .L. peckii (see L. perlatum, p. 693)
7. Fruiting body not lavender-tinged when young . 8
8. Typically growing in grass, prairies, and other open areas; either paracapillitium present or
capillitium with small round pits . 9
8. Not as above; typically growing in woods and at their edges, under trees, on roadsides, etc. 11
9. Sterile base very small or rudimentary; spore case rupturing through an apical pore; usually
densely gregarious or in clusters . Vascellum curtisii (see V. pratense, p. 695)
9. Not as above . 10
10. Fruiting body typically with a large mouth at maturity (the top disintegrating); very common
on lawns, golf courses, etc.Vascellum pratense & others, p. 695
10. Fruiting body typically rupturing through a slit or tear at maturity; widespread but not common
.(see Bovista & Disciseda, p. 696)
11. Spines on exterior of young fruiting body 2-6 mm long and often joined at their tips . 12
11. Spines absent, or if present not as above (usually shorter and sparser) . 13
12. Spines white becoming brown, leaving small scars or pockmarks on the inner peridium when
they fall off.L. americanum (see L. puleherrimum, p. 694)
12. Spines remaining white until they fall off, not leaving scars (i.e., inner peridium smooth) ....
.L. puleherrimum, p. 694
13. Outer layer of peridium (skin) sloughing off in sheets or chunks at maturity; known only from
the Pacific Northwest; rare (?) .L. nettyana (see L. marginatum, p. 694)
13. Not as above; very common and widespread . 14
14. Mature spore mass olive-brown to brown; spines leaving pockmarks on inner peridium when
they fall away (but marks may disappear, leaving inner peridium smooth) L.perlatum, p. 693
14. Mature spore mass purple-brown; spines not leaving scars .
.L. umbrinum (see L. perlatum, p. 693)

Lycoperdon pyriforme (Pear-S haped P uffball)

FRUITING BODY pear-shaped to nearly round, but usually with a stemlike sterile
base; 1.5-5 cm high and sometimes almost as broad in the widest part. Peridium (skin)
whitish to pale brown when young, yellowish to dark rusty-brown in age; at first smooth
or with a few small scattered spines on top, then becoming finely cracked to form small
patches or minute granules or particles (making it rough to the touch), this rough outer
layer slowly but eventually falling away to expose the smooth inner layer in which an
apical pore or tear is very slow to form. STERILE BASE small or well-developed,
spongy when fresh, occupying the stemlike base (if base is present); chambers very small;
conspicuous white mycelial threads (rhizomorphs) usually radiating from the base and
connected to others in the surrounding wood or humus. SPORE MASS at first firm and
white, then yellow to olive and finally deep olive-brown and powdery. Spores 3^4.5
microns, round, smooth.
HABITAT: Scattered to densely gregarious or clustered on stumps, rotting logs, sawdust,
and in lignin-rich humus; widely distributed and common, fruiting mostly in the fall and
winter in our area, but old bleached-out fruiting bodies can be found most any time. It
sometimes forms dense clusters “as large as a loaf of bread ” (to borrow a phrase from
Alexander Smith).
Lycoperdon pyriforme, immature specimens. Note absence of prominent spines, the narrowed
sterile base, plus the clustered growth habit and white mycelial threads.

EDIBILITY: Edible when young, but only worth collecting when it occurs in quantity.
In my fickle fungal opinion it is one of the better puffballs, but is not as good as “a loaf of
bread” and is apt to be bitter if not absolutely white and firm inside.
COMMENTS: The tendency to grow on rotting wood is a distinctive feature of this pear-
shaped puffball, but it often appears to be terrestrial (when growing from buried wood
or humus rich in lignin). The white rhizomorphs or “roots” that emanate from the base of
the fruiting body plus the narrowed or stemlike base and absence of prominent spines
are also good fieldmarks. It is one of the few Lycoperdons that occurs in sufficient quantity
to merit collecting for the table. Other species: L.pedicellatum also grows on rotten wood,
but has longer spines and ornamented spores.

Lycoperdon foetidum (Dark Puffball)

FRUITING BODY pear-shaped to nearly round, but usually with a narrowed base; 1.5-6
cm high and 1.5^1 cm broad. Outer layer of peridium (skin) composed of fine pointed
black to dark brown spines (especially dense on top) interspersed with granular material,
the spines persisting or eventually falling away; background (inner layer) thin, grayish-
tan to yellowish (usually distinctly yellow in age), developing an apical pore or tear at
maturity. STERILE BASE well-developed, chambered and spongy when fresh; exterior
usually paler than the rest of fruiting body, at least when young. SPORE MASS white
when young, then yellow, finally dull cinnamon-brown to dark brown or sepia, and pow¬
dery. Spores 4-5 microns, round, minutely spiny.

Lycoperdon foetidum, immature specimens. Note the dark brown to blackish color—its principal
fieldmark. A pore forms at the top in old age.
LYCOPERDON 693

HABITAT: Solitary, scattered, or in groups in humus and debris in deep woods along
the west coast (also Europe). Fairly common in our area from fall through early spring—
especially under conifers—but easily overlooked because of its dark color.
EDIBILITY: Presumably edible when firm and white inside; I haven’t tried it.
COMMENTS: Also known as L. nigrescens, this attractive puffball is our only Lyco-
perdon with dark brown to black spines when immature (several species may be quite
dark in age, however—see L. perlatum). The dark spines contrast nicely with the white
flesh, and the yellowish background color that develops in age is also distinctive. The
species epithet is something of a misnomer, for I have never found it to have an odor other
than the usual slightly unpleasant smell that all ripening puffballs develop.

Lycoperdonperlatum (Common Puffball; Gemmed Puffball)

FRUITING BODY pear-shaped or top-shaped (broader above) or with a flattened top,
or at times practically round with a narrowed, often wrinkled stemlike base; 1.5-6 (9) cm
broad, 3-7 (10) cm high. Outer layer of peridium(skin) consisting of slender, short, cone-
shaped spines interspersed with smaller spines or granules, the larger ones leaving scars
or pockmarks when they fall off; spines white to gray or in one form brown. Inner layer
at first with scars, but often smooth in old age, white to tan becoming yellowish-brown
to grayish-brown or dark brown in old age; eventually rupturing through a pore at the
top. STERILE BASE large, well-developed, chambered, forming a stalklike base beneath
the spore mass; at first white and spongy, then yellow, olive, brown, or chocolate-colored.
SPORE MASS firm and white but soon becoming soft and turning yellow, then olive, and
finally becoming dark olive-brown to chocolate-brown or brown and powdery. Spores
3.5-4.5 microns, round, minutely spiny.
HABITAT: Solitary, scattered, gregarious, or clustered on ground in woods and under
trees, along roads, or sometimes in the open; very widely distributed. It is probably the
most abundant woodland puffball in North America, and our region is no exception. It
can be found most any time in our area, but usually fruits in the fall or winter.
EDIBILITY: Edible when firm and white inside, but specimens showing the slightest
traces of yellow should not be eaten. It occasionally occurs in enough quantity to merit
collecting, but in my experience it is bland at best and bitter at worst.

Lycoperdon perlatum, maturing specimens. If you look closely you can see some of the spines and
the small scars they leave when they fall off. Younger specimens usually lack the apical pore that is
beginning to form in these; older ones are often smooth (without scars or pockmarks).
694 LYCOPERDALES & ALLIES

COMMENTS: Also known as L. gemmatum, this cosmopolitan puffball is easily told

by its white to gray or brownish, slender, cone-shaped spines which leave small scars behind
when they fall off. As pointed out in the description, however, the pockmarks may
eventually disappear. It is one of the most variable of all puffballs, especially in color and
size, but the well-developed sterile base, dark olive-brown mature spore mass, and
ubiquitousness help to identify it. Very large specimens are occasionally found, but these
can be distinguished from Calvatia by the apical pore which forms in old age. There are
several similar species, including: L. umbrinum (-L. molle), widely distributed and
common, with spines that do not leave scars plus a dark brown to purple-brown mature
spore mass and a purplish-tinged sterile base at maturity; L. muscorum, small and rare,
found in deep moss; L. peckii, with lavender-tinged spines when young; andL. rimulatum,
with purplish spores and a nearly smooth peridium. See also L. pyriforme.

Ly coperdon pulcherrimum (Long-S pined Puffball)

FRUITING BODY more or less pear-shaped or rounded above with a narrowed, stem¬
like, often wrinkled base; 2-5 cm broad and high. Outer layer of peridium (skin) with a
dense coating of long (3-6 mm), slender, white spines which typically form numerous
fascicles (by uniting at their tips); spines eventually wearing away to expose the smooth
brown to dark purple-brown inner layer; pore forming at the top in age. STERILE BASE
occupying up to one half the height of fruiting body, chambered, white when young, brown
to purple-brown in age. SPORE MASS white and firm at first, becoming yellow and
eventually dark purple-brown and powdery. S pores 4-4.5 microns, round, minutely spiny,
with a long pedicel (“tail”) 10-13 microns in length.
HABITAT: Solitary or in small groups in humus or on very rotten wood, usually under
hardwoods. It is not uncommon in eastern North America (especially the southern states),
and I occasionally find it (or something very similar) in our area in the fall and early winter.

EDIBILITY: Edible when young, but too small and infrequent to be of value.
COMMENTS: The long white spines that are frequently joined at their tips distinguish
this small puffball from the more common L. perlatum and other small species. L. echi-
natum (now called L. americanum by some puffball pundits) is a similar but more common
species with white spines that soon turn brown and leave small marks or reticulations on
the inner layer of the spore case when they fall off. A third puffball that often has united
—albeit shorter—spines, Bovistella radicata, strikes a much different pose: it is larger
(3-10 (14) cm), with a thick, rooting base and preference for disturbed or open ground.
Though widespread, it is not particularly common, at least in our area.

Ly coperdon marginatum (Peeling Puffball)

FRUITING BODY at first round to somewhat flattened, at maturity often broader than
it is tall, the underside usually wrinkled (at least in age) and sometimes with a short, rooting
base; 1-5 cm in diameter. Peridium (skin) white or tinged pinkish when young, the outer
layer composed of short erect warts or spines which peel off in sheets, exposing the smooth
to slightly scurfy or faintly pitted, pale to dark olive-brown inner layer; an apical pore
eventually forming. STERILE BASE chambered, usually well-developed but sometimes
inconspicuous. SPORE MASS white and firm at first, then olive to grayish-brown and
eventually powdery. Spores 3.5-5.5 microns, round, minutely ornamented or smooth,
sometimes with a broken pedicel (“tail”).
HABITAT: Scattered or in groups or clusters on ground, usually in sandy soil or waste
places; widely distributed. I find it regularly in sandy soil under oak, pine, madrone, and
manzanita, usually in the late fall and winter.
 Lycoperdon marginatum, maturing specimens. Note how the outer layer of the spore case is peeling
away in sheets. Exterior is white or pinkish-tinged when immature, browner in age.

EDIBILITY: Edible when firm and white inside. In Mexico, this species and L. mizte-
corum are used to induce auditory hallucinations. It is known as “gi-i-sa-wa,” meaning
“fungus of the second quality” {L. miztecorum being the fungus of first quality). H owever,
no intoxicating substances were found when these puffballs were analyzed.
COMMENTS: The peculiar manner in which the outer layer of the peridium separates
completely from the inner layer and peels off in sheets is the hallmark of this pleasing
puffball. L. candidum is a synonym./,, nettyana, known only from the Pacific Northwest,
also peels off in sheets, but is more or less pear-shaped (with a well-developed, stalklike
sterile base). Vascellum (-Lycoperdon) curtisii is also quite similar, but usually grows
in grass and has little or no sterile base. L. rimulatum (see comments under L. perlatum)
sometimes peels off in sheets, but has a smooth (spineless) peridium.

Vascellum pratense (Field P uffball)

FRUITING BODY typically top-shaped (broader above) or round with a narrowed base,
often wrinkled somewhat below; 2-6 cm high and 2A cm broad. Outer layer of peridium
(skin) scurfy from a coating of particles or fine spines (especially on top) which disappear
in age, exposing the smooth inner layer; white when young, yellowish-tan to metallic
brown in age (or the inner layer grayish); rupturing at the top to form a large pore that soon
widens so that only the base and lower portion of the spore case remain, forming a“bowl.”
STERILE BASE usually well-developed, chambered, white becoming brownish or
purplish in age, separated from the spore mass by a thin membrane (but see comments).
SPORE MASS white but rather soft when young, then greenish-yellow and finally olive-
brown and powdery. Spores 3-5 microns, round, minutely warted, more or less apiculate.
HABITAT: Widely scattered to gregarious in grassy places—lawns, golf courses, pastures,
etc; widely distributed, but especially common on the west coast(along with V. lloydianum
— see comments). In our mild climate it occurs year-round, but is most abundant in the
fall. I have also seen large fruitings in Oregon.
EDIBILITY: Edible when immature, but of mediocre quality.

Vascellum pratense is a small grassland puffball with a conspicuous sterile base. Specimens at left
are immature; bowl-shaped specimen at right is quite old.
Hi
696 LYCOPERDALES & ALLIES

COMMENTS: This undistinguished grassland puffball can be distinguished by its modest

size, the presence of a sterile base and soft, deciduous, scurfy spines or granules, and the
bowl- or urnlike remains of ruptured specimens. It is often found in the company of Bo-
vista plumbea, which is rounder and smoother and lacks a sterile base. It used to be called
Lycoperdon hiemale, but was transferred to the genus Vascellum because of microscopic
differences. H owever, now that taxonomists have settled upon its genus name, they cannot
seem to agree on a species epithet. S ome hold out for V. pratense, others for V. depressum,
while Alexander Smith has described the common western variant as a “new” species, V.
lloydianum, because the membrane separating the spore mass from the sterile base is not
very distinct. It hardly matters what you call it, however, since it belongs to that vast army
of fungi that are “better neglected than collected.” Other species: V .(-Lycoperdon) curtisii
is a small (0.5-2 cm) widespread species with numerous spines when young and a distinct
apical pore at maturity. It somewhat resembles Lycoperdon marginatum when fresh, but
usually grows gregariously in grass.

BOVISTA & DISCISEDA (Tumbling Puffballs)

Small or occasionally medium-sized puffballs found mostly in grassy or open areas. FRUITING
BODY more or less round to somewhat flattened or irregular; peridium two-layered, rupturing
to forma large mouth at the top (Bovista), or often rupturing basaWy (Disciseda); inner layer usually
thin and papery when mature; outer layer usually lacking distinct spines or warts, persistent in
Disciseda, disappearing in Bovista. STERILE BASE typically absent or rudimentary. SPORE
MASS firm and white becoming powdery and brown to yellowish. Spores round to oval or elliptical,
minutely warted or spiny or smooth, often pedicellate. Capillitium present, typically branched.

THESE small (marble- to baseball-sized) puffballs usually lack the sterile base charac¬
teristic of Lycoperdon and are smaller and/ or smoother than most Calvatia species. In
old age they often break loose from the soil so that the thin, lustrous, papery spore cases
tumble about freely in the wind.
Bovista species release their spores through an apical pore or large mouth, and are
probably the most common puffballs of our lawns and pastures. Disciseda, on the other
hand, is as rare as it is bizarre. In the most common species, D. Candida, the tough outer
peridium splits around its “equator” and the papery inner layer ruptures basally, then
the spore case breaks loose and flips over so as to resemble an acorn in a cup.
Bovistas are edible when firm and white inside, but are bland at best and bitter at worst.
Discisedas are too rare to be of food value and are not normally found until they are old
and powdery. One representative of each genus is described here.

Key to Bovista & Disciseda

1. Outer layer of peridium (skin) persistent, often incrusted with sand or dirt particles and often
forming an acornlike cup around the inner layer; spores typically released through a small
pore or tear at maturity; not common.D. Candida & others, p. 698
1. Not as above; outer layer of peridium not persistent, or if persistent then typically rupturing
radially or forming a large mouth at top of spore case . 2
2. Fruiting body bright yellow to golden-orange when young B. colorata(see B. plumbea, p. 697)
2. Not as above . 3
3. Fruiting body typically buried in ground except for the apex or mouth, even at maturity .. 4
3. Not as above (but fruiting body may be buried or half-buried when very young) . 5
4. Outer peridium often persisting for a long time; found in grass or prairies in north-central U nited
States and Canada . Gastrosporium simplex
4. Not as above; found mainly in eastern North America under trees, along roads, in bare soil
and waste places, etc.B. minor (see B. plumbea, p. 697)
BOVISTA & DISCISEDA 697

5. Sterile base present .B. dakotense (see B. plumbea, below)

5. Sterile base absent or rudimentary . 6
6. Fruiting body small, with crowded spines and scurf when young, rupturing through an apical
pore at maturity; usually clustered or densely gregarious . (see Lycoperdon & Allies, p. 690)
6. Not as above; fruiting body smooth or with small warts or granules when young . 7
7. Fruiting body small (less than 2 cm broad) and round, but usually with a narrowed or pinched,
rooting base; rupturing through an apical pore at maturity .
..B. pusilla & B. longispora (see B. plumbea, below)
7. Not as above; usually rupturing through a large mouth or tear at maturity . 8
8. Fruiting body typically 1-3 (6) cm broad and attached to ground by a small patch of dirt-bound
fibers when young .B. plumbea & others, below
8. Fruiting body typically 3-9 cm broad, usually attached to ground by a small cord or “root”
when young ....9
9. Spore case typically dark brown to bronze-brown or coppery when mature; common in North
America .B. pila (see B. plumbea, below)
9. Spore case typically gray to lead-colored when mature; common in Eurasia, uncommon or
absent (?) in North America .. . B. nigrescens (see B. plumbea, below)

Bovistaplumbea (Tumbling Puffball; Tumble Ball)

FRUITING BODY round or slightly flattened, \A (8) cm in diameter, usually with a
small patch of dirt (held together by fibers) at base. Outer layer of peridium(skin) white,
smooth, felty, or with small flattened scales, peeling away or shrivelling up in age to reveal
the smooth, papery inner layer which is blue-gray to purplish-brown or lead-colored in
age and usually has a metallic luster; inner layer rupturing at the top to form a large
circular “mouth.” STERILE BASE absent. SPORE MASS white and firm at first, then
yellow-olive and mushy, finally powdery and olive-brown to deep chocolate-brown.
Spores 5-7 * 4-5 microns, oval or broadly elliptical to nearly round, minutely spiny to
nearly smooth, with a long pointed pedicel (“tail”) 8-14 microns in length.
HABITAT: Solitary, scattered, or gregarious in grassy places; widely distributed and
very common. In our area it fruits mainly in the fall and winter in pastures, year-round
on lawns, golf courses, and cemeteries. The thin, papery windblown spore cases can be
found most any time.
EDIBILITY: Edible when immature, but mediocre. It is bland, and a great many would
be needed for a meal.
COMMENTS: This paltry puffball is a common fungal feature of our lawns and pastures.
In the immature stage it can be distinguished from Vascellum pratense and other grass-
loving puffballs by its small size, smooth white skin, and absence of a sterile base, while
older specimens are easily told by their lustrous, paper-thin spore cases. Another wide¬
spread cosmopolitan species,B.pila, is slightly larger(3-9 cm in diameter) and attached to
the ground by a small cord or “root.” It has a dark brown to bronze mature spore case,
smooth apedicellate (tail-less) spores, and grows in cultivated and manured soil as well as
in grass and open woods. I have found it several times in our area, but it is not nearly as

Bovista plumbea, immature specimens. This common small round puffball of lawns and pastures
lacks a sterile base. Note the patch of dirt-bound fibers below. B. pila (not illustrated) is similar but
slightly larger and has a rootlike cord at the base rather than a patch of fibers.
Bovista plumbea, mature specimens. Note metallic lustre, thin skin, and the large mouth that is
starting to form in each one.

numerous as B. plumbea. Another larger species with a cordlike root, B. nigrescens, is

lead-gray when mature and is of questionable occurrence in North America. A cosmopoli¬
tan species, B. pusilla (-Lycoperdon pusillum), is one of our smallest puffballs. It is
marble-sized and more or less round above and pinched below to a cordlike root. It also
lacks a sterile base and the spores are not pedicellate. It usually grows in grass, bare ground,
vacant lots, sandy soil, etc.; it is also known asB. californica. Other small species include:
B. longispora, similar to B. pusilla, but with a slight sterile base and elliptical spores;
B. leucoderma, reddish- to bronze-brown when mature with a patch of fibers at the base
when young; B. minor, rare, with a trash- or dirt-covered spore case, found in poor soils
and usually developing underground; B. colorata (-Lycoperdon coloratum), easily told
by its bright yellow to golden-orange color; and B. dakotense, one of the few members
of the genus to have a sterile base (leading to confusion with Lycoperdon, but usually
growing in grassy or open areas). None of these are worth eating.

Disciseda Candida (Acorn Puffball)

FRUITING BODY round to somewhat flattened or irregular, 0.8-3 cm broad. Outer
layer of peridium (skin) dirt-incrusted, composed of white mycelial matter that is cottony
when young but tough in age; inner layer minutely granulose or scurfy, pallid to tan or
grayish, forming a basal pore at maturity. Outer layer rupturing around its periphery,
the upper portion retaining the inner layer (spore case) within it, and detaching and flip¬
ping over so as to resemble an acorn in its cup(thereby causing the pore to appear apical).
STERILE BASE absent. SPORE MASS white when very young, soon yellowish to olive,
finally olive-brown to brown and powdery. Spores 3.5-4.5 microns, round, minutely
warted, with a short stubby pedicel (“tail”).
HABITAT: Solitary or in groups in pastures and other grassy or open areas, along paths,
in barnyards, etc; usually developing at or beneath ground level and then flipping over or
breaking free to tumble about in the wind. It is widely distributed, but not common. It
has been found in southern California on sandy and grassy soil. In eastern North America
it is sometimes found with Arachnion alburn^see the key toLycoperdales). D. subterranea
(see comments) has been reported from Washington, Colorado, and Wyoming.
EDIBILITY: W orthless—it is rarely found when white inside, and is insubstantial anyway.
COMMENTS: This peculiar puffball is easily told by its small size and acornlike mature
fruiting body. Other species: D. subterranea is similar, but has a grayish to bluish-gray
spore case and larger spores, while D. pedicellata has spores with long “tails.” Several
other species occur in sandy or exposed soil in California and the Southwest, but are rarely
collected and difficult to identify because they are not necessarily acornlike. They have very
thin spore cases (as in Bovista) that usually retain vestiges of the outer peridium. Some
of them are: D. ater, which has a blackish outer peridium; D, luteola, originally described
from Lake Tahoe, California, with a yellowish to dull yellow-brown spore case; andD.
brandegeei, a desert species with smooth spores.

698
 699

GEASTRUM, ASTRAEUS, & MYRIOSTOMA

(Earthstars)
Small to medium-sized, mostly terrestrial fungi. FRUITING BODY round or flattened when
young (usually underground in this stage), but at maturity the outer wall(peridium)splitting into
several starlike rays which curl back (and often under) the spore case. SPORE CASE (inner peri¬
dium) smooth or roughened, rupturing through an apical pore (Geastrum & Astraeus) or through
several pores (Myriostoma), or irregularly (Astraeus). SPORE MASS white when very young
(underground), brown and powdery when mature. STALK and STERILE BASEabsent, butspore
case sometimes mounted on a pedicel (short stalk) or pedicels. Spores round and warted or
occasionally smooth. Capillitium present, branched in Astraeus, unbranched in Geastrum and
Myriostoma.

EARTHSTARS are modified puffballs in which the thick outer skin splits into starlike
rays which unfold and often recurve, exposing the spore case (inner skin) to the elements.
Some puffballs and earthballs (e.g., Scleroderma geaster, Mycenastrum corium) rupture
in starlike fashion, but only the earthstars have a discrete spore case intact within the rays.
Some earthstars are hygroscopic; the rays open in wet weather to expose the spore case
to raindrops, and then close up in dry weather to protect it. This phenomenon can be
observed at home by placing a closed-up earthstar in a shallow bowl of water. If hygro¬
scopic, it will open up in a few minutes—even if it is a year or two old! Earthstars are also
accomplished acrobats, coming in an amazing assortment of unorthdox and extra¬
ordinary poses. Geastrum fornicatum in particular is so prodigious a contortionist
that it is difficult to find two that are alike.
The earthstars are comprised of three genera: Geastrum (-Geaster), Astraeus, and
Myriostoma. Geastrum embraces a sizable number of small to medium-sized, difficult-
to-identify earthstars that rupture through a distinct apical pore and have small spores
(typically less than 7 microns long). Some Geastrums are hygroscopic, but most are not.
Astraeus, on the other hand, is always hygroscopic. It includes two species (one quite
large) that rupture irregularly or through a large apical pore or slit, and have relatively
large spores (typically more than 7 microns in diameter). The third genus, Myriostoma,
includes a single rare species whose spore case is perforated by many holes (like a salt-
shaker) instead of just one. Most puffball pundits place the latter two genera alongside the
earthballs in the Sclerodermatales because of certain microscopic features. However, they
are grouped here with Geastrum because of their superficial similarity.

Fruiting body development in Geastrum fornicatum. Earthstars resemble puffballs when very young
(specimen at bottom left), but their outer skin splits into rays and unfolds (bottom right), revealing
the inner skin or spore case (specimens at top). See p. 701 for another photo of this species.
700 LYCOPERDALES & ALLIES

Earthstars occur in a variety of habitats (pastures, woods, waste places, etc.), but like
many of the Gasteromycetes, they are especially prevalent and diverse in arid and semi-
arid regions. The Southwest, for instance, is unusually rich in Geastrum species. They
are real crowdpleasers, but have no culinary value because of their tough or woody
texture.* Several earthstars are described and/ or keyed out here, but in the case of Geas¬
trum, they represent only a fraction of the total number.

Key to Geastrum, Astraeus, & Myriostoma

1. Spore case typically rupturing irregularly or through a single apical pore (mouth at top) . . 2
1. Spore case typically with more than one (usually several) pores at maturity . 18
2. Rays distinctly hygroscopic (i.e., folding over the spore case in dry weather to protect it and
unfolding when moistened); spore case usually sessile (not seated on a short stalk) .3
2. Rays not hygroscopic; spore case may or may not be seated on a short stalk . 7
3. Fruiting body 5-15 cm or more broad when expanded; spore case 2.5-5 cm broad, rupturing
irregularly (i.e., not forming a distinct pore) .Astraeuspteridis, p. 706
3. Fruiting body smaller and/or rupturing through an apical pore or slit.4
4. Spore case roughened by numerous particles; apical pore often irregular or poorly defined or
merely a slit; underside (exterior) of rays often with blackish fibrils; spores over 7 microns
in diameter .Astraeus hygrometricus, p. 705
4. Not with above features; apical pore usually well-defined and spores less than 7 microns . . 5
5. Apical pore (mouth) with distinct radial grooves .
.Geastrum drummondii & others (see Astraeus hygrometricus, p. 705)
5. Apical pore not grooved (but may be fibrillose) . 6
6. Apical pore fibrillose . Geastrum mammosum & others (see Astraeus hygrometricus, p.705)
6. Apical pore more or less naked . . Geastrum floriforme (see Astraeus hygrometricus, p. 705)
7. Spore case mounted on a distinct pedicel (short stalk) . 8
7. Spore case sessile (stalkless) or nearly so . 15
8. Mouth of spore case “beaked” (i.e., forming a raised cone), the beak longitudinally grooved 9
8. Not as above; mouth not beaked, or if beaked, then not grooved . 11
9. Pedicel with a distinct collar around base of spore case .
.Geastrum striatum (see G. pectinatum, p. 702)
9. Not as above . 10
10. “Beak” long; pedicel also long (2-6 mm) and slender.Geastrum pectinatum, p. 702
10. Not as above . Geastrum nanum & others (see G. pectinatum, p. 702)
11. Fruiting body small, typically less than 3 (4) cm broad when expanded; spore case typically
3-10 mm broad . 12
11. Fruiting body larger, typically 2-6 cm or more broad; spore case typically 1-2.5 cm broad 13
12. Fruiting body usually with 8-12 rays .Geastrum minimum (see G. fornicatum, p. 701)
12. Fruiting body usually with 4-6 rays .Geastrum quadrifidum (see G. fornicatum, p. 701)
13. Fruiting body with 4-5 (6) rays (but the tips may split) that are usually attached to a “cup” of
tissue, mycelium, and debris; fruiting body poised on the tips of its rays when mature; apical
pore or mouth more or less same color as rest of spore case . . . Geastrum fornicatum, p. 701
13. Fruiting body usually with 6-12 rays, or if with fewer rays then not as above . 14
14. Mouth area usually delimited (set off from rest of spore case), sometimes paler in color than rest
of spore case.Geastrum limbatum (see G. pectinatum, p. 702)
14. Mouth area not clearly delimited . Geastrum rufescens (see G. pectinatum, p. 702)

*Captain Charles Mcllvaine, as usual, casts a dissenting opinion, sayingof Astraeus hygrometricus: “when young**
it is, when cooked, soft and creamy inside. The outer part is tough and semi-glutinous but of pleasant texture. It
has not a marked flavor, but makes a succulent dish.” (Does this make any sense to you?)

**Immature earthstars are not commonly found because they’re inconspicuous and often develop underground.
GEASTRUM, ASTRAEUS, & MYRIOSTOMA 701

15. Fleshy upper (inner) layer of rays often breaking away to form a broad cup or saucer around
the base of spore case .Geastrum triplex, p. 703
15. Not as above; spore case not seated in a saucer . 16
16. Mouth area sharply delimited (i.e., with a circular zone around it) . 17
16. Mouth not sharply delimited .Geastrum fimbriatum (see G. saccatum, p. 703)
17. Mouth with distinct radial grooves Geastrum campestre (see Astraeus hygrometricus, p. 705)
17. Mouth not grooved, but often silky-fibrillose . Geastrum saccatum, p. 703
18. Spore case mounted on several pedicels (short stalks) .Myriostoma coliforme, p. 704
18. Spore case sessile (stalkless) .Geastrumpluriosteum (see Myriostoma coliforme, p. 704)

Geastrumfornicatum (Acrobatic Earthstar)

FRUITING BODY round or flattened when young and incrusted with debris, the outer
wall (peridium) then splitting into 3-6 (usually 4 or 5) rays which peel back and under until
they’re more or less erect. Mature fruiting body 3-6 cm broad and 5-10 cm high, usually
poised on the pointed tips of its rays, the tips joined to a concave mycelial mat or “cup”
which has a grayish interior and debris-incrusted exterior and may be broken into rays
itself. RAYS not hygroscopic; underside usually smooth and tan to brown; upper surface
chocolate-brown to dark brown or nearly blackish, but peeling off in patches to reveal the
smooth, paler brown or tan undersurface. SPORE CASE 1-2.5 cm broad, round or
somewhat flattened to urn-shaped, mounted on a pedicel (short stalk), often with a
compressed collar underneath it (at the top of the pedicel); surface brown to dark choco¬
late brown, finely velvety; rupturing at the top to form a fairly large, irregularly torn or
lacerated pore. SPORE MASS chocolate-brown to blackish-brown and powdery at
maturity. Spores 3-5 microns, round, warted.
HABITAT: Scattered to densely gregarious in humus under trees, on rubbish, around
stables and other waste places, etc.; widely distributed, but apparently rare except in
southern California and the Southwest, where it is quite common. In our area I have
found it in only one locality—under oaks where there is run-off from a bathing area for
horses—but it fruits there by the hundreds and can be found every month of the year. In
New Mexico it is quite common under juniper.

Geastrum fornicatum, mature specimens perched on the tips of their rays. Note how the rays peel
in patches and how the specimens at left are firmly attached to a mat or cup of mycelium and debris.
See p. 699 for another photograph of this acrobatic species.
702 LYCOPERDALES & ALLIES

EDIBILITY: Inedible, but makes an extraordinary ornament or conversation piece (it

is easily dried).
COMMENTS: One of the most distinctive of all fleshy fungi, this earthstar is a true
pleasure to find. Mature individuals defy description, but look something like a cross
between a flat tire and a ballet dancer. The tendency of the rays to stand erect on their tips
(like a ballet dancer) plus the membranous mycelial cup (“flat tire”) to which they’re
attached are the principal fieldmarks. Because of the likeness to a human figure, it
was named Fungus anthropomorphus when first described in 1688. Specimens are some¬
times found with only one or two “arms” extended, as if they were putting out tentative
“feelers” before committing themselves completely (and irrevocably) to exposure. Other
species: G. quadrifidum (also called G. coronatum) is a similar but somewhat smaller
species with a well-defined, conical mouth that is paler than the rest of the spore case; it is
also equipped with a mycelial “cup” and is widely distributed. G. minimum, which may or
may not be a diminutive form of G. coronatum, has8-12 rays and usually lacks a mycelial
cup; it occurs commonly intheSouthwestunderpinyonandjuniper. For other species with
a pedicel beneath the spore case, see G. pectinatum.

Geastrumpectinatum (Beaked Earthstar)

FRUITING BODY round to flattened when young, the outer wall (peridium) then split¬
ting into 5-12 rays(usually6-10) which peel back and under or stand upright; 3-6 cm broad
when expanded. RAYS not hygroscopic, copiously incrusted with mycelial matter and
debris on underside (exterior); upper surface brown to grayish-brown, fleshy, cracking
irregularly or flaking off as it dries, revealing the smooth undersurface. SPORE CASE
0.5-2 cm broad, round to somewhat urn-shaped, mounted on a prominent pedicel; sur¬
face more or less smooth, lead-colored to purple-brown, but often appearing paler from a
hoary sheen; underside sometimes striate or grooved near the pedicel; rupturing to form
a prominent apical pore that is beaklike (more or less conical) and strongly grooved or
furrowed longitudinally. Pedicel slender, 2-6 mm long, sometimes with a collar or ring
around its middle or base. SPORE MASS brown and powdery when mature. Spores
4-6 (7) microns, round, warted.
HABITAT: Solitary or in groups in humus under trees, especially conifers such as juniper,
or less commonly in the open; widely distributed but not common, fruiting after rains but
persisting for weeks ormonths. I have found it(or something very similar) inNew Mexico.
EDIBILITY: Too tough to be edible.
COMMENTS: The prominent pedicel and “beaked” mouth of the spore case distinguish
this earthstar from most others. The rays are not hygroscopic, and their tips are not joined
to a mycelial “cup” as in G. fornicatum. Other earthstars with a pedicel and grooved
mouth include: G. striatum (-G. bryantii), similar but typically with a prominent collar
at the base of the spore case (where it meets the pedicel); G. nanum(-G. schmidelii), also
similar but much smaller (up to 2.5 cm broad) and more common, with a shorter “beak”
and a short, thick pedicel; G. smithii, common in the Southwest, withaflattened-conical
mouth seated in a well-defined, depressed area; and G. xerophilum, also common in the
Southwest (but usually growing in the desert), with a short pedicel(if any) and a granular
spore case. T wo species with a non-grooved mouth should also be mentioned: G. limbatum
is a medium-sized to fairly large species with a well-defined fibrillose (but not grooved
or beaked) mouth and 6-12 rays (more than G. fornicatum—see photo on p. 703). Itcomes
in a striking dark brown to black version that is common in the Southwest, and a paler
(pinkish-buff to brown) form that is more widely distributed. G. rufescens, on the other
hand, has a poorly-defined, often fringed or torn mouth and a short or indistinct
Left: Geastrum limbatum (see comments under G. pectinatum). Note the numerous rays and the
pedicel (short stalk) below spore case. Right: Geastrum triplex, mature specimen. The “saucer”
around the spore case is distinctive.

pedicel. It is fairly large (5-10 cm broad) and is more common in eastern North America
than in the West. All of these are better eyed then fried.

Geastrum triplex (Saucered Earthstar)

FRUITING BODY round to flattened or bulblike when young, the outer wall splitting
at maturity into 4-8 rays which unfold and then bend under the spore case; 3-10 cm broad
when expanded. RAYS not hygroscopic, usually free of adhering debris on underside;
upper surface with a rather thick, fleshy, pinkish to tan or brown layer that cracks into
patches, the central region usually breaking loose to form a broad cup or saucer around
the spore case. SPORE CASE 1-3 cm broad, round or flattened, sessile (not mounted on
a stalk); pale to dark tan, grayish, or even reddish-brown, rupturing through an apical
pore which is often paler, slightly raised, and radially fibrillose. SPORE MASS deep
brown to smoky-brown and powdery when mature. Spores 3.5-4.5 microns, round,
warted.
HABITAT: Solitary or in groups in forest humus and under trees (usually hardwoods);
widely distributed and common in some regions. I have found it only twice in our area
(in the fall and winter), but have seen large fruitings under aspen in the Southwest.
EDIBILITY: Supposedly edible when immature (white inside), but like all earthstars,
rarely found in that stage, and too tough and fibrous to eat when older.
COMMENTS: Also known as G. indicum, this relatively large, attractive earthstar is
distinguished by the shallow cup or “saucer” that often forms around the base of the spore
case. There seems to be a great deal of variation in the extent to which the fleshy upper
surface of the rays cracks, and those which don’t crack and lack a saucer can be confused
with G. saccatum and G. fimbriatum (which, however, are often smaller). For other
relatively large, non-hygroscopic earthstars, see comments under G. pectinatum.

Geastrum saccatum (Sessile Earthstar)

FRUITING BODY round, flattened, or bulblike when young, the outer wall splitting
at maturity into 4-8 rays which unfold and then bend back under the spore case; (1.5) 2-5
cm broad when expanded. RAYS not hygroscopic, rather rubbery when fresh, relatively
clean and buff to pale tan on underside; upper surface fleshy, pallid becoming pale tan,
pinkish, or ochre-brown; sometimes cracking. SPORE CASE 0.5-2 cm broad, round to
somewhat flattened, smooth, sessile (without a stalk or pedicel), papery, pallid to buff,
dull grayish, or brownish; rupturing through an apical pore that is often paler in color
and set off from the rest of the spore case by a circular line, ridge, or groove; pore usually
raised and fibrillose. SPORE MASS brown and powdery when mature. Spores 3.5-5
microns, round, warted.
703
Left: Geastrum saccatum lacks a pedicel and “saucer,” but features a circular zone around the pore
at top. The rays often bend under the spore case as shown in photo at right. Right: Geastrumfim-
briatum resembles G. saccatum, but lacks circular zone (see comments under latter species).

HABITAT: Solitary, scattered, or in groups in humus under trees (especially conifers),

in woods, or occasionally in the open; widely distributed. This species and G.fimbriatum
(see comments) seem to be the most common Geastrums in our area. They fruit mainly
in the fall and winter, but persist for weeks or months without decaying.
EDIBILITY: Inedible in the mature state in which it is usually found.
COMMENTS: The spore case in this species has neither the pedicel (short stalk) of G.
fornicatum and G. pectinatum, nor the saucer of G. triplex, and the rays are not hygro¬
scopic. There are several similar species, including: G. fimbriatum (-G. sessile), often
slightly darker, with a torn or fringed apical pore that is not set off by a circle, usually
growing under trees and widely distributed; and G. rufescens and G. xerophilum, both
of which may have a slight pedicel (see comments under G. pectinatum for more details
on these species, and also note the hygroscopic Geastrums discussed under Astraeus
hygrometricus, most of which lack a pedicel).

Myriostoma coliforme (Saltshaker Earthstar)

FRUITING BODY round to flattened when young, the outer wall (peridium) splitting
into 5-14 pointed rays which open out; 2-12 cm broad when expanded. RAYS not hygro¬
scopic; more or less smooth on underside (exterior), but often covered with dirt near base;
inner (upper) surface with a brown to cinnamon fleshy layer that weathers or peels away,
revealing the paler (buff) undersurface. SPORE CASE 1 -5 cm broad, round to somewhat
flattened, usually mounted on several slender, short, sometimes inconspicuous pedicels
(columns); surface minutely roughened, brown to silvery-brown or grayish in age, rup¬
turing through several to many mouths or pores, mostly on the upper portion(the number
of mouths often corresponding to the number of pedicels). SPORE MASS powdery and
brown at maturity. Spores4-6 microns, round, warted.
HABITAT: Solitary or in groups in sandy soil, sometimes under trees, fruiting after
heavy rains but persisting year-round; widely distributed. It is usually listed as rare, but
is not uncommon in New Mexico, where I have seen it in the summer and fall.
EDIBILITY: Not edible.
COMMENTS: This fungal rarity can be recognized with clarity by its saltshaker-like
spore case mounted on several short columns. Most taxonomists place it alongside
Astraeus in the Sclerodermatales because of certain microscopic features. Geastrum
pluriosteum is a rare southwestern earthstar that forms one to several pores on a stalkless
spore case.
704
Astraeus hygrometricus, mature specimens. Ten minutes before this picture was taken, the fully
expanded individuals were curled up into tight “fists” like the one in foreground. However, a quick
“bath” caused the “fists” to unclench. Note the roughened surface of the spore case.

Astraeus hygrometricus (Hygroscopic Earthstar)

FRUITING BODY round to flattened when young, the outer wall splitting at maturity
into 6-15 pointed rays; 1-5 (8) cm broad when fully expanded. RAYS hygroscopic (un¬
folding in wet weather and closing up over the spore case in dry weather), often unequal in
length, tough and leathery when moist, hard when dry; underside (exterior) fibrillose,
usually with adhering sand and sometimes with black hairlike threads (rhizomorphs) at
base; upper surface (interior) at first smooth, but often developing numerous cracks,
pale tan darkening to brown, gray, or blackish-brown. SPORE CASE 0.5-3 cm broad,
round or somewhat flattened, sessile (without a stalk), thin and papery; surface whitish
to grayish or brownish and roughened by particles (and/ or finely reticulate); rupturing
at maturity through an irregular or poorly defined apical pore or slit. SPORE MASS
brown to cocoa-brown and powdery when mature. Spores 7-11 microns, round, finely
warted.
HABITAT: Solitary, scattered, or gregarious in old fields, sand or sandy soil, pastures,
roadsides, waste places, etc.; worldwide in distribution. It grows from sea level to above
timberline, from verdant pastures to desert wastelands. In our area it fruits mainly in the
fall, but the tough fruiting bodies persist for months without decaying. I have seen hundreds
of spiderlike fruiting bodies in Arizona under pinyon pine, but when the weathered,
blackened outer skins (rays) are found without their spore cases, they look like old tire
patches.
EDIBILITY: Indisputably inedible because of its toughness.
COMMENTS: This veritable barometer is the most theatrical of all the earthstars. A few
minutes’ immersion in water will open up old, dried-up specimens that seem as tightly
closed as clenched fists. It is distinguished from A. pteridis by its smaller size (but see
comments under that species). There are also several hygroscopic species of Geastrum
that have spores less than7.5 microns in diameter. They grow mainly in arid climates(e.g.,
the deserts of the West), but a few are more cosmopolitan. Several of these Geastrums can
be distinguished from A. hygrometricus by the presence of a prominently grooved apical
pore, e.g., G. drummondii, with a granular spore case; G. umbUicatum, with a smooth-
spore case; and G. campestre, with a roughened spore case and sometimes a short pedicel
(stalk). Others have a fibrillose or fringed pore, e.g., G. mammosum, especially common
in the Southwest under pinyon and juniper; G. recolligens, very similar, but found in sand
or waste places; and G. arenarium, also in sand or deserts, but only slightly hygroscopic.
Still others have a naked mouth (merely a puncture or slit) like that of A. hygrometricus.
One of these, G.floriforme(-G. delicatus) is widely distributed and occurs in our area, but
can be separated from A. hygrometricus by its small(up to 1.5 cm broad), smoother spore
case and smaller spores.
705
Astraeus pteridis, mature specimen. This species is much larger than A. hygrometricus and often
has a more pronounced crazy-quilt pattern on its rays. Note how the spore case disintegrates irregu¬
larly. (Joel Leivick)

Astraeus pteridis (Giant Hygroscopic Earthstar)

FRUITING BODY round or somewhat flattened when young, the outer wall then split¬
ting into 6 or more rays; 5-15 cm or more broad when fully expanded. RAYS hygroscopic
(unfolding in wet weather, closing up over the spore case in dry weather), thick (3-6 mm),
tough and leathery becoming hard or woody when dry; exterior (underside) with a thin
coating of matted brownish fibrils which may or may not wear away; upper surface (in¬
terior) often conspicuously cracked or fissured transversely to form a checked pattern;
tan when fresh (but dark brown in the cracks), darker overall in age (but weathered speci¬
mens sometimes whitened). SPORE CASE2.5-5 cm broad, round or somewhat flattened,
sessile (not seated on a stalk); at first roughened, veined, or fibrillose, but soon smooth or
merely roughened; grayish-brown to brown; rupturing irregularly in age, i.e., not forming
a distinct pore. SPORE MASS dark brown and powdery when mature. Spores 8-12
microns, round, warted.
HABITAT: Solitary, widely scattered, or in groups on ground, usually along roads, rail¬
road tracks, in waste places, old fields, etc., but sometimes in woods; known only from
western North America. In our area it is fairly common year-round.
EDIBILITY: Much too leathery or woody to be edible.
COMMENTS: The thick rays which are frequently checkered like a crooked crossword
puzzle plus the large size and irregular rupturing of the spore case distinguish this im¬
pressive species from its more mundane cousin, A. hygrometricus, and other hygroscopic
earthstars. However, the two species seem to intergrade in our area (e.g., you can find
fairly large, uncheckered specimens), and consequently some earthstar authorities “re¬
duce” A. pteridis to a variety of A. hygrometricus. For some inexplicable reason, the
fructifications remind me of “tribbles”—those lovable but relentlessly prolific creatures
of “Star Trek” renown.

706
 707

SCLERODERMA (Earthballs)
Medium-sized to fairly large fungi found in humus, soil, sand, or on rotten wood. FRUITING
BODY more or less round to oval, lobed, or with a stemlike base; rupturing irregularly in old age
or splitting into starlike rays or sometimes forming an apical pore or tear. PERIDIUM usually
one-layered and typically thick, tough, and rigid when fresh; smooth or wrinkled or with scales,
typically yellowish to brown but sometimes whitish. STERILE BASE absent, but a stalklike base
composed of tough mycelial fibers often present. SPORE MASS initially white and very firm,
but in most cases soon becoming gray to purple-black (sometimes marbled with paler veins) and
remaining firm, then eventually becoming powdery and sometimes browner. Spores more or less
round, spiny and/ or reticulate, not borne in a hymenium. Capillitium absent or rudimentary.

ALSO known as the thick-skinned or hard-skinned puffballs, the earthballs superficially

resemble the true puffballs(Calvalia, Lycoperdon, etc.), but differ in several fundamental
respects. Their peridium (skin) is typically hard (rindlike) when fresh and tough or
leathery in age, and the spore mass becomes colored (usually purple-gray to black) at
an early age while remaining firm. In the puffballs, on the other hand, the peridium is
often thin and papery in age and the spore mass becomes soft or mushy as it ripens
(darkens). There are other differences as well. In Scleroderma there are no specialized
threadlike cells (capillitium) in the spore mass, the exterior of the spore case lacks the spines
or soft particles found in many puffballs, and there is no sterile base, though a stalklike or
rootlike mass of tough mycelial fibers is often present. These differences have led most
taxonomists to group the Sclerodermas in a separate order, the Sclerodermatales, along
with several other genera such as Pisolithus.
As their name implies, earthballs are often buried or partially buried before maturity,
leading to possible confusion with the false truffles (Hymenogastrales & Allies). The
spore mass, however, becomes powdery in old age and the thick, tough peridium is dis¬
tinctive. The deer truffles (Elaphomyces) also have a thick peridium, but tend to grow
deeper in the ground, lack a distinct basal point of attachment, and bear their spores in asci.
Sclerodermas are common and ubiquitous, but are not as conspicuous as the true
puffballs. Their nomenclature is very confused. Over 150 species have been described
but in a 1970 monograph, Gaston Guzman condenses them into a mere 21 widespread,
polymorphic species. However, even with only 21 species to choose from, identification is
difficult without a microscopic examination of the spores. The genus is divided into three
“sections” based on whether the spores are echinulate (spiny), reticulate (covered with a
network of ridges), or a compromise between the two, and species in the different sections
resemble each other closely.
Many Sclerodermas are thought to be mycorrhizal, but they can grow almost any¬
where—in woods or at their edges, under planted trees and shrubs, in sand and asphalt,
along roads and trails, on exposed hillsides, around old stumps, etc. In our mild climate
they occur year-round, but seem to favor warm weather. Three representative species are
described here (one from each “section”) and several others are discussed. Sedecula, a rare,
smooth-spored, underground genus possibly related to Scleroderma, is also keyed out.

Key to Scleroderma
1. Spore mass (interior) divided into large irregular chambers by tough cordlike veins (which may
turn into coarse black mycelial cords at base); fruiting body 2-9 cm broad, oval to cushion¬
shaped; exterior often pitted or split in age, whitish to grayish to olive-buff or yellowish (but
often blackening in age); spore mass black and powdery to slightly gelatinous when mature;
spores 20-28.5 * 11-16 microns, smooth; widespread under western conifers (especially in
mountains), but rather rare; usually growing underground .Sedeculapulvinata
1. Not as above; mature spores ornamented; found underground or above; common . 2
2. Fruiting body growing underground, typically without an obvious basal point of attachment to
substrate; peridium (skin) sometimes marbled in cross-section; spores borne inside asci (but
asci soon disintegrating) . (set Elaphomyces, p. 862)
2. Not as above; fruiting above the ground, or if underground then usually with an obvious base or
point of attachment; peridium not marbled in cross-section; spores borne on basidia .... 3
708 LYCOPERDALES & ALLIES

3. Peridium (skin) very thick (3-10 mm), rupturing into starlike lobes in old age; fruiting body
medium-sized to large; spores partially reticulate .S. geaster& others, p. 710
3. Not as above; either peridium thinner (averaging 1-4 mm) or not rupturing into starlike lobes;
fruiting body fairly small to medium-sized (rarely large) .4
4. Fruiting body with a long “stalk” composed of tough strands and fibers that totals at least half
(and usually three-quarters) of the fruiting body’s total height; spores reticulate; found in
sand dunes or sandy soil .S'. macrorhizon(see S. citrinum, below)
4. N ot as above (“stalk” if present shorter and/ or habitat different) . 5
5. Peridium (skin) covered with prominent inherent rosette-like scales (i.e., each scale often with
a central wart); widespread, but especially common in forests (see Color Plate 190).6
5. Not as above; peridium typically smooth (at least when young) but often becoming fissured
or cracking and peeling to form scales in age . 7
6. Peridium (skin) rather thin (typically lessthan2 mm), usually rupturing in old age through a pore
or slit at top; spores spiny.S'. verrucosum & S. areolatum (see S. citrinum, below)
6. Peridium fairly thick (usually 1-4 mm), rupturing into lobes in age or forming an irregular
pore; spores reticulate ..S', citrinum, below
7. Spores spiny but not reticulate . S.cepagroup, p. 709
7. Spores reticulate or partially reticulate . S. bovista& others (see S. cepa group, p. 709)

Scleroderma citrinum (Common Earthball) Color Plate 190

FRUITING BODY 2-10 (12) cm broad and 2-6 cm high, round or somewhat flattened,
the underside often with a stemlike base composed of ridges, mycelial fibers, and/ or
adhering debris; eventually cracking into lobes to form an irregular pore, the lobes not
normally bending outward or unfolding appreciably. PERIDIUM (skin) hard and fairly
rigid (rindlike), 1 -3 mm thick, white when sectioned but usually staining pinkish (if fresh);
surface yellow-brown to dingy yellow to ochre or tan and cracked or arranged into
prominent, inherent scales which often have a smaller, central wart. SPORE MASS
white when very young, soon gray to purple-gray with whitish veins often running through
it, then dark purple-gray to purple-black or black and still solid and firm; eventually
becoming powdery and blackish-brown. Spores 8-13 microns, round, strongly reticulate.
HABITAT: Solitary, scattered, or in groups or clumps on ground and rotten wood,
usually in forests but sometimes in gardens or in sandy or disturbed soil; widespread and
the most common of all Scleroderma species. Although it is more typically a northern
than a southern species, I have seen enormous fruitings in the Great Smoky Mountains
of Tennessee. In our area I have found it in a garden in October, but it is rare—at least
in comparison to other species.
EDIBILITY: Poisonous! If eaten raw or consumed in quantity it causes nausea, vomiting,
diarrhea, and even chills or cold sweats. Nevertheless, it has been used by some unscrupu¬
lous individuals to adulterate truffles.
COMMENTS: Also known as S. aurantium and S. vulgare, this earthball is easily
recognized by its scaly, yellow-brown, rindlike skin that tends to stain pinkish when cut,
and the purple-black spore mass (see color plate). It might be mistaken for an edible puff¬
ball, but is only rarely found while still white inside. The scales often looked embossed
(i.e., with a central wart), a character that helps distinguish it from scaly specimens of
S. cepa, S. bovista, etc. (see comments under S. cepa). Other species: S. macrorhizon has a
long “stalk” composed of tough strands and fibers and grows exclusively in sand (often
in sand dunes!). It is widely distributed in northern latitudes and has spores similar to
those of S. citrinum, but slightly larger. S', verrucosum and S. areolatum (-S. lycoper-
doides) are smallish species with a thin (up to 1 mm thick), delicately scaly peridium that
usually ruptures in old age through an apical pore or tear. Both have spiny rather than
reticulate spores, and a stalklike, fibrous base, and are widely distributed under hard¬
woods or sometimes conifers. (In S. verrucosum the spores are 7-11 microns in diameter;
in S. areolatum they are 10-18 microns.)
 "'•N

Scleroderma cepa group. Note thick, relatively smooth skin and dark (purple-black) interior when
young. Also note how the older specimen at right is beginning to split open. These specimens stained
burgundy when rubbed, especially near base (staining is visible in middle specimen).

Scleroderma cepa group (Smooth Earthball)

FRUITING BODY 1.5-8 (10) cm broad and/or high, more or less round to somewhat
flattened or lobed (often broader than it is high), the underside often—but not always—
with a rootlike or stalklike base made up of tough mycelial fibers and adhering debris;
eventually rupturing irregularly or splitting at the top into lobes which may peel back
slightly. PERIDIUM (skin) hard and fairly tough (rindlike) when fresh, 1-3 mm thick,
white in cross-section but often staining reddish-pink to vinaceous when cut (if fresh).
Surface whitish or pallid, soon becoming buff, yellowish, straw-colored, or brownish,
often staining reddish or vinaceous when rubbed (especially the underside), then dis¬
coloring brownish; smooth when young, often becoming areolate (cracked to form scales)
in age or where exposed to light (especially on top). SPORE MASS white and very firm
when young, soon becoming black or purple-black (sometimes with paler veins) while
remaining firm, then eventually becoming powdery and somewhat paler or browner.
Spores 7-12 microns, round, spiny but not at all reticulate.
HABITAT: Solitary, scattered, or gregarious under trees (both hardwoods and conifers),
in woods or cultivated areas, along roads, in gardens, lawns, etc.; widely distributed.
In our area this species and its numerous look-alikes (see comments) are common year-
round in a variety of habitats, but seem especially prevalent underpineand/ oreucalyptus.
EDIBILITY: Poisonous! Chanterelle picker and Russula-kicker Bob Sellers of Santa
Cruz, California, ate a small piece of S. laeve (see comments) under the impression that all
“puffballs” were edible. Twenty minutes later he broke into a cold sweat, felt nauseous,
and started vomiting. After expelling the offender, he recovered quickly.
COMMENTS: There are a number of microscopically distinct Sclerodermas that will
more or less fit the above description. Collectively they can be recognized by their pallid
to buff or yellow-brown peridium that is initially smooth but may crack into scales as it

Scleroderma cepa group. These specimens differ from those in above photo in their more highly
developed “stalk” composed of tough fibers. Microscopically, however, they are indistinguishable.
Left: Fairly young specimens. Right: An older individual which has split into lobes or “rays.”
710 LYCOPERDALES & ALLIES

is exposed (the scales are not inherent as in S. citrinum), by their tough rindlike skin, and
by their firm purple-black spore mass when young. Features such as the tendency of the
peridium to stain reddish when rubbed and the presence or absence of a “stalk” composed
of mycelial fibers seem to vary according to environmental conditions. Other species:
S. flavidum is considered by some Scleroderma-scholars to be a form of S. cepa. It is
microscopically identical, but does not usually bruise reddish and normally splits open
into starlike lobes which bend back to expose the spore mass. S. laeve closely resembles
the “true” S. cepa, but has slightly larger spores (9-15 microns); it appears to be the most
common Scleroderma in our area. S. albidum is also similar, but has even larger spores
(12-17 microns). S’, reae, which favors arid habitats and ruptures irregularly, can be dis¬
tinguished microscopically by its partially reticulate spores (9-18 microns). S.floridanum
also has partially reticulate spores, but is tropical and subtropical and ruptures stellately
(i.e., splits into starlike lobes). There are also four farflung Sclerodermas with completely
reticulate spores: S. bovista, which sometimes develops blackish spots or stains in old
age and has spores 9.5-16 microns broad;S. fuscum, which favors conifers and has spores
averaging 13.5-19.5 microns; S. hypogaeum (-S. arenicola), a common vinaceous-
staining conifer-lover that often grows underground and has an unusually thick peridium
and spores 18-23 microns broad; andS. michiganense, which has spores like-S', hypogaeum
and often grows underground, but has a thinner peridium and favors hardwoods. All of
the above species are widespread and vary considerably in size, form, and scaliness, and all
but the latter species may stain reddish or vinaceous when bruised. They should be con¬
sidered poisonous until proven otherwise. Also see the species listed under S. citrinum.

Scleroderma geaster (Dead Man’s Hand)

FRUITING BODY 4-15 cm broad or high when closed, 12-30 cm broad when expanded
(after rupturing); at first round to oval (but often somewhat flattened or irregularly lobed),
eventually splitting into several (usually 4-8) coarse, irregular, thick rays which curl back
in starlike fashion to expose the spore mass. PERIDIUM (skin) very thick(3-10 mm) and
firm and tough when fresh; exterior whitish becoming yellowish, straw-colored, or
brownish, roughened or somewhat hairy, often with adhering debris and often fissured
or cracked into scales, especially in age; interior (upper side of rays) brownish becoming
blackened and empty in old age; base often attached to the soil by a mycelial “root” of
tough fibers. SPORE MASS at first firm and pallid, soon dark gray to purple-black
or black (and still firm), eventually becoming brown to dark brown or purple-brown

Scleroderma geaster (-S. polyrhizon), mature but rather small specimens that have split into rays.
In this stage they might be mistaken for old cup fungi, but usually contain traces of spore powder.
 ■V

Scleroderma geaster (=S. polyrhizon), young specimens which have yet to split open. Note how thick
the skin is! These were found along a road with Pisolithus tinctorius {p. 712).

and powdery. Spores (5) 6-11 (12) microns, round, with warts or spines that often form
incomplete lines or ridges (i.e., partially reticulate). Peridium containing few if any thick-
walled hyphae.
HABITAT: Solitary, scattered, or in groups on hillsides, along roads, in ditches, poor
soil, sand, asphalt, gravel, etc.; often buried or partially buried before maturity. It is
widely distributed and quite common in our area, especially in sandy soil. It usually ap¬
pears in the fall, but the thick tough skins take a long time to decompose.
EDIBILITY: Poisonous? This is one fleshy fungus I’ve never been tempted to eat!
COMMENTS: The large size and tough skin that splits into coarse, thick rays are the
telltale traits of this bizarre fungus. It is also known (more correctly) as S. polyrhizon
or S. polyrhizum. It rivals Pisolithus tinctorius (“Dead Man’s Foot”) for grotesque¬
ness. The two thrive in similar milieu—asphalt, sand, poor soil, etc.—and make a well-
matched if not exactly charming couple. It might be mistaken for a large cup fungus (e.g.,
Sarcosphaera), but it usually has traces of the powdery spore mass to distinguish it. The
rays lack the intact inner spore case characteristic of the earthstars, and there may or
may not be a stemlike base of mycelial fibers. It can also be confused with Mycenastrum
corium, but that puffball has a thick white felty outer peridium and a smooth, purple-
brown inner one which splits into lobes at maturity. Other species: S. texense (-S. fur-
furellum) is a very similar, thick-skinned species with a scalier (often shingled) exterior
and thick-walled hyphae in the peridium; it hasa moresoutherly(tropicaland subtropical)
distribution but also occurs on the west coast. S.flavidum(see comments under the S. cepa
group) also splits in starlike fashion, but is muchsmaller, withathinner(about 1 mm thick)
peridium that is tawny and more or less smooth.

PISOLITHUS & DICTYOCEPHALOS (Oddballs)

Medium-sized to fairly large terrestrial fungi found mostly in poor soil. FRUITING BODY
variously shaped at maturity. PERIDIUM variously colored and rupturing irregularly in old age.
STERILE BASE absent, but a stalk often present. VOLVA absent (Pisolithus) or often present
(Dictyocephalos). SPORE MASS either composed of numerous small capsules (peridioles) or
divided up into numerous "cells” or chambers; brown and powdery at maturity, the peridioles or
“cell” walls mostly disintegrating. Spores more or less round, warty or spiny. Capillitium absent.

THESE two oddballs are easily distinguished from puffballs and earthballs by the struc¬
ture of their spore mass. In Pisolithus there are hundreds of small spore-containing cap¬
sules (peridioles) imbedded in the fruiting body. The peridioles disintegrate, however,
so they are best seen by making a lengthwise section of a fairly young specimen. The

711
712 LYCOPERDALES & ALLIES

fruiting body is internally sticky when young, but at maturity it becomes crumbly and
protrudes from the ground like a dusty stump, half-rotted root, or ball of dried-up dung.
Dictyocephalos is just as unsightly, but has a long, woody stalk. It is likely to be mistaken
for a stalked puffball at first glance, but its spore mass is divided into numerous “cells”
or chambers. Most of the chamber walls disintegrate, but the lowermost ones can often
be seen after the spore mass has dispersed. In each genus there is a single variable species
with a worldwide distribution.

Key to Pisolithus & Dictyocephalos

1. Fruiting body with a long, tough or woody stalk; volva often present at base of stalk; spore mass
chambered when young; found mostly in deserts .Dictyocephalos attenuatus, p. 713
1. Fruiting body lacking a volva and true stalk (but often with an elongated stalklike sterile base);
spores borne in numerous lentil-like capsules imbedded in upper part of fruiting body (best
seen when young); widespread in poor soils, along roads, etc. . . Pisolithus tinctorius, below

Pisolithus tinctorius (Dead Man’s Foot) Color Plates 185,189

FRUITING BODY 5-30 cm or more high and 4-20 cm broad, at first round to pear-
shaped, then club-shaped (i.e., usually but not always with a narrowed, rooting base or
“stalk”), finally breaking up in age or taking on the appearance of a large dusty root or
stump; odor mild to aromatic or unpleasant, depending on the age. SPORE CASE with
a thin, brittle peridium (skin) that is variously colored (but usually yellowish, purplish,
olive-black, or brown) and often lustrous and soon ruptures irregularly or flakes away.
U pper portion of fruiting body containing hundreds of seedlike peridioles which gradually
disintegrate, turning into a crumbly or dusty mass of brown spores (the disintegration
process starts at the top of fruiting body and proceeds downward). Peridioles small {2A
mm long), elongated to oval or circular, whitish to greenish-yellow, yellow, brownish, or
vinaceous, at first imbedded in a sticky dark or blackish substance which dries out and
becomes brittle or crumbly at maturity. Lower portion of fruiting body typically con¬
sisting of a fibrous, stalklike, persistent, sterile, rooting base which may have coarse
greenish-yellow mycelial fibers attached. VOLVA absent. SPORE MASS powdery when
mature, dark brown to cinnamon-brown. Spores 7-12 microns, round, warty or spiny.
HABITAT: Solitary, widely scattered, or in small groups on or along roads, in waste
places, and in hardpacked, poor, sandy, or gravelly soil; very widely distributed, but
especially common in California and the Pacific Northwest (the largest fruitings I’ve seen
were in the Siskiyou Mountains of northern California). In our area it fruits chiefly in
the late summer and fall, but is slow to decay and can be found most any time. It forms
mycorrhiza with a wide range of trees and shrubs(oak, pine, etc.) and is said to be of value
in reforestation projects.
EDIBILITY: Not recommended. In Europe it is known as the “Bohemian Truffle” and
used as an aromatic seasoning when unripe, and in China it is employed medicinally. It
is scarcely appetizing, however, and should be tried very cautiously, if at all. The name
tinctorius reflects its use as a dye—a variety of rich colors (mostly browns and golds, but
also blacks and dark blues) can be obtained, depending on the mordants used and the
type of soil in which it is found.
COMMENTS: This dusty monstrosity is among the most distinctive and memorable of
all the fleshy fungi. Young specimens are somewhat puffball-like (i.e., pear- or club-
shaped), but are easily told by the hundreds of seedlike peridioles they contain (see Color
Plate 185). In this stage the peridium often has a beautiful metallic luster. Older spe¬
cimens, on the other hand, can be recognized from a great distance by their dusty
brown stumplike stance. Fresh specimens will stain practically anything they come into
Pisolithus tinctorius. When young this monstrosity is puffball-shaped, but its outer skin ruptures
(left), exposing the powdery brown spore mass. Old specimens protrude from the ground like dusty
stumps (right), and bear little resemblance to puffballs or any other fleshy fungi. For other views of
this distinctive fungus (including a close-up of the lentil-like spore capsules), see the color plates.

contact with, and the dry spore dust coats everything in the vicinity. This species has a
penchant for adversity, inevitably fruiting in poor soil, ditches, chaparral, or road cuts
(often in the company of “Dead Man’s Hand,” Scleroderma geaster), or even bursting up
through asphalt. As might be expected, it is disdained by many mycologists (“It is the most
objectionable of all fungi“This ugly, stinky fungus...”), but to me it is one of the most
enthralling—if not beautiful—of all fungi (I call it “Dead Man’s Foot” not to demean it,
but because it looks like one). Its wide distribution and variable features have resulted
in a plethora of aliases, including/*. arenarius,P. arrhizus, and Polysaccumpisocarpium.

Dictyocephalos attenuatus (Stalked Oddball)

FRUITING BODY beginning underground as an “egg” completely encased in a mem¬
brane, the stalk then elongating rapidly and the ruptured outer membrane forming a volva
at its base. Mature fruiting body consisting of a spore case mounted on a tough stalk.
SPORE CASE round to somewhat flattened or even depressed on top, or irregular(e.g.,
sometimes with several lobes), 2-6 cm high and 5-13 cm broad, seated on a disclike enlarge¬
ment of the stalk apex which is white to brownish in old specimens. Outer peridium(skin)
fleshy to slightly gelatinous when young, becoming tougher and developing brown to
reddish scales or warts which vary from small, flat, and more or less persistent to large,
pyramidal and deciduous; lower margin sometimes with a hanging “veil” composed of
adhering tissue stripped from the stalk during expansion. Inner layer of peridium 1 -2 mm
thick, usually paler( whitish to buff) than outer peridium, but sometimes darkeninginage.
Upper portion of spore case breaking up into irregular pieces which fall away; lower
portion tougher and persisting as a more or less bowl-shaped structure. STALK (5)
10-40 (52) cm long, 2-5 cm thick at apex, equal or more often tapered downward (rarely
thicker below or bulbous), the base sometimes pointed and free of the volva, sometimes
with 1 -2 rootlike mycelial cords attached; solid (or with insect cavities), sometimes forked
(with one spore case on each branch), fleshy-tough becoming woody when dry; at first
white, but becoming brown in age; often grooved, ridged, and/ or flattened and sometimes
twisted; often cracking, peeling, or curling back to form scales or sometimes even an
irregular “ring” or “annulus.” VOLVA at base of stalk saclike, but occasionally poorly
developed or absent; white to tan when fresh, 3-11 cm high; tough, hard, or chalky in age.

713
Dictyocephalos attenuatus. Left: Two typical dusty, mature specimens. The one on the left has lost
most of its spore case but retains its volva, while the one on the right still has its spore case but has
lost its volva. Note the well-developed stalk in both specimens. Right: A close-up of the lower part of
a mature spore mass, showing the hairy or fibrous remains of the walls that originally divided the
spore mass into numerous small chambers (a feature which separates it from the stalked puffballs).

SPORE MASS divided up into numerous whitish-walled “cells” (peridioles), but all but
the lowermost “cell” walls disintegrating, so that the mature spore mass is brown and
powdery, usually with an unpleasant odor (like decaying fish); lowermost “cell” walls
often persisting after the spores have been dispersed as flattened, pointed “teeth” on the
inner surface of the spore case, giving it a pitted or reticulate appearance. Spores 5-7
microns, round or nearly round, warted or spiny.
HABITAT: Solitary, widely scattered, or in small groups(often in tufts of 2-5 individuals
and occasionally two emerging from a single volva) in barren sandy soil or clay, waste
places, etc. It is widespread in the arid and semi-arid parts of the West, but is most com¬
mon in the deserts of southern California, the Four Corners area, and the Southwest—
often near or under saltbush (Atriplex) or along washes. It fruits after seasonal rains, but
the fruiting bodies are practically impervious to bacterial and fungal decay, persisting
for as long as 30 years in their natural environment! Mature specimens can protrude
from the soil like dusty roots (in the grand tradition of Pisolithus tinctorius), or remain
underground, depending on the hardness of the soil and weather conditions. W. H. Long,
who gathered a gaggle of them near Antelope Valley, California, reports that developing
specimens raised hard blocks of soil weighing 15 pounds each!
EDIBILITY: Much too crusty, musty, and dusty to be worthwhile, but possibly useful
as a dye (see Pisolithus tinctorius).
COMMENTS: As can be surmised from the description, this “oddball,” like Pisolithus
tinctorius, exhibits a great deal of variation in size, shape, color, and degree of scaliness
—in other words, it is not difficult to identify but is easy to misidentify. Its polymorphism
has resulted in the naming of dozens of “new” species, when in fact there is only one highly
variable one. It can be distinguished from the stalked puffballs and other desert fungi by

714
DICTYOCEPHALOS 715

its chambered spore mass, the pitted-reticulate upper surface of old (empty) spore cases,
the long and tough or woody stalk, plus the frequent presence of a volva and irregular
breaking up of the peridium (a distinct apical pore is not formed). Its closest relative,
Pisolithus tinctorius, lacks the well-developed stalk and volva, and does not usually
grow in such desolate places.

Stalked Puffballs
spores
TULOSTOMATALES
Small to medium-sized puffballs often found in arid habitats or in sand or poor soil. FRUITING
BODY with a spore case mounted on a well-developed, differentiated stalk. SPORE CASE typi¬
cally with a two-layered peridium, rupturing apically, peripherally, stellately, or irregularly.
STERILE BASE absent. STALK not percurrent; usually tough or woody and often hairy or scaly
(but gelatinous in Calostoma). VOLVA present or absent. SPORE MASS powdery at maturity
and buff to rusty-salmon to cinnamon-brown or dark brown. Spores typically round and orna¬
mented, but occasionally smooth. Capillitium usually present, well-developed.

THESE are puffballs with a clearly differentiated, well-defined stalk. As in the true
puffballs (Lycoperdales), the mature spore mass is powdery and the peridium (skin) that
encloses it is composed of at least two layers. The presence of a stalk can lead to confusion
with the gastroid agarics (Podaxales & Allies, p. 724), but in those fungi the stalk is per¬
current, i.e., it extends through the spore mass to the top of the spore case or “cap,”
just like the stalk of a gilled mushroom. In the stalked puffballs, on the other hand, the
stalk terminates at the spore case rather than extending through it.
Tulostoma, with over 30 species, is the largest and most cosmopolitan genus of stalked
puffballs, yet it is by no means common. It can aptly be characterized as “a puffball on a
stick” because that’s just what it looks like, with its slender stalk and its small round spore
case with an apical pore. There are six other genera of stalked puffballs, none with more
than five North American species and several with only one. Calostoma is the most out¬
landish of the lot, with its brightly colored gelatinous fruiting body. Battarrea is also
unmistakable, for its outer peridium forms a volva at the base of the stalk and the spore
case ruptures around its rim or through several holes. Chlamydopus, like Tulostoma,
forms an apical pore, but lacks the ball-and-socket relationship of stalk to spore case
that characterizes that genus. Schizostoma ruptures stellately (that is, splits into several
lobes), while Phellorina and Queletia rupture irregularly. An eighth genus, Podaxis, is
often treated as a stalked puffball, but in this book is grouped with the gastroid agarics
because it has a percurrent stalk.
With the notable exception of Calostoma, the stalked puffballs tend to grow inenviron¬
ments that most fleshy fungi find inhospitable—sand, poor soil, waste places, etc. Several
like it hot and are found strictly in the desert, where they form a fairly large percentage of
the fungi in the arid wastelands of the Great Basin and Southwest. Calostoma, as already
noted, is the exception. It prefers the humid forests of eastern North America and is
particularly common in the southern Appalachians.
None of the stalked puffballs are known to be poisonous. None are known to be edible
either, because most of them spend their youth underground (a necessary adaptation to
their harsh environment). As a result, they are invariably encountered after their spore
mass is mature and powdery (and thus inedible), and the stalks are much too tough to eat,
unless you’re starving and barefoot (in other words, you’re better off chewing on a shoe!).
Six stalked puffballs are fully described here, and several others are keyed out.
716 TULOSTOMATALES

Key to the Tulostomatales

1. Stalk (and often spore case) with a viscid or gelatinous outer layer when fresh; spore case rup¬
turing through an apical pore at maturity; confined to humid regions, especially southeastern
North America . 2
1. Not as above . 4
2. Spore case red when fresh (but may fade to yellow or buff) . . Calostoma cinnabarina, p. 718
2. Spore case yellow to buff when fresh (but mouth may be red) . 3
3. Spore case with a viscid or gelatinous outer layer when fresh .
. Calostoma lutescens (see C. cinnabarina, p. 718)
3. Spore case not viscid . . . Calostoma ravenelii& C. microsporum (see C. cinnabarina, p. 718)
4. Spore case rupturing around its lower periphery, the top falling off like a lid in old age (as shown
in photo on p. 717); stalk usually long, often scaly at maturity .
..Battarreaphalloides & others, p. 717
4. Not as above; spore case not rupturing around its lower periphery .5
5. Cap cylindrical (shaggy mane-shaped); stalk extending into (and often through) the spore
mass .(see Podaxis, p. 725)
5. Not as above . 6
6. Spore case typically rupturing through a single small mouth at the top (an apical pore) ... 7
6. Spore case rupturing through several pores or by splitting into lobes or by irregular disinte¬
gration . 11
7. Stalk comprised of a mass of tough rootlike fibers; spore mass often purplish or black when
young and firm (but often browner when powdery) .(see Scleroderma, p. 707)
7. Not as above . 8
8. Stalk thickest at apex, not inserted into the spore case like a ball-and-socket; volva present
when fresh; found mostly in deserts .Chlamydopus meyenianus, p. 721
8. Stalk sometimes thicker above but usually equal, with a ball-and-socket-like attachment to
the spore case (but not necessarily separating easily); volva present or absent; found in many
habitats, including deserts . 9
9. Fruiting body medium-sized (spore case typically 1.2-3.5 cm broad; stalk typically 0.5-2 cm
thick) .Tulostoma macrocephalum & others, p. 720
9. Fruiting body usually rather small and slender (spore case typically up to 2 cm broad; stalk
typically 2-6 mm thick) . 10
10. Stalk with a volva at base and one or more “roots” below the volva; spore case white or whitish;
found in desert .Tulostoma cretaceum (see T. brumale group, p. 719)
10. Not with above features; volva present or absent; found in many habitats (including desert)
.Tulostoma brumale group & many others, p. 719

11. Spore case rupturing through several holes (pores); volva usually present when fresh; found
in deserts .Battarrea digueti (see B. phalloides, p. 717)
11. Spore case disintegrating, rupturing irregularly, or splitting into several lobes; volva present
or absent; found in many habitats (including deserts) . 12
12. Stalk continuous with spore case (i.e., bottom of spore case merely an expansion of stalk) 13
12. Stalk and spore case distinct, with a ball-and-socket-like attachment. 15
13. Fruiting body full of spore-containing capsules when young and protruding from ground like a
dusty stump in old age or spore mass composed of numerous cells or chambers (when young)
whose walls usually give a pitted, reticulate, or hairy appearance to inside surface of bottom of
spore case in old specimens; volva present or absent (see Pisolithus& Dictyocephalos, p. 711)
13. Not as above; spore mass lacking capsules, chambers, or “cells”; volva typically absent . . 14
14. Stalk composed of tough rootlike strands or fibers; spore mass often purplish to black when
young and firm (but often browner when powdery); found in many habitats .
.. (see Scleroderma, p. 707)
14. Not as above; found in deserts . Phellorina strobilina, p. 723
15. Spore case rupturing Stellately (i.e., splitting into several starlike rays or lobes) when old; found
in deserts .Schizostoma laceratum (see Chlamydopus meyenianus, p. 721)
15. Spore case rupturing irregularly or disintegrating; very rare (reported from Pennsylvania)
.Queletia mirabilis (see Chlamydopus meyenianus, p. 721)
Battarrea phalloides. Left: A mature specimen in which the spore case has ruptured around its
perimeter and the top has fallen off. Note large volva at base. Right: Old specimens, some of which
have lost their volvas. Note how stalks vary considerably in thickness and degree of scaliness.

Battarrea phalloides (Scaly-Stalked Puffball) Color Plate 187

FRUITING BODY initially underground and completely enclosed by a membrane
(outer peridium) which ruptures around its periphery, the stalk then bursting through and
rapidly elongating. Mature fruiting body composed of a spore case (the inner peridium)
mounted on a long stalk, with the ruptured outer peridium leaving a volva at base of stalk
and sometimes a volval patch on top of the spore case. SPORE CASE 3-5 (6) cm broad
and 1-3.5 cm high, typically rounded (convex) on top, with a flat or even depressed (con¬
cave) underside that somewhat resembles a “veil”; surface smooth, pallid or whitish,
soon splitting around its lower periphery so that the upper (rounded) portion falls off,
thereby exposing the spore mass; lower portion remaining intact as a disc beneath the
spore mass and sometimes persisting (after spore mass has dispersed) as a disclike “cap”
or sliding down the stalk to form a “ring.” STALK 10-40 cm long, 0.4-1.5 cm thick, not
percurrent, equal or tapered downward (but volva-encased base may be swollen); rigid
and tough or even woody, hollow, white to brownish or rusty-brown, longitudinally
striate or grooved and covered with fibers that often peel or split to form fine to very coarse
needlelike, ribbonlike, or shaggy scales. VOLVA at base of stalk saclike, whitish or
dirt-incrusted, often loose, buried in ground and soon rotting away. SPORE MASS
rusty-brown and powdery when mature, adhering to everything it comes in contact with;
odor sometimes unpleasant in old age. Spores (4) 5-8 (10) microns, round to broadly
elliptical, warted. Capillitium of two types: simple, short hyphae up to 10 microns long,
and elators (elongated cells with internal spirals) up to 100 microns long.
HABITAT: Solitary, widely scattered, or in groups in sand, poor soil, and waste places
(but often under trees in deep shade); very widely distributed, but in the United States
apparently confined to the West. It is generally regarded as rare, but in the deserts and
sagebrush country of the Great Basin and Southwest it is relatively common for a desert
fungus. It also occurs along the California coast and Orson Miller reports it from Alaska.
In our area it fruits during the rainy season but persists for months without decaying.
EDIBILITY: Inedible

717
Battarrea phalloides, old specimens bereft of volva and spores. Only the bottom of the spore case
remains, forming a thin “cap” on the stem. In this condition they are as light as straws.

COMMENTS: The fibrous-scaly stalk, presence of a volva (which, however, often rots
away or can be left behind in the ground), and curious manner in which the spore case
ruptures around its periphery are the telltale traits of this intriguing fungus. The flattened
or concave, veil-like underside of the spore case is also unique. The upper portion or“lid”
usually detaches completely; it can often be found on the ground nearby, but is some¬
times blown away by the wind. The volval patch (when present) may tear away from the
“lid,” leaving a gaping hole as shown in the photograph. The stature of the fruiting body
varies considerably according to soil and weather conditions. Some specimens(especially
the coastal ones) have relatively thick, coarsely scaly stalks, whereas others (particularly
desert dwellers) have longer, thinner stalks. All sorts of intergradations can be found,
however. Some authorities claim that the “true” B. phalloides of Europe has a gelatinous¬
layered volva in the egg stage. If this is really the case, then the American species, which
lacks the gelatinous layer, is more correctly called B. stevenii. (Since the “eggs” develop
deep in the ground, they are rarely encountered and it is difficult to know whether or not
this distinction is a valid one.) Another questionably distinct species, B. laciniata, has a
more persistent, multi-layered volva with inner, concentrically-arranged “leaflets”
around the stalk base. It usually has a white to buff or reddish volval patch on top of the
spore case and tends to be more robust (stalk 2A cm thick at apex and spore case 4-8 cm
broad, 2-4 cm high). It occurs in southern California and the Southwest, usually in “rich
loamy alkaline soil” in the open desert (not under trees). Still another species, B. digueti,
is definitely distinct, for it does not rupture by peripheral cleavage. Instead, its outer
peridium ruptures apically (thus never leaving a volval patch) and its spore case remains
intact, with several holes forming in the upper (convex) portion. It also has longer elators
than B. phalloides, but in other respects (shape and overall appearance) is quite similar.
It appears to be strictly a desert fungus. I have found it in Baja California, and it is also
known from southern California, mainland Mexico, and the Southwest.

Calostoma cinnabarina (Red Slimy-Stalked Puffball) Color Plate 188

FRUITING BODY consisting of a spore case mounted ona thick, short, fluted stalk(the
spore case often barely protruding above the ground). SPORE CASE 0.5-2 cm broad,
more or less round to somewhat oval, at first covered by the outer peridium, which is
composed of a thick, gelatinous or slimy, translucent layer and a thin, bright red inner
layer; outer peridium breaking up into pieces and falling away (the pieces of the inner
layer often imbedded in the gelatinous pulp of the outer layer so as to resemble small
tomato or pomegranate seeds). Inner peridium persistent, not gelatinous, at first red and
scurfy or powdery, but often fading to orange, yellow, or buff as the scurf wears away;
rupturing through a crosslike apical pore (mouth) formed by 4-5 dark red to scarlet ridges
or “teeth.” STALK 1.5-4 cm long, 0.5-2 cm thick, typically thick and short, more or less
equal, spongy, with an outer gelatinous layer when fresh; otherwise composed of firm,
branching strands that form a ridged, netted, and/ or pitted pattern; ochraceous to red or
cinnabar-red to yellow-brown, often fading in age. VOLV A absent. SPORE MASS white
at first, becoming powdery and buff-colored at maturity. Spores 14-28 * 6-11 microns,
elliptical to oblong, pitted. Capillitium present when young but soon disintegrating.

718
Calostoma cinnabarina. Since these mature specimens are dried out, the gelatinous layers evident in
the color plate do not show. The bright red color (when fresh) and thick, fibrous stalk are distinctive.

HABITAT: Solitary to gregarious in soil and humus in woods, along roadcuts, under
trees, etc.; fairly common in the late summer and fall in the southern and eastern United
States, especially at higher elevations. It occurs as far west as Texas, and is to be looked
for in southern Arizona and New Mexico, where several “eastern” mushrooms occur.
EDIBILITY: Unknown—but too small and slimy to merit experimentation.
COMMENTS: In a group (the puffballs) not noted for its bright colors, the genus Calo¬
stoma stands out. The complex structure of the outer and inner peridiums (each has at
least two layers) and peculiar microscopic features such as the absence of capillitium in
the mature spore mass have made it difficult for taxonomists to relate it to other puffball
genera. The thick gelatinous outer layer, which can be likened to a universal veil, dis¬
tinguishes Calostoma from other stalked puffballs, and the powdery spore mass separates
it from the stinkhorns, which may also be slimy. C. cinnabarina is the only red Calostoma.
Others—all southeastern in distribution-—include: C. lutescens, with a longer stalk that
usually elevates the spore case above the ground, and a light to bright yellow spore case
(except for the red mouth or pore) that usually has a wide collar at its base formed by the
outer peridium; C. ravenelii, a smaller species with a gelatinous-fibered stalk and non-
gelatinous spore case that is tan to yellowish (with a red mouth) and often decorated with
warts left by the outer peridium; and C. microsporum, rather similar to C. ravenelii but
slightly larger, with smaller spores (up to 11 microns long).

Tulostoma brumale group (Common Stalked Puffball)

FRUITING BODY consisting of a small spore case mounted on a stalk. SPORE CASE
round (but often somewhat collapsed or flattened), 1 -2 cm broad, the underside shrinking
as it dries and often pulling away from the stalk to form a collar. Outer layer of peridium
(skin) brownish, but covered with sand and other debris, peeling away to reveal the inner
layer, but often remaining intact around the base. Inner layer smooth or with minute
particles, tan to pinkish-buff to buff, gray, or whitish, developing an apical pore at ma¬
turity; pore usually encircled by a raised collar or tube. STALK 1-6 cm long, 2-4 (6) mm
thick, usually slender, equal except for a small bulb at base; tough but usually pliant,
covered at first with coarse brown to rusty-brown fibers, but sometimes smoother and/ or
paler in age; longitudinally striate, joined to spore case via a ball-and-socketarrangement.
VOLVA absent or inconspicuous. SPOREMASS rusty-salmon and powdery at maturity.
Spores 3-5 microns, more or less round, minutely warted. Capillitium well-developed.
HABITAT: Scattered or in groups in sand and gravel, sandy soil, and other waste places
(even in the mortar of brick walls!); very widely distributed, but not particularly common.

719
Tulostoma berteroanum (see comments under T. brumale group) is probably the most common
stalked puffball in coastal California. Left: Stocky specimens. Right: More slender ones. Note the
small pore that forms at the top of each spore case.

In our area this species and its close relatives (see comments) fruit mainly in the fall and
winter, but the weathered fruiting bodies can be found most any time.
EDIBILITY: Inedible when mature, and rarely found in the immature stage.
COMMENTS: Tulostoma species are easily recognized by their slender stalk, small
spore case, and rusty-salmon spores. The fruiting body develops underground as in Bat-
tarrea, and the stalks of mature specimens are often buried so that only the spore case
is visible. The absence of a volva in most species plus the small size and formation of an
apical pore distinguish Tulostoma from Battarrea, and the stalk is not as consistently
thickened at the apex as it is in Chlamydopus. There are several closely related Tulostomas
that will more or less fit the above description, including T. berteroanum (see photo), the
most common in our area. They are partial to sandy soil and are best differentiated
microscopically (in other words, leave them to the Tulostoma taxonomists). Some of the
more widespread species are: T. striatum, which has ridged spores and an outer peridium
that leaves an acornlike cup around the lower half of the spore case; T. simulans, in which
the spore case is persistently covered by adhering sand particles and other debris; T.
campestre, also sand-incrusted, but lacking a well-defined tube around the apical pore; and
T. fibrillosum, a fairly common species which also lacks a tube but has practically smooth
spores and a longer, coarsely hairy stalk.
There are also several species apparently endemic to the Southwest (or at least to arid
regions), including: T. involucratum, common, with a membranous (not granular) outer
peridium that often forms a frilled cup around the inner peridium plus a tubular apical
pore; T. opacum, a rare species with large spores (7-11 microns); T. meristostoma, small,
with a slit or irregular tear instead of a raised apical pore or tube even when freshen many
species the pore becomes slitlike or irregularly torn after weathering); T. cretaceum, a
common and highly distinctive sand-loving species with a chalk-white spore case and a
stalk that tapers downward to a volva beneath which are one or more rootlike processes
(at least in most specimens); and T. excentricum, with an off-center tubular mouth and
inconspicuous volva plus a spore case which is not white unless weathered. All of these
species differ from Chlamydopus meyenianus in the ball-and-socket relationship between
the spore case and stalk (though the two may be firmly attached), and all are rather small
and slender-stemmed. For larger species, see T. macrocephalum.

Tulostoma macrocephalum (Fat-Headed Tulostoma)

FRUITING BODY consisting of a spore case mounted on a stalk. SPORE CASE 1.2-3
cm broad and 0.8-1.5 cm high, nearly round to somewhat flattened (usually broader than
it is high), firmly attached to the stalk. Outer layer of peridium (skin) grayish-white to buff
and soon falling away to reveal the inner layer, but often remaining intact around the base.
Inner layer smooth, dingy whitish, tough, developing a short tubular apical pore(mouth)

720
Left: Tulostoma macrocephalum has a thicker stem and larger “head” than most Tulostomas.
Right: Chlamydopus meyenianus, rather small specimens in which the rough outer peridium(skin)
has completely worn away, leaving the beautiful smooth inner layer. Note volva in specimen on right.

at maturity. STALK 5-15 cm long, 0.5-1.5 cm thick, usually rather long, equal or tapered
slightly toward the base, which usually has a bulb; tough or woody, usually scaly and/ or
transversely cracked. VOLVA absent or present only as a small fragile dirt-incrusted
sack or collar at base of stalk. SPORE MASS rusty-salmon to cinnamon-brown and
powdery at maturity. Spores 4-5.5 microns, round, warted. Capillitium present.

HABITAT: Solitary to gregarious in sand or sandy soil in arid regions, fruiting most any
time if rainfall is sufficient. It was originally found in the gypsum dunes of White Sands
National Monument in southern New Mexico (I have seen it there), but has also turned
up in southern California and can probably be found throughout the Southwest.
EDIBILITY: Inedible because of its toughness.
COMMENTS: The broad head, scaly stalk, and relatively large size (for a Tulostoma)
are the hallmarks of this distinctive stalked puffball. The presence of an apical pore dis¬
tinguishes it from Battarrea, and the equal or only slightly tapered stalk separates it from
Chlamydopus. It is not likely to be found by the average mushroom hunter, but any mush¬
room willing to grow in gypsum crystals warrants attention. Another robust southwestern
species, T. lysocephalum, has a coarser, dirtier appearance and its soil-incrusted “head”
topples off easily and is often found beside the stalk. It grows under mesquite and other
desert shrubs, but is not common.

Chlamydopus meyenianus (Desert Stalked Puffball)

FRUITING BODY beginning as an underground “egg” encased in an outer peridium,
the stalk then elongating rapidly; outer peridium rupturing around its periphery, the
lower half forming a volva at base of stalk, the upper portion forming a patch of warty,
brittle tissue that breaks up and disintegrates or occasionally clings to the top of the spore
722 TULOSTOMATALES

case. Mature fruiting body consisting of a spore case (inner peridium) mounted on a stalk.
SPORE CASE round to somewhat flattened, 1-3.5 cm broad and 0.5-2 cm high, tough,
persistent, attached firmly to the stalk; surface smooth or slightly roughened, pallid to
buff, pinkish-buff, yellowish, or sometimes cinnamon, developing an apical pore at
maturity which may enlarge in age to form a“mouth.” STALK4-15 (35) cm long,0.2-1.5
(3.5) cm thick at apex, tapering downward, usually solid and tough or rather woody, often
curved and/ or flattened; usually longitudinally striate or grooved, smooth to silky-
fibrillose or sometimes fibrillose-scaly; colored more or less like spore case or browner;
not percurrent. VOLVA at base of stalk saclike, two-layered, thick, often rotting away
or staying behind in the ground; white to brownish, usually incrusted with dirt or sand.
SPORE MASS rusty to ochraceous to brown and powdery at maturity. Spores 5.5-9
microns, round, spiny (or rarely smooth). Capillitium present.
HABITAT: Solitary to scattered or in small groups in sandy, gravelly, or volcanic soil,
sand dunes, gypsum flats, and other barren places; sometimes also in adobe soil. It is
found throughout the arid and semi-arid regions of the world (Australia, North Africa,
etc.) and was originally described from Peru. It is widespread in western North America,
but is common only in the Southwest—and then only sporadically. I have seen it near the
Painted Hills in eastern Oregon and outside San Bernardino in southern California. It
fruits after heavy rains but, like other stalked puffballs, persists for months afterward.
EDIBILITY: Worthless—it is much too tough and fibrous, even as a substitute for beef
jerky.
COMMENTS: Like other desert-dwelling stalked puffballs, this species is quite variable
in size and appearance but still easy to recognize. When still covered by the warty outer
peridium it is reminiscent of Phellorina strobilina, but the presence of a volva (in most
specimens) distinguishes it. Once it has shed its outer coat (as shown in photo on previous
page), it resembles the cosmopolitan genus Tulostoma, but is slightly larger, lacks the
ball-and-socket connection of spore case to stalk typical of that genus, and is characteristi¬
cally thickened at the apex of the stalk and tapered downward. The presence of a volva is
also noteworthy, but it decays more quickly than the rest of the fruiting body and is not
always evident in older specimens. The tendency of the spore case to rupture through a
single pore distinguishes it from Battarrea, which also has a volva. Two other distinctive
stalked puffballs should also be mentioned. Schizostoma laceratum is a desert dweller
whose spore case ruptures along sutures to form rays or lobes which subsequently spread
out somewhat (see photo). The spore case is attached firmly to the stalk via a ball-and-
socket arrangement (as in Tulostoma) and there is no volva, though the base of the stalk
may split to form a free rim or collar. Queletia mirabilis resembles Schizostoma, but
has a spore case that ruptures irregularly. It is widely distributed but rare—there is one
report of it from Pennsylvania.

Schizostoma laceratum (see comments above) is easily distinguished from other stalked puffballs
by the way in which its spore case ruptures into rays in old age.
Phellorina strobilina. This desert dweller releases its spores through general disintegration of the
spore case, leaving an urnlike structure behind (specimen at far right). Note presence of large warts
in the youngest individual (far left).

Phellorina strobilina (Urnlike Stalked Puffball)

FRUITING BODY consisting of a spore case which narrows downward into a distinct
stalk. SPORE CASE round to oval or pear-shaped, 1.5-6 cm broad and/ or high. Outer
layer of peridium (skin) scaly, the scales erect or pyramidal and often quite large on top,
usually smaller and more or less shingled below; white when fresh but soon becoming
brown to cinnamon and rupturing irregularly and falling away in pieces, exposing the
inner layer. Inner peridium a cup- or urnlike expansion of the stalk apex(i.e., completely
continuous with the stalk), pinkish-buff to cinnamon, with a large (2-5 cm) irregular
mouth forming at the top in age. STALK 2-5 cm long, 0.8-2 cm thick at apex, dilated at
apex and tapered downward, but with a bulb at the base; firm, solid, woody, brown or
cinnamon and scaly like the outer peridium of the spore case (but scales may wear away
in age); not percurrent. VOLVA typically absent, but the basal bulb sometimes splitting
or peeling to form a volva-like collar or rim. SPORE MASS powdery when mature and
cinnamon to rusty-brown. Spores 5-7 x 4.5-6 microns, round or nearly round, warty
or spiny. Capillitium present but often scanty.
HABITAT: Solitary or in small groups (occasionally two or three growing from a com¬
mon base) in desert soil, usually near shrubs; widely distributed in the warm, arid regions
of the world. It occurs throughout the Southwest after heavy rains but is not common and
only rarely fruits in large numbers. I have seen it outside Reno, Nevada and near Tucson,
Arizona as well as in India.
EDIBILITY: Much too woody to be edible.
COMMENTS: Also known as P. inquinans, this peculiar stalked puffball is reminiscent
of the common desert fungus Podaxis pistillaris when young, but its stalk is not per¬
current (i.e., it terminates at the base of the spore case). The scales flake off the spore case
or “cap” and the top eventually ruptures to form a large mouth so that old specimens
resemble wine goblets. These might be confused with Dictyocephalos attenuatus, but
the spore mass lacks the chambers characteristic of that species and there is normally no
volva at the base of the stalk.
723
 spores
Gastroid Agarics
PODAXALES & Allies
THIS motley assortment of bizarre and inelegant fungi combine the features of an agaric
(presence of a cap and stalk and a spore mass composed of chambers, plates, or rudi¬
mentary gills) with those of a Gasteromycete (lack of forcible spore discharge and a
frequently enclosed spore mass). Most of them look like gilled mushrooms that have
failed to open or develop properly, and that is precisely what they are most frequently
mistaken for. A few, such as Montagnea arenarius, have a fully exposed spore mass at
maturity, but they lack true gills and do not forcibly discharge their spores. Those with
an enclosed spore mass might be mistaken for stalked puffballs, but their stalk is percurrent
(i.e., it extends all the way through the spore mass to the top of the cap, just as in a gilled
mushroom). In the stalked puffballs, on the other hand, the stalk terminates at the
bottom of the spore case or “cap.”
The gastroid agarics are sometimes called “secotioid” fungi because many of them were
originally grouped in a single genus, Secotium. To facilitate identification they are retained
together here as an artificial subgroup of the Gasteromycetes. It should be realized, how¬
ever, that they are thought to have evolved independently from each other, and that
toadstool taxonomists try to express their true relationships by scattering them among the
various families of gilled mushrooms which they resemble. Most of them thrive in severe
environments (e.g., deserts or high mountains), suggesting that they are indeed
metamorphosed agarics which, through a process of adaptation and specialization, have
lost the ability or necessity to forcibly discharge their spores (hence they also go by the
unflattering moniker “reduced agarics”). Podaxispistillaris, a common desert fungus, is a
good example: it resembles the shaggy mane (Coprinus comatus), but its cap remains
closed to protect the developing spores from the desert heat, and the mature spores are
thick-walled to prevent moisture loss. Taxonomists still argue about its relationship to
Coprinus, but most of the other gastroid agarics have clearcut counterparts among the
gilled mushrooms—e.g., Longula is closely related to Agaricus, Montagnea to Coprinus,
Brauniellula to Chroogomphus, Macowanites and Arcangeliella to Russula and Lac-
tarius, and so forth. Some gastroid agarics are closely related to the false truffles. As
defined here, however, the false truffles lack a stalk and/or grow underground.
The gastroid agarics are particularly prominent and diverse in western North America.
They fruit after seasonal rains but the tougher types persist, like the stalked puffballs,
for weeks or even months before decaying. Some are said to be edible when very young,
but as a rule they are better observed or preserved than served.

Key to the Podaxales& Allies

1. Spore mass composed of tubes or empty tubular chambers .2
1. Spore mass solid, slimy, or powdery or with rudimentary gills, plates, or cavities.3
2. Fruiting body resembling a misshapen or aborted bolete; spore mass (tubes) exposed at
maturity, lining the underside of cap.(see Gastroboletus, p. 544)
2. Spore mass enclosed by a peridium (skin); not as above (see Hymenogastrales& Allies, p. 739)
3. Fruiting body averaging 3-8 cm high, narrowly to broadly club-shaped or with an oval cap;
peridium (skin) of “cap” often wearing away in age; spore mass greenish-brown to brown and
often slimy, or becoming powdery in old age; found on living or dead wood (especially pine)
and other debris in humid regions (e.g., southeastern U.S.) . Rhopalogaster transversarium
3. Fruiting body not as above, or if similar then habitat different .4
4. Fruiting body shaggy-mane-like; cap oval to cylindrical (much taller than it is wide), usually with
shaggy scales or fibrils which may, however, wear off in age; spore mass enclosed, at least until
very old age; found in deserts and other hot, arid environments .Podaxis, p. 725
4. Not as above; differently shaped (including narrowly conical), etc.5
PODAXALES & ALLIES 725

5. Fruiting body Russula- or Lactarius-like: mature spore mass or “gills” white to yellowish, ochra-
ceous, or pinkish; stalk also pale in color; fruiting body crisp, brittle orfragile, the stalk typically
snapping open cleanly like a piece of chalk; sporescolorless under the microscope, withamyloid
ornamentation; found in forests or under trees Macowanites, Arcangeliella, & Allies, p. 736
5. Spore mass brown to black at maturity, or if paler then not as above .6
6. Flesh in stalk orange to ochraceous to vinaceous when fresh and mature spore mass smoky
to gray or black; found under conifers .Brauniellula, p. 732
6. Not as above (but flesh may stain yellow). 7
7. Flesh white in upper part of stalk, bright yellow in base; spore mass minutely chambered; found
under conifers .Gomphogasler leucosarx (see Brauniellula nancyae, p. 732)
7. Not as above .8
8. Growing on hardwoods in Midwest; spore mass or“gills” white; cap usually with small brownish
scales.Lentinus tigrinus (gastroid form) (see Lentinus & Lentinellus, p. 141)
8. Growing on ground, or if on wood then not as above .9
9. Cap narrowly conical and not expanding appreciably (see photo on p. 734, top right); usually
found in grass or alpine meadows .Thaxterogaster, Nivatogastrium, & Allies, p. 733
9. Not as above . 10
10. Fruiting body Agaricus-\\ke or Coprinus-like; mature spore mass or “gills” dark brown to
black; terrestrial .Endoptychum & Allies, p. 727
10. Not as above (mature spore mass may be paler brown or grayish); found on wood or ground 11
11. Fruiting body somewhat Lepiota- or puffball-like, at least when young; spore mass often white
at first but yellowish to brown or grayish and often powdery in age; fruiting body sometimes
broadly conical but not narrowly conical; growing mainly in grass, cultivated earth, deserts,
open places, etc.; terrestrial .Endoptychum & Allies, p. Ill
11. Not as above; spore mass soon yellow-brown to cinnamon-brown or reddish-brown; cap very
narrowly conical, or if not then usually found on wood or on ground in forests and under
trees (including eucalyptus) .Thaxterogaster, Nivatogastrium, & Allies, p. 733

PODAXIS
THIS genus occurs throughout the hotter parts of the world. It displays several interesting
protective adaptations which enable it to survive in its hostile environment, e.g., the spores
are thick-walled to prevent moisture loss and the cap never opens out. The single species
described here is easily recognized by its tough, shaggy mane-like fruiting body and
powdery mature spore mass. The presence ofcapillitium(seecomments)isalsodistinctive.

Podaxispistillaris (Desert Shaggy Mane)

FRUITING BODY with a cap and stalk. CAP 2-15 cm high and 1-4 cm broad, oval to
cylindrical or sometimes somewhat twisted; surface dry, pure white to tan, yellow-brown,
or brown; typically breaking up to form shaggy fibrils or scales which may eventually peel
or wear away to reveal the smooth undersurface; eventually tearing radially or irregularly,
the margin usually remaining attached to the stalk for a long time. SPORE MASS con¬
sisting of contorted plates or rudimentary gills; at first white, soon darkening to yellowish,
then reddish, and finally reddish-brown to dark brown to blackish; powdery at maturity
(but liquefying slightly under certain conditions). STALK 4-15 (26) cm long,0.2-1 (1.5) cm
thick, equal above, the base usually swollen; extending into the spore mass and often
percurrent; usually longand slender, rather tough and fibrillose to raggedly scaly(but often
smoother in age); solid or hollow, often twisted-striate; white, or in age often discolored
like the cap. VEIL not clearly differentiated from cap and stalk tissue. VOLVA absent.
SPORE PRINT not obtainable; spores averaging (7) 10-16 (20) * (5)9-15 microns, but
sometimes much larger (to 36 microns!); round to broadly elliptical, pear-shaped, or
irregular, smooth, thick-walled, with an apical germ pore. Capillitium present.
Podaxispistillaris looks like a shaggy mane, but lacks gills and grows in the desert. Stalk is percurrent
(i.e., it extends into the spore case or “cap”). Left: Typical specimens with a shaggy or scaly cap. Right:
Weathered specimens in which the scaly layer has worn away, revealing the smooth undersurface.

HABITAT: Solitary or scattered to gregarious or clustered onground in deserts, washes,

lawns, gardens, irrigated fields, etc.; occurring throughout the warm, dry parts of the
world (known from every continent except Antarctica). It is the most prominent desert
fungus of the western United States, showing a preference for sandy soil but also coming
up in clay or adobe. In California it has been found from below sea level to over 5,000
feet (in the Panamint Mountains). It usually fruits between April and October, but the
fruiting bodies persist for months and consequently can be found most any time. I have
seen it growing outside the Taj Mahal in India, accompanied by Agaricus bitorquis.
EDIBILITY: Said to be edible when very young (white inside), but tough and dusty in age.
COMMENTS: This distinctive denizen of the desert looks like a shaggy mane (Coprinus
comatus) but does not have gills, does not normally deliquesce, and has a powdery spore
mass when mature. To the feverish, sun-fried, dust-incrusted fungophile driving deli¬
riously across the monotonous mushroom-meager desert while dreaming of cool coastal
pine forests bulging with boletes and flower-filled mountain meadows overflowing with
Agaricus, it is likely to be mistaken for a miraculous mirage or wistful hallucination.
To the methodical mushroom taxonomist, on the other hand, it is an evolutionary
anomaly: the stalk is percurrent as in other gastroid agarics, but the spore mass contains
capillitium as in the stalked puffballs. (Some taxonomists consider it to be a “gastroid”
relative of Coprinus, while others sequester it in an exclusive order of its own, the Podax-
ales.) The percurrent stalk distinguishes it from the similarly-shaped Phellorina strobilina
(a stalked puffball), and the shape and habitat separate it from other gastroid agarics and
stalked puffballs (but see Rhopalogaster transversarium in key on p. 724). It’s interesting
to note that, like the shaggy mane, it sometimes grows with Agaricus bitorquis. Perhaps
since the latter fruits underground, it has a greater temperature range than either the
shaggy mane (which likes it cool) or the desert shaggy mane (which likes it hot). Other
species: P. argentinus, with an olive-brown to yellow-brown mature spore mass, andP.
microsporus, with a reddish-brown mature spore mass, are both very similar but have
much smaller spores (5-9 microns long). P. longii also has smaller spores, but has a large
and robust fruiting body (17^45 cm tall, stalk 1-2.5 cm thick at apex). All of these occur in
the Southwest, but are not nearly as common as P. pistillaris.
 727

ENDOPTYCHUM& Allies
Medium-sized to fairly large, terrestrial fungi found mostly in arid places. FRUITING BODY with
a cap and stalk. CAP variously shaped. Flesh usually white when fresh, but often staining or dis¬
coloring. SPORE MASS typically composed of plates, irregularly contorted gills that may branch
to form cavities, etc.; enclosed or exposed, typically brown to deep brown or black at maturity
and sometimes powdery. STALK percurrent, long to very short. VEIL and/ or VOLVA typically
present when fresh, but sometimes disintegrating or easily overlooked. SPORE PRINT not ob¬
tainable. Spores yellow-brown to dark brown under the microscope, round to elliptical, smooth.
Capillitium absent.

THESE dark-spored fungi are remarkably reminiscent of Agaricus and Coprinus, but
do not forcibly discharge their spores. Five genera are treated here. The central genus,
Endoptychum, may someday be divided into two genera because some of its species (e.g.,
E. depressum) are obviously related to Agaricus, while others (e.g., E. agaricoides) may
be closer to the parasol mushrooms (Lepiota and Chlorophyllum). In Endoptychum the
spore mass is typically enclosed by the cap and becomes rather powdery in old age. A
similar genus, Longula, usually has an exposed spore mass at maturity. It also resem¬
bles Agaricus, but has congested or disfigured “gills.” Neosecotium is a small genus
which, like Endoptychum, is probably related to the Lepiotas. The other two genera,
Montagnea and Gyrophragmium, have blackish spores and very small, disclike caps with
a fully exposed spore mass. They are thought to be related to the genus Coprinus, although
the name Gyrophragmium has also been used for Agaricus-hkt forms.
Most of the above fungi fruit in hot, open, arid or semi-arid habitats (E. depressum,
however, favors mountain conifers). None are known to be poisonous, but they are usually
found after the spore mass has matured and the fruiting body has toughened.

Key to Endoptychum & Allies

1. Cap thin and disclike (less than 4 cm broad), with spore-bearing plates or “gills” hanging from
its underside or margin; mature spore mass black .2
1. Spore mass enclosed for a long time, or if exposed then not as above .3
2. Spore-bearing plates or “gills” attached to underside of cap; rare .
. Gyrophragmium californicum (see Montagnea arenarius, below)
2. Spore-bearing plates hanging from margin of cap; fairly common Montagnea arenarius, below
3. Found with mountain conifers (or aspen); mature spore mass chocolate-brown to black; young
fruiting body often with a sweet odor when broken open . Endoptychum depressum, p. 730
3. Not with above features; usually found in deserts, lawns, gardens, open areas, etc.4
4. Mature spore mass usually exposed and brown to blackish .Longula texensis, p. 729
4. Spore mass usually enclosed until very old age, white becoming yellowish to brownish (but
never chocolate-brown or blackish) . 5
5. Spore mass containing minute peridioles (spore-bearing capsules) which look like grains of
sand; stalk separating easily from cap .(see Lycoperdales & Allies, p. 677)
5. Not as above .Endoptychum agaricoides & others, p. 731

Montagnea arenarius (Gastroid Coprinus)

FRUITING BODY at first deeply buried and enclosed in a tough membrane, then expan¬
ding as the ruptured outer skin forms a volva at base of stalk. Mature fruiting body with
a stalk and disclike cap (the cap first appearing oval, but all but the disc (center) soon
splitting into spore-bearing plates. CAP 1-3.5 cm broad, merely a thin disclike expansion
of the stalk; convex becoming plane or depressed, persistent; surface smooth, white to
grayish, buff, or occasionally straw-colored, often with a volval patch or remnants; mar¬
gin often tattered or fringed in age. Flesh (in stalk) white when fresh. SPORE MASS ex¬
posed at maturity, composed of thin plates which hang from the margin of the disclike cap
Left: Montagnea arenarius, small specimens. Note how central disc or “cap” has “gills” radiating
from its edge. Right: Gyrophragmium californicum (see comments under Montagnea arenarius), a
rare mushroom whose “gills” are attached to underside of cap. Both species have a volva when fresh.

and are entirely free from the stalk; plates often wavy or curled, up to 3.5 (6) cm long,
reddish-black to blackish at maturity, eventually falling off the cap or disintegrating,
but not deliquescing. STALK (5) 8-30 cm long, 0.2-1.5 (2.5) cm thick, percurrent, more
or less equal or often tapering downward, hollow, tough or almost woody when old and
dry (but very light); smooth or longitudinally fibrillose-striate, often splitting or cracking
into fibrillose or shaggy scales, white to buff or sometimes discoloring darker in old age.
VEIL absent or rudimentary. VOLVA at base of stalk saclike, usually buried in soil,
loose (often remaining in ground), two-layered, the outer layer white and ample, the
inner layer composed of tough fibers. SPORE PRINT not obtainable; spores (7.5) 12-20
(28) * (4.5) 6-11 (14) microns, elliptical to nearly round, smooth, with a germ pore. Capil-
litium absent.
HABITAT: Solitary, scattered, or gregarious in sandy soil, old fields, and other waste
places; widely distributed and fairly common in the arid and semi-arid parts of western
North America (from Mexico and Texas to California and eastern Washington). It has
been found high up on the slopes of Mount Shasta, and by the hundreds in fields in eastern
Oregon. In California it is quite common inland. I have yet to find it in our area, but the
very similar Gyrophragmium californicum (see comments) does occur rarely in sandy soil.
EDIBILITY: Like myself, too thin and tough to be of value.
COMMENTS: This odd but interesting fungus can be told by its long stalk and thin
disclike cap from whose margin the spore-bearing plates are suspended. The presence
of a volva is also distinctive, but the volva is deeply buried and easily left behind in the
ground (or sometimes rots away). The size and stature vary considerably—those from
Oregon and north-central California being rather small and slender (usually less than 20
cm tall), and those from southern California and the Southwest being somewhat taller
(20-30 cm) and often thicker. The blackish spores and thin spore-bearing plates suggest
that Montagnea evolved from Coprinus, but the fruiting body does not deliquesce, nor
does it have true gills. A similar but much rarer fungus, Gyrophragmium calif ornicum, has
dark brown to blackish spore-bearing plates that hang from the underside of the disclike
cap rather than from the margin. It also has a loose volva plus a double-layered partial
veil that either disappears or forms an annulus (ring) on the stalk. It is known only from
the San Francisco Bay region of California, and appears to be quite rare. In both of these

728
MONTAGNEA 729

species the volva and spore mass disintegrate or rot away, while the light woody stalk
and thin disclike cap remain intact and persist for months without decaying. In this
condition they can be mistaken for old specimens of Battarrea phalloides (a stalked puff¬
ball) whose spore mass has dispersed. In the latter species, however, the disc at the top of
the stalk represents the underside of the old spore case rather than the cap.

Longula texensis (Gastroid Agaricus)

FRUITING BODY with a cap and stalk. CAP 3-9 cm broad, oval to round or broadly
convex or somewhat flattened; surface dry, white to buff becoming tan to ochre or
brownish in old age, smooth or often breaking up to form white to brown fibrillose scales
or warts that may wear away in age; becoming fragile as it dries out, often rupturing longi¬
tudinally in age (especially near margin); margin at first joined to cap by a veil. Flesh firm,
white, but bruised areas often stained yellowish (or sometimes pinkish-stained in stalk).
SPORE MASS composed of crowded, convoluted, folded or branched plates and/ or
cavities; free from the stalk (at least in age); brownish becoming deep chocolate-brown
to blackish at maturity and usually exposed or partially exposed in age. STALK 2-10 cm
long, 1.5-3.5 (5) cm thick, equal or thicker at base, smooth or longitudinally striate, firm
and solid when fresh, becoming tougher or even woody as it dries out; white or colored like
cap; percurrent. VEIL two-layered, at first continuous with the cap margin and stalk, but in
age usually pulling away from the stalk or separating from the cap to form a superior ring on
stalk. VOLVA present or absent (usually intergrown with stalk and therefore inconspi¬
cuous). SPORE PRINT not obtainable; spores 6-7.5 * 5-6.5 microns, nearly round,
smooth. Capillitium absent.
HABITAT: Solitary, scattered, or gregarious in poor soil, waste places, irrigated fields,
lawns, disturbed areas, etc.; apparently widespread in the arid and semi-arid areas of the
West (from Texas to California and north at least to Oregon), fruiting after rains or
artificial watering. It used to be common in the San Francisco Bay area, but has apparently
been pushed out by development. It can still be found with regularity in the Sacramento and
San Joaquin Valleys, and has also been found in Santa Clara and San Mateo counties.
EDIBIIT Y: U nknown. It has probably been eaten as an Agaricus button, but I can find no
specific information on it. According to Rick Kerrigan, it is easily cultivated.

Longula texensis looks like an aborted or deformed Agaricus. Note wide variation in size and shape.
At left a stout button has been sliced open to show the dark spore mass inside; at far right are two slim
buttons; at center are three small, mature, dried-up individuals (two with a fully exposed spore mass).
Longula texensis. These large specimens were found near Coalinga, California. The cap can be scaly
or smooth (see photo on previous page).

COMMENTS: Originally called Longia (or Gyrophragmium) texensis, this mushroom

is closely allied to Agaricus, and is likely to be mistaken for that genus because of the veil
(which sometimes forms an annulus or ring), cap, stalk, and dark spore mass. However, it
lacks true gills and the spores are not forcibly discharged. When the veil breaks or pulls
away from the stalk or the cap splits radially, the spore mass is exposed to the elements,
whereas in Endoptychum the spore mass usually remains enclosed. A giant version of this
species, L. texensis var. major, occurs throughout the range of the typical variety. Its cap is
6-12 or more cm broad and often quite scaly, and the stalk is 10-25 cm long and 2.5 cm
or more thick (see photograph above).

Endoptychum depressum Color Plate 84

(Mountain Gastroid Agaricus)
FRUITING BODY with a cap and stalk. CAP 3-15 cm broad, rounded or convex or
flattened on top, often centrally depressed in age; surface smooth or occasionally scaly,
white to dingy whitish or buff, often staining yellow or amber when bruised and in age
(and sometimes staining vinaceous-brown near bottom); margin depressed around the
stalk and joined to it by a veil, sometimes pulling away from stalk in age or the cap rup¬
turing irregularly. Flesh white, firm becoming tough in age, often bruising yellow; odor
usually sweet (like almond extract or anise) when young but often unpleasant or musty at
maturity. SPORE MASS composed of branched or contorted, crowded plates (rudi¬
mentary gills) or elongated chambers, pallid soon becoming chocolate-brown to blackish-
brown, usually powdery at maturity and not normally exposed except in old age. ST ALK
1 -4 cm or more long, 0.8-3 cm thick, equal or tapered, usually short (but see comments),

Endoptychum depressum, the widespread short-stalked form. Specimen at right isbeingviewed from
top, the specimen next to it from the bottom. Specimen at far left has been cut open to show the dark
mature spore mass. Specimen next to it is being viewed from side. See color plate for long-stalked form.
ENDOPTYCHUM 731

firm, white or stained like the cap, smooth, percurrent. VEIL rather tough, covering the
juncture of cap margin and stalk, not normally rupturing but sometimes breaking away
from the stalk in age. VOLVA typically absent. SPORE PRINT not obtainable; spores
variable in size but usually averaging 6-10 * 5.5-8 microns, round to broadly elliptical,
smooth, thick-walled. Capillitium absent.
HABITAT: Solitary to gregarious or clustered in duff or soil under conifers (or some¬
times aspen); common in the mountains of western North America, especially in the late
summer and fall. The short-stemmed form typical of the Sierra Nevada often develops
underground but usually pokes through the surface by maturity. A long-stemmed version
(see comments) is sometimes abundant under ponderosa pine in the Southwest.
EDIBILITY: Said to be edible (use only firm specimens); I haven’t tried it.
COMMENTS: This mushroom is closely related to the section Arvenses of the genus
Agaricus by virtue of its sweet odor when young, tendency to stain or age yellow, and
dark spores. However, the veil doesn’t normally break until after the spores are mature(if
it breaks at all) and the gills are misshapen to practically absent. The stalk is usually quite
short (as shown in the black-and-white photo), but the long-stemmed variety (which may
or may not be distinct) shown in the color plate is the prevalent one in the Southwest.
Several other species may occur in the West, but have not been formally described. The
names Secotium and Gyrophragmium are used by some authors instead of Endoptychum.

Endoptychum agaricoides (Gastroid Lepiota)

FRUITING BODY with a round to oval to pear-shaped, broadly conical, or heart-shaped
(narrowed at top) cap and a short stalk. CAP (1) 2-7 (10) cm broad, 2.5-10 (12) cm high;
surface dry, white when young and fresh, but discoloring buff to tan, ochre, or brownish
with age; fibrillose, but often breaking up to form scales which give it a shredded ap¬
pearance; margin typically joined to the stalk. Flesh (in stalk and cap) white when fresh,
but in one form staining vinaceous-brown when exposed. SPORE MASS composed of
distorted chambers and/or plates (rudimentary gills) crowded within the cap; at first
white and fleshy, then turning yellowish and finally becoming brown to yellow-brown in
old age, and sometimes powdery. STALK percurrent, sometimes entirely internal (a
vertical column that extends to the top of cap), but usually extending slightly below the cap
(up to 2 cm), usually thickest at base of cap; white when fresh, but often discoloring like
the cap in age; usually with a mycelial cord at base. VEIL not clearly differentiated from

This immature Endoptychum agaricoides has been sliced open to show the percurrent stalk and white
spore mass. Later the spores turn yellowish or brown and may become powdery. Note the Lepiota-
like scales on cap.
732 PODAXALES & ALLIES

cap (i.e., underside or “margin” of cap joined to stalk). VOLVA absent. SPORE PRINT
not obtainable; spores 6-9 (11) * 5-7 microns, elliptical, smooth, with a thick inner wall
that has an apical pore. Capillitium absent.
HABITAT: Solitary, scattered, or ingroups or clusters in lawns, fields, flower beds, waste
places, etc.; widely distributed. It is fairly common in the Southwest and Rocky Mountain
region in the summer and early fall, but I have not seen it in coastal California. It is one of
the few gastroid agarics to be found in eastern North America.

EDIBILITY: Said to be edible when young; I haven’t tried it.

COMMENTS: Also known as Secotium agaricoides, this unsung, farflung fungus looks
like an unexpanded Agaricus or Lepiota, but has only rudimentary gills and never opens
out. The rather pale spore color and thick inner spore wall with a pore suggests a closer
relationship to Lepiota or Chlorophyllum than to Agaricus. It does not appear to have
much in common with other Endoptychums(e.g., E. depressum), but is the “type” species
of the genus. Other species: E. arizonicum of the Southwest is very similar, but has much
larger spores; Neosecotium macrosporum of the Midwest, Southwest, and southern
California also has larger spores, but they are ornamented rather than smooth and the
fruiting body is usually slightly smaller(\A cm high).

BRAUNIELLULA
EASILY recognized by its orangish to vinaceous flesh and gastroid appearance, this
small genus is clearly related to the pine spikes (Chroogomphus). Like the latter, it has
grayish-black spores and amyloid flesh and grows exclusively with conifers. As there is
only one common species, a key hardly seems necessary.

Brauniellula nancyae (Gastroid Pine Spike)

FRUITING BODY with a cap and short stalk. CAP (0.5) 1-6 cm broad, rounded to
convex or lobed, or sometimes flattened on top; surface dry to very slightly viscid, fibril-
lose, ochre to pale or dull orange beneath a layer of flattened grayish to brownish-gray
fibrils, often becoming vinaceous in age; margin often lobed, at first attached to the stalk by
the veil, separating from the veil in age but remaining incurved. Flesh thick, firm, pale
orange to ochraceous when fresh. SPORE MASS composed of crowded, convoluted
plates and/ or cavities; ochraceous to pale orange, becoming darker (grayish to nearly
black) as spores mature; usually remaining covered by the veil, but sometimes partially
exposed. STALK 0.3-2 cm long, 0.5-1 cm thick, equal or narrowed below, often so short
that it scarcely protrudes below the cap; solid, firm, ochraceous to dull orange, usually
fibrillose, often becoming vinaceous or reddish-stained in age and/or at base. VEIL
fibrillose, ochraceous or becoming vinaceous. VOLVA absent. SPORE PRINT not
obtainable; spores 16-20 * 6.5-9 microns, narrowly elliptical to spindle-shaped, smooth.
HABIT AT: S olitary or scattered to densely gregarious or clustered in duff under conifers
(especially fir and pine) in the summer and early fall; known only from western North

Brauniellula nancyae (-B. albipes) is common under mountain conifers throughout the West. Note
short stalk. It is closely related to Chroogomphus. (Herb Saylor)
BRAUNIELLULA 733

America. It is common in the Sierra Nevada and Cascades as well as in Idaho; it is also
said to occur in northern Arizona.
EDIBILITY: Unknown, but probably edible.
COMMENTS: This curious mountain fungus is an almost exact replica of a Chroo-
gomphus, except that it has only rudimentary gills (if any) and does not open out, and is
often buried or half-buried in the humus layer. Note that it is the same color as Chroo-
gomphus (dull or pale orange to ochraceous, becoming vinaceous in age) and has the same
spore color (gray to black) and habitat. The grayish fibrils on the cap are suggestive of C.
leptocystis (see comments under C. tomentosus). According to Eric Gerry, B. albipes
is an earlier (more correct) name for B. nancyae. Other gastroid members of the Chroo-
gomphus-{2im\\y (Gomphidiaceae) include: an undescribed species (or perhaps just a
“freak”) found under a pine in southern California (larger than B. nancyae, and lacking
the grayish fibrils on the cap); and Gomphogaster leucosarx, described from Idaho, which
has white flesh in the upper stalk and a bright yellow stem base (like the agaric genus
Gomphidius), but which has a minutely chambered spore mass instead of gills.

THAXTEROGASTER, NIVATOGASTRIUM, & Allies

Medium-sized to small fungi found on wood (Nivatogastrium) or ground (Thaxterogaster &
others). FRUITING BODY with a cap and stalk. CAP narrowly conical to convex, plane, or cen¬
trally depressed, viscid or dry. SPORE MASS enclosed by the cap but occasionally exposed at
maturity, composed of rudimentary gills or plates that sometimes branch to form cavities, ochre
to brown, rusty-brown, or reddish-brown at maturity, not normally becoming powdery. STALK
usually well-developed (but sometimes very short), percurrent. VEIL absent or if present then
thin and fibrillose. VOLVA typically absent. SPORE PRINT not obtainable. Spores tawny to
brown under the microscope, mostly elliptical, smooth or roughened. Sphaerocystsandcapillitium
absent.

SIX miscellaneous genera of gastroid agarics are treated here. All have brown spores
and are thought to have evolved independently from various brown-spored gilled
mushrooms. Nivatogastrium, for instance, is closely related to Pholiota; like that genus,
it grows on wood. Thaxterogaster, on the other hand, is terrestrial and allied to Cor-
tinarius; Weraroa and Galeropsis, with their distinctive pointed caps, may be derived
from Psilocybe or possibly Conocybe; Setchelliogaster and Gastrocybe are both sug¬
gestive of the Bolbitiaceae. All six are small genera and none are worth eating. Three
characteristic species of our western mountains are described here.

Key to Thaxterogaster, Nivatogastrium, & Allies

1. Growing on wood (on mountain conifers) ..Nivatogastrium nubigenum, p. 735
1. Growing on ground . 2
2. Cap very narrowly conical (like a dunce cap); usually found in grass or mountains . 3
2. Not as above .4
3. Found on lawns; fruiting body soon dissolving or collapsing .
.Gastrocybe lateritia {see Weraroa cucullata, p. 734)
3. Usually found in mountains (but often among grasses); fruiting body not dissolving quickly
.Weraroa cucullata & others, p. 734
4. Associated principally (if not exclusively) with eucalyptus; stalk slender .
. Setchelliogaster tenuipesisee Weraroa cucullata, p. 734)
4. Not as above; associated mainly with conifers . 5
5. Veil thick and/ or persistent; gills present; spore print obtainable . . . (see Cortinarius, p. 417)
5. Veil fibrillose or not well-developed, persisting or disappearing; spore mass composed of
contorted plates or cavities; spore print not obtainable .Thaxterogasterpingue, p. 734
Left: Thaxterogaster pingue. Note percurrent stalk in sliced specimen on left. The stalk can be very
short or fairly long depending on environmental conditions. Right: Weraroa cucullaia is easily told
by its small size and narrow, pointed “head.”

Thaxterogaster pingue (Gastroid Cortinarius)

FRUITING BODY with a cap and short stalk. CAP 1-5 cm broad, rounded or obtuse to
convex or lobed, the top often flattened somewhat in age; surface slightly viscid to slimy
when moist, smooth, buff to olive-yellow to dingy yellow-brown or dark brown. Flesh
firm, white or tinged cap color. SPORE MASS composed of crowded, contorted plates
and/or small cavities; yellowish becoming dull brown to pale or dark cinnamon-brown.
STALK 0. 5-4.5 cm long, 0.8-2.5 cm thick, very short or rudimentary in some forms, well-
developed in others; equal or swollen at base; percurrent (the portion in the spore mass
usually quite narrow and unbranched); smooth, often viscid near base when moist, dull
yellowish to buff or colored like the cap, often purple or lilac where exposed to light. VEIL
fibrillose or cobwebby, persisting or disappearing. VOLVA absent. SPORE PRINT
not obtainable; spores (12) 14-16.5 * 8-9.5 microns, elliptical to somewhat oblong, warty
and/or wrinkled.
HABITAT: Solitary to gregarious in duff under conifers in western North America; fairly
common in the summer and fall, especially at higher elevations. It is one of the most fre¬
quently encountered gastroid agarics of the Rockies (usually under spruce). In the Sierra
Nevada and Cascades it favors fir. In our area it is rare—one specimen of local origin
was brought to me, but its collector did not note the habitat.
EDIBILITY: Unknown—but hardly tempting.
COMMENTS: This species is best recognized by its viscid cap, dingy yellowish-brown
to brown overall color, and frequently violet-tinged stalk. It is a variable species insofar
as shape is concerned—specimens growing in regions of high rainfall are apt to fruit above
the ground and have a well-developed stalk, while those growing in drier habitats are more
likely to remain buried in the duff and have a short stalk that barely protrudes from the cap
(i.e., the fruiting body can be roundish and Rhizopogon-\ike). Thaxterogaster is thought
to be closely related to the agaric genus Cortinarius. Several other species of Thaxtero¬
gaster occur in the United States, some of them undescribed.

Weraroa cucullata (Gastroid Liberty Cap)

FRUITING BODY with a cap and stalk. CAP 1^4 cm high, 0.4-1.5 cm broad, narrowly
conical with a pointed apex; surface dry to slightly viscid, smooth or fibrillose, often
wrinkled, yellow to yellowish-buff to tawny or yellow-brown; margin often splitting in

734
WERAROA 735

age and retracting slightly from the stalk. SPORE MASS composed of crisped or con¬
torted gills that may or may not branch to form cavities; brown to reddish-brown. ST ALK
long and thin (5-12 cm long, 1 -4 mm thick), equal or with a small bulb at base, hollow or
stuffed, rather tough, smooth or fibrillose, colored like cap, dry. VEIL (partial) absent
or evanescent. VOLVA absent. SPORE PRINT not obtainable; spores 11-14 (16) x 6-8
(10) microns, elliptical, smooth, with a germ pore; tawny to brown under the microscope.
HABITAT: Scattered to gregarious on ground, usually among grasses in wet alpine
meadows; widely distributed in the mountains of western North America. It occurs in the
Sierra Nevada and I have found it in the southern Rockies in August.
EDIBILITY: Utterly and irrefutably inconsequential.
COMMENTS: The narrowly conical, pointed cap perched on a long thin stalk plus the
rudimentary gills make this a most distinctive mushroom. It is reminiscent of a liberty cap
(Psilocybe semilanceata), but does not have true gills. Galeropsis cucullata is an older
name for it. Galeropsis polytrichoides, found in the same habitats, may be a smaller-
spored version of the same species. G. angusticeps is an eastern species. Gastrocybe
lateritia is also similar but grows on lawns and, like Conocybe, collapses or dissolves
very quickly (within a few hours). Also deserving mention is Setchelliogasler tenuipes,
a small species probably related to the Bolbitiaceae. It has a convex to round or cylindrical,
yellow-brown to brown or reddish-brown, non-viscid cap with a chambered or gill-like,
exposed spore mass and a short or long but slender stalk. It is associated with eucalyptus
and is either especially fond of erudite settings or is only sought for by “academia nuts.”
(It has been found on the Stanford and University of California campuses.)

Nivatogastrium nubigenum (Gastroid Pholiota)

FRUITING BODY with a cap and stalk. CAP (1)2-8 cm broad, nearly round to convex,
sometimes expanding in age, the center often flattened or slightly depressed; surface
smooth or slightly fibrillose, ochraceous or tawny to dingy yellowish, buff, or white
(usually fading to whitish in age), slightly viscid when wet, sometimes pitted; margin at
first incurved and often lobed, remaining attached to the stalk or sometimes pulling away
in old age. Flesh white in the cap and columella (that portion of the stalk surrounded by the
spore mass), usually brown in the stalk; rather soft but tough (especially in the stalk);
odor mild or fragrant (like bubble gum). SPORE MASS brown to cinnamon-brown when

Nivatogastrium nubigenum grows on dead mountain conifers in the spring. Spore mass is composed
of contorted gills and is sometimes exposed in old age (large specimen). Note pale color.
736 PODAXALES & ALLIES

mature, composed of irregularly contorted gills that often form chambers; exposed only
in old age if at all. STALK 0.5-2.5 cm long, 0.5-2 cm thick, equal or thicker at either end,
percurrent, usually short and stout, quite tough (difficult to section); white or stained
brownish to rusty-bfown, not viscid. VEIL present as whitish fibrillose tissue that extends
from the cap margin to the stalk; often disappearing in age, not forming an annulus (ring).
VOLVA absent. SPORE PRINT not obtainable; spores(3)7-9 (12) * (3)5.5-6.5 microns,
elliptical, smooth, brown in mass, honey-colored under the microscope.
HABITAT: Solitary to gregarious or in small clusters on rotting conifers, often near
melting snow or shortly after the snow disappears; fairly common in the mountains of
the West in the spring and early summer, especially on fir and lodgepole pine. I have seen
large fruitings in the Sierra Nevada and Cascades, but have yet to find it on the coast.
EDIBILITY: I can find no information on it.
COMMENTS: This distinctive member of the “snowbank” mushroom flora is easily
identified by its pale color, growth on wood, and irregularly contorted or chambered
brown “gills” which remain enclosed by the cap or are exposed only in old age. Like other
gastroid or “reduced” agarics, it is apt to be mistaken for an unopened gilled mushroom.
Its occurrence on wood and brown “gills” plus the color and shape of the spores relate it
to the agaric genus Pholiota. Another P/20/iota-relative, N. wrightii, has been found in
the mountains of southern California.

MACOWANITES, ARCANGELIELLA, & Allies

Small to medium-sized, terrestrial woodland fungi with a crisp and brittle or fragile texture.
FRUITING BODY with a cap and stalk. CAP variously shaped but never conical, dull or brightly
colored; margin frequently joined to or touching the stalk, at least when young. Flesh white or
pallid. SPORE MASS white to yellow, pinkish, or ochraceous; usually composed of contorted
gills which may branch to form cavities; enclosed or exposed. Latex present in Arcangeliella, other¬
wise absent. STALK often short, percurrent, white or pale-colored, brittle (typically snapping
open cleanly like a piece of chalk). VEIL and VOLVA absent. SPORE PRINT not obtainable.
Spores hyaline (colorless) under the microscope, round to elliptical, ornamented with amyloid
warts, spines, or ridges. Sphaerocysts usually present in some part of tissue. Capillitium absent.

THESE curious fungi look like disfigured specimens of Russula and Lactarius, which is
essentially what they are. Like those genera, they have white to ochre gills, a crisp or brittle
texture, and ornamented amyloid spores. However, as in other gastroid agarics, the
cap does not expand fully and the gills are irregularly contorted or aborted and have lost
the ability to forcibly discharge spores. Furthermore, while they are not truly hypogeous
(subterranean), they exhibit a tendency in that direction, i.e., they are frequently only
partially exposed at maturity. In other words, they are intermediate in aspect between the
Russulaceae (Russula and Lactarius) and certain genera of false truffles (Martellia,
Gymnomyces, and Zelleromyces). (The latter are microscopically similar but have gone
completely underground, lost their stalk, and typically have a chambered rather than
gilled spore mass.)
In Macowanites the fruiting body is Russula-like (i.e., the cap is sometimes brightly
colored and there is no latex). In Arcangeliella, on the other hand, a latex is usually present
as in Lactarius (the latex is best seen by slicing open a fresh fruiting body). A third genus,
Elasmomyces, closely resembles Macowanites, but has spores which are not modified for
forcible discharge (thus it is a step closer to the false truffles) and lacks sphaerocysts in the
gills. (Macowanites has sphaerocysts in the gills and its spores are modified for forcible
discharge though they are not actually discharged.)
Like Russula and Lactarius, these mushrooms are strictly mycorrhizal. They are
MACOWANITES, ARCANGELIELLA, & ALLIES 737

especially common and diverse under conifers in the mountains of western N orth America,
but several (e.g., Macowanites magnus) occur in our area. They are not often collected
for food and those with an acrid (peppery) taste or unpleasant odor should be avoided.
Over 25 species of Macowanites have been described from the U nited States. Most of them
are difficult to identify without a microscope. Arcangeliella and Elasmomyces have
fewer species, but are still not easy to identify. One Macowanites and one Arcangeliella
are described here.

Key to Macowanites, Arcangeliella, & Allies

1. Taste very acrid (peppery), but sometimes latently so . 2
1. Taste mild or distinctive but not acrid .4
2. Fruiting body exuding a latex (milk or juice) when cut open, at least when fresh and moist (but
latex sometimes very scant) . 3
2. Latex absent .9
3. Stalk usually well-developed; latex usually copious; found mainly in coastal forests .
.Arcangeliella variegata (see A. crassa, p. 738)
3. Stalk usually short, often poorly developed; latex copious or scanty; found mainly under moun¬
tain conifers (in Sierra Nevada, Cascades, etc.) .Arcangeliella crassa & others, p. 738
4. Fruiting body medium-sized to fairly large (cap 3-14 cm broad, stalk 3-7 cm long and often 3 cm
thick); cap dull-colored (cinnamon-buff to pale tan to dark brown); odor usually unpleasant
in age .Macowanites magnus {set M. americanus, below)
4. Not as above . 5
5. Odor antiseptic (like iodoform) or chlorine-like, at least at maturity .6
5. Odor mild or distinctive but not as above . 7
6. Odor chlorine-like; taste unpleasant Macowanites chlorinosmus (see M. americanus, below)
6. Odor antiseptic; taste mild .Macowanites iodiolens (see M. americanus, below)
7. Odor sweet, at least in age or when dried . 8
7. Odor mild or unpleasant but not sweet .9
8. Cap whitish .Elasmomyces odoratus {see Macowanites americanus, below)
8. Cap tan to rusty-brown . Arcangeliella camphorata (see A. crassa, p. 738)
9. Associated primarily with hardwoods (oak, etc.); cap whitish to bright red or reddish .... 10
9. Associated primarily with conifers; cap whitish to yellow, blue, olive, vinaceous, salmon, rosy,
purplish, vinaceous, or mixtures thereof . 11
10. Spore mass composed of contorted or rudimentary gills .
.Elasmomyces russuloides & others (see Macowanites americanus, below)
10. Spore mass minutely chambered, not normally exposed .... (seeMartellia & Allies, p. 742)
11. Taste acrid (peppery) .Macowanites luteolus & others (see M. americanus, below)
11. Taste mild or at least not acrid . Macowanites americanus & many others, below

Macowanites americanus (Gastroid Russula)

FRUITING BODY with a cap and stalk. CAP 1 -5 cm broad, convex or irregularly lobed,
but often expanding in age to plane or even centrally depressed; surface viscid when wet,
smooth or often breaking up into scales(especiallyatcenter); color variable: lilac, purplish,
vinaceous, olive, blue, yellow, or even whitish, or often mixtures thereof; margin not
typically joined to the stalk but often touching it when young. Flesh brittle, crisp, white;
odor and taste mild. SPORE MASS consisting of irregular or distorted, often veined or
even chambered gills that are whitish at first but soon become ochre to dull yellow-orange;
usually attached to stalk. STALK 1-3 cm long, 0.3-1.5 cm thick, more or less equal, firm,
brittle, smooth, white, percurrent. VEIL and VOLVA absent. SPORE PRINT not
obtainable; spores 8.5-13.5 * 8-12 microns, broadly elliptical to round, with amyloid warts
and ridges, modified for forcible discharge but not actually discharged.
738 PODAXALES & ALLIES

HABITAT: Solitary to gregarious or clustered in duff under conifers (sometimes buried);

fairly common and widespread in western North America, especially under spruce, fir,
Douglas-fir, and pine. It fruits mainly in the summer and fall and is said to be the most
common member of the genus. I have seen it in northern California.

EDIBILITY: Said to be edible—but be sure of your identification.

COMMENTS: This conifer-lover looks like a deformed or aborted Russula, which is
essentially what it is! The spores are not forcibly discharged and hence a spore print is
unobtainable. There are more than 20 other members of the genus in North America,
including: M. iodiolens, very similar but with a distinct iodoform (antiseptic) odor, found
under conifers; M. subolivaceus, a mild-tasting species wtih a white or olive-tinged cap
and smaller spores, found in the Sierra Nevada under conifers; M. chlorinosmus, with a
whitish to yellow cap, deep ochre spore mass, chlorine odor, and unpleasant taste, found
under conifers; M. luteolus, a dull or pale yellowish, acrid-tasting species found in coastal
California and the Pacific Northwest under conifers; M. subrosaceus, a pinkish-capped,
ochraceous-staining species; M. (-Elasmomyces) roseipes, with a pinkish to salmon or
vinaceous cap whose margin never touches the stalk, favoring spruce and fir;M. alpinus,
a white to creamy-buff montane species; and M. magnus, a large species (cap 3-14 cm
broad, stalk 3-7 cm long and up to 3 cm thick) with a viscid tan to dark reddish-brown cap
and an unpleasant odor, originally collected in Santa Clara County, California. The
closely related genus Elasmomyces has spores which are not modified for forcible dis¬
charge and seems to be more common under oak than conifers, at least in California. Its
members include: E. russuloides, with a red to reddish-brown cap (at least in age), widely
distributed; E. pilosus, which tends to grow underground in association with oak; and
E. odoratus of Washington, with a strong fragrant odor. None of these are worth eating.

Arcangeliella crassa (Gastroid Milk Cap)

FRUITING BODY with a cap and stalk. CAP 2-8 cm broad, convex when young, be¬
coming plane to shallowly depressed or irregular in age; surface pale buff to pinkish-buff,
sometimes darkening slightly (to cinnamon-buff) in old age, dry or slightly viscid, smooth;
margin typically remaining attached to the stalk, but often not quite reaching it in some
places, and in age sometimes separating slightly from it. Flesh thick, crisp, brittle, white to
pinkish-buff; odor often unpleasant in age; taste very acrid (peppery). Latex present
but often scanty (especially in dry weather), white, unchanging. SPORE MASS com¬
prised of contorted plates or rudimentary, often chambered gills; pale pinkish-buff to
ochre-buff, usually completely enclosed by the cap but occasionally exposed when old,
especially near the stalk. STALK 0.5-2.5 cm long, 0.5-1.5 (2) cm thick, usually stubby and
poorly-developed, often off-center; solid or hollow, crisp, brittle, dry, smooth, percurrent;
whitish or colored like the cap. VEIL and VOLV A absent. SPORE PRINT not obtainable;
spores 7.5-11 * 5.5-8 microns, elliptical, with strongly amyloid ridges (reticulate).

Left: Macowanites sp. (perhaps Macowanites magnus), a Russula-like fungus with deformed gills.
Right: Arcangeliella crassa has short stalk plus white latex which is not visible here. (Herb Saylor)
ARCANGELIELLA 739

HABITAT: Solitary to gregarious in duff under conifers and in mixed woods; known only
from the mountains of western North America. It is sometimes common (along with A.
tenax—see comments) in the Sierra Nevada in the spring, summer, and fall. Like many
gastroid agarics, it does not seem to occur on the coast.
EDIBILITY: Unknown, but the acrid taste is a deterrent.
COMMENTS: The genus Arcangeliella is closely related to Lactarius, and this species
looks like an aborted milk cap. The overall buff to pinkish-buff color and white acrid
latex are important field characters. A. tenax and A. lactarioides (the latter with a
gelatinous cap cuticle) have also been described from the mountains of Oregon and Cali¬
fornia; they are very similar if not the same. Other species: A. variegata is a coastal species
with a grayish-buff to olive-buff cap that is often yellowish-spotted in age plus a copious
white to almost clear, acrid latex and well-developed white stalk. It occurs in our area but
is more common northward. Still another species, A. (-Elasmomyces) camphorata of
Washington, is remarkable for its strong fragrant odor like that of candy caps (Lactarius
fragilis and relatives), especially when dry. As in candy caps, its latex is often absent, but
the odor and overall tan to rusty-brown color distinguish it. See also Zelleromyces(under
Martellia & Allies).

False Truffles
spores

HYMENOGASTRALES & Allies

THE false truffles, deprecatingly dubbed “squirrel food” by at least one jaded fungophile
I know, are easy-to-overlook, difficult-to-identify, subterranean, tuberlike fungi. They
are neither delicious (unless you’re a squirrel) nor conspicuous (unless you’re a squirrel)
nor attractive (unless you’re a squirrel), but they are interesting from an evolutionary
standpoint (unless you’re a squirrel) because many are thought to have evolved from the
Hymenomycetes (e.g., agarics and boletes) or perhaps to have given rise to them.
The fruiting body of a false truffle is typically potatolike: round to oval or knobby with
a tough or cartilaginous to rubbery or gelatinous interior. The spore mass or gleba (in¬
terior) is often composed of small chambers that give it a spongelike appearance. The
spores are borne on the walls of the chambers or sometimes within the chambers them¬
selves. The gleba usually remains intact through maturity rather than becoming powdery
and dispersing. These features, plus the fact that most false truffles are hypogeous (sub¬
terranean) or erumpent (i.e., they “erupt” through the surface of the ground at maturity)
distinguish them from the puffballs and earthballs, which have a powdery spore mass in
old age and usually fruit above the ground.
Some false truffles, such as Truncocolumella, possess an internal stalk or columella—
a branched or unbranched central column which penetrates the spore massand sometimes
protrudes slightly beneath it as a rudimentary stalk or “stump.” The columella can be
quite slender, so you should always make several sections of the fruiting body so as not to
overlook it. The columella or “stalk” is not usually percurrent as in the gastroid agarics,
and in many false truffles (including the most common genus, Rhizopogon) it is al¬
together lacking.
The false truffles can also be confused with the “true” truffles, which are not as common,
usually have a channelled or marbled or hollow interior, and lack a columella (see lengthy
footnote on p. 844). As might be expected, there are some exceptions (e.g., false truffles
such as Melanogaster and Alpova have veins in the spore mass, and some truffles have a
False truffles look something like puffballs, but usually grow underground and do not have powdery
spores at maturity. The interior is usually divided into chambers, but the chambers can be so minute
that a hand lens is required to see them. This species is Rhizopogon ochraceorubens (p. 755).

chambered interior), and it is sometimes necessary to make a microscopic determination

as to whether the spores are borne inside asci (the true truffles) or on basidia (the false
truffles).*
As defined here, the false truffles are an artificial and heterogeneous group. Several
genera are related to the agarics (e.g., Hymenogaster to the Cortinariaceae, Hydnangium
to Laccaria, Martellia to Russula, and Zelleromyces to Lactarius). A few, like Trunco-
columella, show affinities to the boletes, while the origins and relationships of others
(e.g., Rhizopogon and Alpova) are obscure.
Western North America harbors more than its share of false truffles—more, in fact,
than any other region in the world. Most if not all are mycorrhizal. They are very common,
but since they tend to grow underground, they’re unlikely to be seen unless sought for.
The best way to find them is to take a handrake and sift gently through the humus or needle
duff and upper soil layer (see chapter on truffles for more details), giving special attention
to areas where squirrels, deer, or wild pigs have been digging. You face fierce competition,
however, for those not devoured by animals are eagerly snatched up by mycologists (it
is only in the last fifty years that the false truffles have been studied closely—dozens have
yet to be named and many of the named ones are known only from a single locality).
Relatively few are left in the ground to rot in peace, which may actually be to their advan¬
tage—since their spore mass is not powdery and easily dispersed, they rely on rodents,
insects, rain, and mycologists to spread their spores for them. Many have strong odors
(e.g., like marzipan, garlic, or vanilla) that help to broadcast their presence.
The false truffles are of more interest to the evolutionary taxonomist than the most
revolutionary gastronomist. Some have an unpleasant taste or texture; a few, I am told,
are edible. The overwhelming majority, however, have not been tested (by humans), in
part because field identification can be very difficult. A microscope is often needed to
determine the genus and family as well as the species.
The false truffles are a vast group—there are over 150 species of Rhizopogon alone!
Only a few representatives are depicted here. The keys, which unavoidably resort to
microscopic characteristics, are greatly modified versions of those in How to Know the
Non-Gilled Mushrooms by Alexander Smith, Helen Smith, and Nancy Weber. Those
of you who wish to know more about the false truffles should consult the extensive keys
in that book, the technical literature on the subject (see bibliography), and/or the
squirrel representative nearest you.

Key to the Hymenogastrales & Allies

1. Spore borne inside asci* . (see Tuberales, p. 841)
1. Spores borne on basidia* .. 2
2. Columella present, branched or unbranched, percurrent (extending all the way through spore
mass) or stumplike, sometimes very thin (be sure you slice the fruiting body perpendicular to
the ground and through its center—you may want to make several thin slices so as not to over¬
look the columella) . 3
2. Columella absent (but a stalklike base of tough fibers sometimes present).7

*For some fairly reliable ways to distinguish the Tuberales (truffles) from the false truffles in the field, see the ex¬
tensive footnote at bottom of p. 844.
740
 HYMENOGASTRALES & ALLIES 741

3. Columella short and stumplike or branched, usually prominent; peridium (skin) typically
yellow to olive; spore mass composed of empty tubular chambers, not blueing whencut; spores
smooth, yellowish to pale brown under the microscope; associated mainly if not exclusively
with Douglas-fir . Truncocolumella, p. 752
3. Not as above .4
4. Outline of gills and cap present in longitudinal (perpendicular) section (see Agaricales, p. 58)
4. Not as above . 5
5. Peridium (skin) 2-5 mm thick; columella fairlv thick (sometimes rounded or cushion-shaped),
not percurrent; spore mass lacking chambers but with radiating lines or plates when young;
white becoming brown or black; firm to gooey or in old age powdery .... Radiigera, p. 760
5. Not as above .6
6. Spore mass usually (but not always) greenish to olive-gray or olive-brown, typically tough or
cartilaginous when young but often slimy in old age; columella often translucent (but some¬
times whitish), branched or unbranched; peridium (skin) usually well-developed; spores
smooth or enclosed in a wrinkled outer coat .Hysterangium & Allies, p. 762
6. Not as above . 11
7. Spore mass white even in age, the chambers usually gel-filled or exuding a white latex when cut;
spores with a gelatinous sheath, not amyloid .Leucophleps& Leucogaster, p. 759
7. Mature spore mass darker (may be white when young), or if white then not as above .8
8. Peridium (skin) thick (occasionally 1 mm, usually 2-5) and tough or leathery; spore mass at first
white but soon black, purplish, or dark brown, remaining firm for a long time but finally pow¬
dery, sometimes with veins but lacking gel-filled chambers .(sqq Scleroderma, p. 707)
8. Not as above; peridium usually thinner; spore mass only rarely powdery .9
9. Spore mass typically marbled with paler veins, the chambers usually gelatinous or gel-filled;
spores smooth .Alpova& Melanogaster, p. 756
9. Not as above (but may have some of above features) . 10
10. Spores smooth (not ornamented), elliptical to oblong, sometimes amyloid but not dextrinoid,
spore mass composed of minute chambers; outer surface often (but not always) with mycelial
threads; very common, especially under conifers. Rhizopogon, p. 753
10. Not as above; spores typically ornamented with lines, warts, wrinkles, spines, etc., or showing
dextrinoid “stripes” . 11
11. Spores round, elliptical, or elongated, colorless to yellowish or brown under microscope, not
amyloid; spore mass variously colored (sometimes dark); peridium absent or present ... 12
11. Spores round to elliptical with amyloid or partially amyloid warts, rods, ridges, or spines;
spores usually colorless or nearly so under the microscope; spore mass variously colored but
often white at first and not typically dark brown to blackish; peridium usually present . . 16
12. Spores round to broadly elliptical and ornamented with large warts, cones, or spines; spores
usually colorless under the microscope but sometimes brown; found in many habitats but
especially common under eucalyptus . HydnangiumSc Allies, p. 744
12. Spores nearly round to elliptical, spindle-shaped, or elongated and typically wrinkled, warted, or
longitudinally lined (rarely smooth or honeycombed), yellow-brown to rusty-brown or brown
under the microscope; common in many habitats (including eucalyptus) .13
13. Spores appearing smooth except in iodine solution(and then showing prominent brown bands
or stripes, i.e., dextrinoid); spore mass white to yellow or yellowish . 14
13. Not as above; spores typically ornamented (rarely smooth, but if so then not as above) . . 15
14. Peridium white to brownish; found in Sierra Nevada .Protogautieria substriata
14. Peridium absent or practically so; known from eastern Washington . .. Protogautieria lutea
15. Peridium (skin) often thin and soon wearing away; spore mass quite tough and crisp when
young; spores ornamented with longitudinal lines .Gautieria, p. 746
15. Not as above; peridium usually well-developed, persistent . . . Hymenogaster& Allies, p. 748
16. Outer layer of each spore consisting of amyloid rods imbedded in non-amyloid material;
sphaerocysts absent; fruiting body \-4 cm broad, usually irregular in shape, white becoming
yellowish to olive-yellow or olive-gray; spore mass white and very dry, becoming olive in age;
found under conifers, not common but sometimes fruiting prolifically Mycolevis siccigleba
16. Not as above; spores ornamented with free spines, warts, and/ or ridges; sphaerocysts usually
(but not always) present somewhere in fruiting body; mature spore mass variously colored but
not often olive; widespread and fairly common .Martellia& Allies, p. 742
742 HYMENOGASTRALES & ALLIES

MARTELLIA & Allies

Small to medium-sized woodland fungi usually found underground. FRUITING BODY round
to knobby or lobed (potato-like), variable in color. SPORE MASS (interior) typically composed of
large to small or minute chambers; usually rather crisp when fresh, white to pinkish, ochre, ochre-
brown, or cinnamon-brown. Latex often present in Zelleromyces, otherwise absent. STALK
absent except in a few species; columella present or absent. SPORES round to elliptical, orna¬
mented with amyloid warts, spines, and/or ridges; hyaline (colorless) under the microscope.
Basidia forming a hymemum that lines the walls of the chambers. Sphaerocysts usually present
somewhere in the fruiting body (but sometimes absent in Martellia). Capillitium absent.

THESE are poorly known, difficult-to-identify false truffles with a white to yellow-
or cinnamon-brown spore mass and amyloid spores. Like the gastroid agarics Maco-
wanites, Arcangeliella, and Elasmomyces* they are thought to be closely related to
Russula and Lactarius, but are one step farther removed (i.e., they lack the cap and stalk
of those genera, grow underground, and typically have a chambered rather than plated or
gilled spore mass).
The most common genus, Martellia, lacks a latex and is thought to be related to Russula.
A second genus, Gymnomyces, mimics Martellia but differs microscopically (the central
strata of tissue in the walls between the chambers contain sphaerocysts; in Martellia they
do not). A third genus, Zelleromyces, is probably closer to Lactarius than to Russula
because the fresh fruiting body often exudes a latex (juice) when cut. (A Zelleromyces
recently discovered in Yosemite National Park has a scanty dark red latex and stains
greenish—just like Lactarius rubrilacteus\)
Unless the spores are examined, all three genera are easily confused with other false
truffles such as Hymenogaster and Octavianina. However, theyareoftencrisperorfleshier
(or not as tough) as many false truffles and often paler in color, at least when young.
Martellia and Gymnomyces, and to a lesser extent, Zelleromyces, are fairly common
in our area under oak, madrone, and various conifers. However, they do not normally fruit
in the large numbers characteristic of some of the other false truffles (e.g., Rhizopogon
and Gautieria). All three genera need critical study, especially Martellia, which is fairly
sizable. As the various species are differentiated almost exclusively on microscopic charac¬
teristics, only two are described here.
Key to Martellia & Allies
1. Fruiting body with a scanty dark red latex; wounded tissue usually staining greenish (within
several hours); columella present but stalk absent; spore mass chambered; peridium (skin)
practically absent; known only from Yosemite National Park.
.Zelleromyces sp. (name soon to be published by Harry Thiers and Herb Saylor)
1. Not as above . 2
2. Columella and/ or stalk present, usually percurrent (extending clear through spore mass) . 3
2. Stalk absent; columella absent or rudimentary . 5
3. Fruiting body exuding a latex when cut; columella thin, branching; found mainly under oak
along the west coast .Zelleromyces gardneri
3. Latex absent .4
4. Spore mass rosy to pinkish to pinkish-buff .Gymnomyces socialis, p. 743
4. Mature spore mass more or less cinnamon-buff Martellia cremea (see M. brunnescens, p. 743)
5. Fruiting body exuding a white latex when cut; exterior cinnamon to reddish; spore mass (interior
of fruiting body) pale cinnamon-buff; widespread in eastern North America, especially under
pine . Zelleromyces cinnabarinus
5. Not as above; latex absent . 6
6. Exterior tinged or spotted rose or red . Gymnomyces roseomaculatus (see G. socialis, p. 743)
6. Not as above . 7
* Macowaniles, Arcangeliella, Elasmomyces, Zelleromyces, Gymnomyces, and Martellia constitute the so-called
“astrogastraceous series.” In modern classifications they are usually placed alongside Russula and Lactarius
in the Russuiales and/ or Russulaceae.
Martellia fallax (see comments under M. brunnescens). Specimen at upper right is immature, as
evidenced by white interior. Specimen at upper left is mature and distinctly fragrant. Since many
species share these features, a microscope is usually needed to distinguish them.

7. Walls of the spore-bearing chambers containing sphaerocysts in their central layers (a micro¬
scopic feature); uncommon . Gymnomycesferruginascens& others (see G. socialis, below)
7. Walls of the spore-bearing chambers lacking sphaerocysts in their central layers (sphaero¬
cysts may or may not be present elsewhere) . . Martellia brunnescens & many others, below

Martellia brunnescens
FRUITING BODY 1 -3 cm broad, round to oval or irregularly knobby (potato-like).
Outer surface white when young, developing ochre- to rusty-brown or brown stains in age
or after handling, and usually entirely brownish in old age; smooth or pitted. SPORE
MASS (interior) composed of small chambers, crisp and white when young, but dis¬
coloring brown to rusty-brown when cut (often slowly) and becoming ochre-brown to
brown at maturity; odor often sweet (somewhat reminiscent of vanilla), at least in age.
STALK and columella absent. SPORES 8-11 * 8-9 microns, broadly elliptical to round,
with strongly amyloid warts and spines; hyaline (colorless) under the microscope. Cystidia
absent. Walls of tissue between the chambers lacking sphaerocysts in their central layers.
HABITAT: Solitary, scattered, or in groups in soil or humus (buried) in woods; known
only from western North America. It favors conifers and is one of the most common
Martellias of Oregon and California. I have found it in mixed woods and under Douglas-
fir in April, July, and October.
EDIBILITY: I can find no information on it.
COMMENTS: There are many Martellias that will more or less fit the macroscopic
features of the above description. The sweet odor which develops in age might seem to be
a distinctive feature, but is found in a number of other species (e.g., M.fragrans of Idaho,
which has a strong vanilla-like odor). Another common western species,M. fallax, smells
like Russula fragrantissima (at least to me). It is quite common in our area under hard¬
woods, and differs microscopically from M. brunnescens in having cystidia which are
golden in potassium hydroxide (KOH). Also worth mentioning are M. foetens, an Idaho
species that smells like M. fallax;M. cremea, which has a percurrent columella; andM. cal¬
if ornica, M. boozeri, M. parksii, M. ellipsospora, et al, which differ microscopically.

Gymnomyces socialis
FRUITING BODY 1 -2 cm broad (but several sometimes tightly clustered to form larger
masses), roundish to somewhat irregular. Outer surface pallid to creamy or pinkish,
usually staining ochraceous to rusty-brown when handled or dried; more or less smooth.

743
744 HYMENOGASTRALES & ALLIES

SPORE MASS (interior) composed of minute chambers, rosy-pinkish to pinkish-buff at

maturity. STALK present as a short sterile base (at least in larger specimens) which is
tucked into the depression formed by the lower portion of the peridium(skin), giving rise
to a columella which penetrates the spore mass and is usually percurrent; whitish to
pinkish-buff. SPORES 9-14 * 8-11 microns, round or nearly round, ornamented with
amyloid warts or spines, hyaline (colorless) under the microscope. Walls of tissue between
the chambers containing sphaerocysts in their central layers.
HABITAT: Solitary to gregarious or clustered in soil or humus under oak; known only
from California. According to Herb Saylor, it is fairly common in our area in the late fall,
winter, and spring.
EDIBILITY: Unknown.
COMMENTS: This species can be told by its small size, pinkish-tinged peridium (skin)
and spore mass, and the presence of a columella which usually extends below the spore
mass as a short stalk. The latter feature is unusual for a Gymnomyces, and serves to dis¬
tinguish it from G. roseomaculatus, which also develops pinkish or reddish spots or
tinges and grows under oak. Specimens with a well-developed stalk might be mistaken
for a Macoxvanites or Elasmomyces, but do not have the rudimentary gills and exposed
spore mass of those genera. Other species: G. cinnamomeus and G. ferruginascens both
lack a columella and have a brownish to rusty-ochre or cinnamon exterior, at least in age.
The first occurs in California under oak; the latter is recorded from Idaho under conifers.

HYDNANGIUM & Allies

Small to medium-sized fungi found mostly underground. FRUITING BODY round to irregularly
lobed (potato-like), variously colored. SPORE MASS (interior) typically composed of chambers
or elongated cavities; pinkish in Hydnangium, yellow-orange at maturity in Sclerogaster, variously
(but usually differently) colored in Octavianina. Latex present orabsent. STALK absent orpresent
as a short sterile base; columella present or absent. SPORES usually round or nearly round and
warted or spiny but not amyloid; hyaline (colorless) or pale under the microscope (or sometimes
brown in Octavianina). Capillitium absent.

THREE miscellaneous genera with spiny or warted, non-amyloid spores are treated here.
The most common genus, Hydnangium, is easily told by its pinkish to flesh-colored spore
mass and frequent association with eucalyptus. Sclerogaster also has a distinctively
colored spore mass: bright golden-yellow to orange-yellow at maturity. It favors conifers.
Octavianina, on the other hand, is variable in color and cannot reliably be distinguished
from other false truffles without examining its spores. Its spore mass, however, is not
normally pinkish or orange-yellow.
Hydnangium is thought to be closely related to the agaric genus Laccaria. The affinities
of Sclerogaster and Octavianina are unclear.

Key to Hydnangium & Allies

1. Spore mass pinkish to flesh-colored; peridium (skin) pinkish or whitish; usually (but not
always) associated with eucalyptus .Hydnangium carneum, p. 745
1. Not as above .2
2. Spore mass typically golden-yellow to orange-yellow when mature (may be whitish at first) 3
2. Not as above . Octavianina asterosperma group & others, p. 745
3. Fruiting body 0.5-2.5 cm broad, white to pale yellow; spore mass sometimes powdery in age;
odor often strong at maturity but not like peanut butter; spores 8-11 microns broad; not un¬
common in Southwest and Rocky Mtns., especially under pine . . Sclerogaster xerophilum
3. Not as above; odor resembling peanut butter when mature; known from southern California
.Sclerogaster columnatus
Hydnangium carneum, a pinkish false truffle that favors eucalyptus. Left: Specimens with a well-
developed sterile base or “stalk.” Right: A sectioned specimen lacking a sterile base. (Herb Saylor)

Hydnangium carneum (Rosy Eucalyptus False Truffle)

FRUITING BODY 0. 5-4 cm broad, round to irregularly lobed and/or flattened. Outer
surface often pitted or wrinkled, dry, fibrillose or minutely velvety, pale rose to pinkish,
pinkish-cinnamon, pinkish-buff, or flesh-colored (but often whitish when young and/ or
retaining white areas in age), sometimes also with darker spots. SPORE MASS (interior)
composed of small, often elongated or mazelike chambers that give it a spongy or marbled
appearance; pinkish or rosy (like exterior or darker); odor usually mild. STALK absent
or often present as a fragile whitish to pinkish sterile base which may extend into the spore
mass as a thin columella. SPORES 10-18 microns, round or nearly so, with large non¬
amyloid warts or spines; hyaline (colorless) or tinged yellow under the microscope.
HABITAT: Solitary to gregarious in soil or humus under eucalyptus and possibly other
exotic trees and shrubs (e.g., Leptospermum); widely distributed wherever eucalyptus
is planted or naturalized. In California it fruits from late fall through the spring, but is par¬
ticularly common from January through April. I often find it with Hymenogaster albus
and Hysterangium fuscum. HarryThiers reports finding a large-spored variant underpine.

EDIBILITY: Edible according to one source; I haven’t tried it.

COMMENTS: The pinkish color (at least of the spore mass) and round spiny spores of this
false truffle are strongly suggestive of the agaric genus Laccaria, and it is very likely that
the two are closely related. It is interesting to note that pinkish Laccarias are sometimes
common in our area under acacia, eucalyptus, and other Australian imports. The common
Hydnangium of California has long gone under the names H. roseum and H. soeder-
stroemii, but they are probably synonyms for the wide-ranging and variable H. carneum.
One source of confusion is the presence or absence of a sterile base or “stalk.” Though
often present, it is very easily broken and consequently apt to be overlooked.

Octavianina asterosperma group

FRUITING BODY 1 -3 cm broad, round or more often knobby and pitted (potato-like),
firm, often with white mycelial threads (rhizomorphs) at the base. Outer surface whitish
at first from a thin layer of cottony tissue which wears off in patches, revealing the dingy
brown to grayish to yellowish-olive undersurface; discoloring pale greenish to faintly
bluish or grayish-olive where bruised or rubbed. Peridium (skin) often wavy, whitish to
buff, discoloring like the surface when cut. SPORE MASS (interior) composed of minute
chambers, very firm, dark brown to blackish when mature, with whitish plates of sterile
tissue sometimes visible; latex (fluid) present or absent (see comments). STALK absent,
but a slight sterile base often present as a thickening of the basal peridium, and in some
specimens this sterile base apparently giving rise to a thin columella. SPORES 13-18
microns, round, covered with large, blunt spines; yellowish to brown under the
microscope, not amyloid.
745
746 HYMENOGASTRALES & ALLIES

HABITAT: Solitary to gregarious in soil or humus under trees in woods (usually buried);
widely distributed but rare. Specimens tentatively identified as this species have been
found near Santa Cruz, California, under redwood and Douglas-fir in June and July.
EDIBILITY: Unknown.
COMMENTS: Most species of Octavianina are rare, and this one is no exception. It was
originally collected in Europe and described as exuding a latex when cut. Local material
seems nearly identical to the European version, but shows no sign of a latex. Var. potteri,
described from Michigan, also lacks a latex, but has a paler spore mass and blackens when
bruised. The large, relatively few, non-amyloid warts or spines on the spores are the out¬
standing feature of the species. Hymenogaster utriculatus (see comments under H. sub-
lilacinus) looks quite similar in the field, but has ridged or honeycombed spores. There
are several Octavianinas with a paler (pallid to cinnamon-brown) spore mass, including:
O. papyracea, with an abundant cream-colored latex and a papery peridium when dry,
originally found in northern California under redwood; O. rogersii, with smaller warts
on its spores and no latex; and O. macrospora, with elliptical spores. All of these are rare.

GAUTIERIA
Small to medium-large woodland fungi usually found underground. FRUITING BODY roundish
to oval to lobed or knobby (potato-like). Peridium (outer skin) absent or poorly-developed in many
species. SPORE MASS (interior) composed of small to large cavities or chambers, usually ochre-
yellow to yellow-brown, cinnnamon-brown, or dull brown; firm and tough or rubbery when
fresh, often rubbery-gelatinous and strong-smelling in age. STALK absent or rudimentary;
columella often present, well-developed, and branched. SPORES typically elliptical and yellowish
to rusty-cinnamon under the microscope, longitudinally striate or ridged (or even “winged”), not
amyloid. Basidia borne in a hymenium lining the walls of the chambers. Capillitium absent.

THE ochre to cinnamon to dull brown overall color and tendency of the outer skin to wear
away (thereby exposing the inner cavities) are the main field characters of this distinctive
genus of false truffles. Those few species with a persistent peridium (skin) can be distin¬
guished from Hymenogaster and other false truffles by their brownish, longitudinally
lined spores. Gautieria is a common genus (if you’re looking for truffles), but the various
species are difficult to distinguish from each other and several are still undescribed. One
representative is described here.

Key to Gautieria
1. Peridium (skin) well-developed and persistent, i.e., the spore mass not exposed .2
1. Peridium absent or present only as a thin layer of tissue that soon wears away, i.e., the spore
mass (spore-bearing chambers or cavities) usually exposed at maturity . 3
2. Peridium whitish when young .G. gautierioides (see G. monticola, p. 747)
2. Not as above . G. parksiana (see G. monticola, p. 747)
3. Spore-bearing cavities quite large (2-10 mm broad); columella typically absent or rudimentary;
found under hardwoods .G. morchelliformis (see G. monticola, p. 747)
3. Not as above; spore-bearing cavities often smaller and columella usually present; very common
under conifers, but also occurring with hardwoods . 4
4. Spore mass yellowish to bright yellow-brown or ochre; spores appearing “winged” under the
microscope .G. pterosperma(see G. monticola, p. 747)
4. Spore mass typically rusty-brown or darker or duller; spores longitudinally lined but not
“winged”..G. monticola & others, p. 747
Gautieria monticola. Left: Exterior; note how there is practically no skin, i.e., the chambered spore
mass is visible. R ight: A specimen sliced open to show the interior. N ote the thin branching columella.

Gautieria monticola
FRUITING BODY (1)2-9 cm broad, round to elongated or irregularly knobby (potato¬
like), often with a mycelial cord (rhizomorph) at base. Outer surface pallid when very
young, soon becoming light brown to ochre to rusty-brown or dull brown. Peridium
(skin) very thin, soon wearing away to expose the spore-bearing cavities. SPORE MASS
(interior) composed of numerous small (0.5-3 mm) round or elongated chambers, firm and
rubbery (not fragile) and crisp when fresh, ochre-brown to rusty-brown to dull cinnamon
to dull brown at maturity; odor often strong in age (sometimes reminiscent of decaying
onions). STALK absent, but a distinct columella usually present and often penetrating
to the center of the spore mass; columella whitish or translucent, thin, often branched.
SPORES 10-15 * 6-9 microns, elliptical, longitudinally grooved, rusty-brown to ochre-
yellow under the microscope.
HABITAT: Solitary, scattered, or in groups or clusters in duff under conifers (some¬
times partially exposed at maturity); common in western North America, particularly
under mountain conifers in the late spring, summer, and early fall. I have seen large
numbers in the Sierra Nevada and Cascades in June, and a similar species occurs in our
area under hardwoods. The strong odor that develops in age presumably aids animals
in locating them. The animals then help to disperse the spores.
EDIBILITY: Edible, I am told, but rather rubbery.
COMMENTS: The rubbery brown spore mass composed of numerous irregularly-
shaped cavities plus the absence or near-absence of an enveloping peridium (skin) are
characteristic of this species and its close relatives. Hysterangium species are somewhat
similar, but have a persistent skin and are differently colored. Similar species include; G.
graveolens, a larger-spored conifer-lover; G. pterosperma, fairly common in Oregon and
California, with “winged” spores and yellow-brown to ochre cavities; and G. Candida, with
melon-like spores. Other species: G. morchelliformis is a widespread hardwood-loving
species with very large (up to 10 mm broad) spore-bearing cavities, no peridium, and
little or no columella. I have sniffed it out under oak because, like G. monticola, itdevelops
a strong odor in age. Two small Gautierias with a persistent peridium should also be
mentioned; G. parksiana, whose peridium is yellowish-orange to ochre at first; and G.
gautierioides, whose peridium is whitish, at least initially. Both of these species occur on
the west coast and are likely to be mistaken for a Hymenogaster or Martellia until their
spores are examined.
747
748 HYMENOGASTRALES & ALLIES

HYMENOGASTER & Allies

Small to medium-sized woodland fungi usually found underground. FRUITING BODY roundish
to pear- or cushion-shaped to knobby (potato-like), variously colored. Peridium (skin) present,
usually well-developed and persistent. SPORE MASS (interior) composed of numerous irregular
chambers or tubelike cavities or at times nearly solid (i.e., the chambers very minute); typically
some shade of brown (rusty- to blackish-brown) when mature; fragile to firm, rubbery, or slightly
gelatinous. STALK absent or rudimentary, butawell-developedcolumellaoftenpresent. SPORES
tawny to brown under the microscope, elliptical to nod ulose, usually waned or wrinkled, sometimes
longitudinally ridged (Chamonixia) or in one species coarsely netted; not amyloid. Basidia borne
in a hymenium (except in Destuntzia); hymenium lining the chamber walls. Capillitium absent.

THESE are small to medium-sized false truffles with an ochre to brown or blackish
mature spore mass, a persistent peridium (skin), and warted or wrinkled, non-amyloid
spores. Hymenogaster is a diverse genus that is gradually being split into smaller units
such as Destuntzia * A smaH“satellite” genus, Chamonixia, is also treated here; it inter¬
grades with Hymenogaster. All of these are thought to be related to the Cortinariaceae
or possibly the Boletaceae. None are significant from an edibility standpoint unless you’re
a squirrel or aspire to be one. Three species are described; several others are keyed out.
Key to Hymenogaster & Allies
l. Peridium (skin) white when fresh but staining vinaceous to vinaceous-brown or blackish after
handling (sometimes also with blue-green to yellow-green stains); columella and/or short
stalk usually present, often percurrent; mature spore mass dark brown to blackish; spores
longitudinally ridged; known from California under oak.Chamonixia ambigua, p. 751
1. Not with above features . 2
2. Peridium typically whitish when young but becoming pale lilac or dull bluish, then yellowish to
orangish and finally brownish in age; spores warted, not beaked; common under mountain
and northern conifers (e.g., fir, spruce, lodgepole pine) . H. sublilacinus, p. 749
2. Not with above features . 3
3. Fruiting body staining blue or bluish-green when bruised or cut in half .4
3. Not as above . 6
4. Peridium yellowish, often with reddish stains or becoming reddish to reddish-brown in age;
spores smooth .(see Truncocolumella rubra under T. citrina, p. 752)
4. Not as above; spores longitudinally lined or ridged . 5
5. Columella short and thick, occupying only the base of the fruiting body .
.Chamonixia brevicolumna (see C. ambigua, p. 751)
5. Not as abvoe .Chamonixia caespitosa & others (see C. ambigua, p. 751)
6. Peridium usually becoming pink, reddish, or vinaceous in age or when handled; chambers of
spore mass often (not always!) gel-filled .Destuntzia rubra & others, p. 750
6. Not as above . 7
7. Spore mass (interior) dark brown at maturity (see photo on p. 749); fruiting body often 1.5 cm
or more broad; mature spores somewhat honeycombed (i.e., with ridges and “pits”); found
mainly under conifers, especially Douglas-fir; rare H. utriculatus (see H. sublilacinus, p.749)
7. Not as above; common . 8
8. Spores longitudinally ridged, grooved, or “striped” .9
8. Spores wrinkled or warted, but not as above . 10
9. Spores 9-15 microns long, with an apical pore; peridium light brown to dark yellow-brown or
maroon-brown .Chamonixia caudata (see C. ambigua, p. 751)
9. Not as above; spores usually larger .(see Gautieria, p. 746)
10. Associated with eucalyptus . 11
10. Not as above; found with oak and other trees H. parksii& others (see H. sublilacinus, p. 749)
11. Peridium and spore mass containing dirt; spore mass dark brown and gelatinous at maturity;
rare . Chondrogaster
11. Not as above; peridium white to yellowish; common . . H. albus(see H. sublilacinus, p. 749)

*The genus Destuntzia and its species (see comments under D. rubra) are described in an article by Robert Fogel
and James Trappe, now in press. The names are used here with their permission.
HYMENOGASTER 749

Hymenogaster sublilacinus
FRUITING BODY 1-3 (5.5) cm broad, round to irregularly lobed (potato-like). Outer
surface often fibrillose, variable in color but typically whitish to lilac or dull bluish when
young, becoming yellowish and then orangish and finally brown in age; often staining din¬
gy ochraceous when handled (if older) or bluish (if young). Peridium (skin) often sepa¬
rating rather easily from spore mass. SPORE MASS (interior) composed of small or
minute chambers that give it a sponge-like appearance; pallid when very young but brown
to cinnamon-brown at maturity; odor variable (mild, sweetish, farinaceous, etc.). ST ALK
absent, but sterile tissue usually present as an expansion or thickening of the basal peri¬
dium, which may or may not give rise to a thin, short, branching columella. SPORES
8-13 (15) * 5-8 (9.5) microns, elliptical, warted-wrinkled, brown under the microscope.
HABITAT: Solitary to gregarious in soil or duff under mountain conifers, especially fir,
spruce, and lodgepole or ponderosa pine; fairly common in the Sierra Nevada and other
mountain ranges of the West in the spring, summer, and early fall.
EDIBILITY: Unknown.
COMMENTS: This species appears to be the most common conifer-loving Hymeno¬
gaster, at least in California. The confusing color changes it undergoes as it ages and dries
out has led to the naming of several “new” species (e.g., H. brunnescens, H. diabolus,
H. subochraceus) where only one exists. The brown color of the mature spore mass plus
the warted brown spores separate it from other genera of false truffles. Other species:
H. parksii is very common in our area under oak and other trees. It is small (5-15 mm
broad) and whitish when fresh and has a grayish to ochre-brown or cinnamon-brown
spore mass and prominently beaked spores (see photo at bottom of p. 750).//. gilkeyae
closely resembles H. parksii but differs microcopically. H. mcmurphyi also grows under
oak, but yellows in age or when bruised. H. albus (-H. albellus, H. luteus) is a white
or yellow-staining species that grows under eucalyptus (often with Hydnangium
carneum). H. utriculatus (see photo below) is a conifer-lover with distinctive pitted-
reticulate spores at maturity. Macroscopically it rather resembles the description of
Octavianina asterosperma (dark spore mass, etc.). I have found it in large numbers under
Douglas-fir and redwood in June and July, but it probably fruits throughout the mush¬
room season. Because of its distinctive spores it may deserve a genus of its own. Many
other Hymenogasters occur, but most can only be identified with the aid of a microscope.

Hymenogaster utriculatus has very unusual spores (see comments under H. sublilacinus). Note the
dark spore mass (interior).
Destuntzia rubra is best told by its tendency to develop reddish or pinkish tones as it ages oris handled,
plus its dark, sometimes gelatinous spore mass (interior).

Destuntzia rubra
FRUITING BODY 1-4 cm broad, roundish to oblong or lobed (potato-like). Outer
surface white when young, but staining pink, reddish, or vinaceous (or even bluish) in age
or after handling. Peridium (skin) fairly thick, white when sectioned. SPORE MASS
(interior) olive-brown to grayish-brown becoming darker (sometimes nearly black) in age;
often speckled with white sterile tissue that separates clusters of minute chambers (use
hand lens!); firm or gelatinous (chambers may or may not be gel-filled). STALK and
columella absent, but vinaceous mycelium sometimes attached to base or visible in
humus. SPORES 8-12 * 6.5-9 microns, elliptical, with striate conical warts, brownish
under the microscope. Basidia borne in chambers but not typically forming a hymenium.
HABITAT: Solitary to gregarious in soil or duff (usually buried) in mixed woods and
under conifers; fairly common in our area in the late winter, spring, and early summer (but
probably more widely distributed). It seems to favor Douglas-fir. A similar species,
D. saylorii, is fairly common in the Sierra Nevada.
EDIBILITY: I can find no information on it.
COMMENTS: Formerly known as Hymenogaster ruber, this species and its relatives
can be recognized by their dark, often somewhat gelatinous spore mass and pink- or
reddish-staining exterior. The specimen shown in the photograph was pure white when
collected and entirely vinaceous when I brought it home. Several Rhizopogons stain
reddish, but usually have a more fibrillose exterior, paler interior, and/or smooth spores.
The spore mass of Destuntzia may gelatinize somewhat, leading to confusion with Melano-
gaster, but the chambers are much smaller than in that genus. Other Destuntzias include:
D. saylorii (with 1-spored basidia like D. rubra) and D.fusca of California;/), subborealis
of Idaho; and D. solstitialis, an eastern species (see footnote on p. 748).

Hymenogaster parksii (see comments under H. sublilacinus on previous page). This small whitish
false truffle is very common in our area, especially under oak. The distinctly chambered spore mass
is grayish to brownish and usually lacks a well-developed columella. There are a number of closely
related look-alikes, including H. albus, which commonly grows under eucalyptus.
Chamonixia ambigua is a rare oak-loving species with a dark spore mass and whitish exterior that
darkens when handled. Note presence of columella in sectioned specimen at center.

Chamonixia ambigua
FRUITING BODY (0.5) 1-5 cm broad, more or less pear-shaped to cushion-shaped (i.e.,
round or oval with a narrowed base) to somewhat lobed (especially if growing in clusters).
Outer surface white when fresh, but discoloring vinaceous to dark vinaceous-brown to
blackish-brown after handling or standing, and either staining yellow-green to bluish-
green when bruised or showing such stains, especially near the base; smooth but rather
soft or cottony. Peridium(skin) very thin. SPORE MASS dark brown to vinaceous-brown
to deep grayish-brown when fresh, minutely chambered, the chambers empty so that they
resemble small tubes. ST ALK absent or present as a white or greenish-stained sterile base
which usually gives rise to a columella; columella branched or unbranched, usually
percurrent. SPORES (9) 11-15 (18) x 8-12 microns, elliptical with a prominent “beak”
at one end, longitudinally ridged and slightly warted, the inner wall with an apical pore;
brown to dark brown under the microscope.

HABITAT: Solitary, scattered, or in groups or small clusters in humus and soil (buried)
under live oak; known only from California. Like many subterranean fungi, it is described
as being “rare,” but may very well be common once you know when and where to look for
it. I have found it in June and July in Santa Cruz County.
EDIBILITY: Unknown.
COMMENTS: The genus Chamonixia intergrades with Hymenogaster, but most of its
species can be told in the field by their tendency to stain bluish or to exhibit natural bluish-
green stains. The distinctly tubulose (spongelike) spore mass is reminiscent of a bolete,
and it has been suggested that Chamonixia is related to the boletes (along with Trunco-
columella). C. ambigua may or may not stain bluish when bruised, but typically stains
vinaceous to black when rubbed (within a few minutes) and usually has a percurrent colu¬
mella. C. caespitosa is a widespread species whose skin is white when fresh and rapidly
stains indigo-blue when bruised; it also has larger spores. C. brevicolumna has an olive-
ochre peridium that blues when bruised, plus a short broad rounded columella; it has been
found under conifers in Idaho. Hymenogasterpyriformis is a small blue-staining species
originally collected under oak in California. C. caudata has a yellowish-brown to brown
or maroon peridium and a short whitish stalk, but does not stain blue; it occurs under
oak and other hardwoods in California and Oregon.

751
752 HYMENOGASTRALES & ALLIES

TRUNCOCOLUMELLA
THIS small but common genus is distinguished by its smooth spores, branched or stump¬
like columella, and yellowish to olive color. The columella may extend below the spore
mass to form a short stalk (see photo), leading to possible confusion with the gastroid
agarics, but it is not usually percurrent. Rhizopogon is somewhat similar, but lacks a colu¬
mella, while Hymenogaster and its allies have wrinkled, striate, or warted spores. Trunco-
columella is thought to be closely related to the boletes, but it is not nearly as bolete-looking
as Gastroboletus (p. 544), which has a cap, exposed tube layer, and percurrent stalk.
At least two species of Truncocolumella occur on the west coast. One stains blue when
bruised and is perhaps better placed in Chamonixia (see key to Hymenogaster & Allies);
the other, described below, does not stain blue.

Truncocolumella citrina
FRUITING BODY 2-5 (7) cm broad, 1.5-5 cm high, irregularly rounded to oval, bulblike,
lobed, or kidney-shaped, often with a stubby stalk. Outer surface sometimes whitish when
very young but soon becoming yellow to ochre, greenish-yellow, or grayish-olive, smooth
or sometimes cracking to form a few scales or patches. Flesh(inperidiumand stalk) yellow
to buff, firm. SPORE MASS (interior) yellow-brown to olive-gray, becoming darker
(sometimes blackish) in age; composed of small empty tubular chambers, completely
enclosed by the peridium; firm when young but often quite gelatinous in old age. STALK
often (but not always) present as a short, thick, narrowed base; columella present also,
either stumplike (i.e., a broad basal region of sterile tissue) or branched and extending
into the spore mass for a considerable distance; yellow to buff. SPORES 6-10 * 3.5-5
microns, elliptical, smooth, yellowish to light brown under the microscope.
HABITAT: Solitary to gregarious under Douglas-fir (and possibly other western coni¬
fers); common, especially in the summer and fall, but fruiting practically year-round in
our area. Like most false truffles it develops underground, but often surfaces at maturity
or is dug up by rodents.
EDIBILITY: I can find no published information on it, but ethnomycologist Jim Jacobs
had a bizarre experience after putting some in an omelet: although they had little or no

Truncocolumella citrina is particularly common under Douglas-fir. Note narrowed stalklike base
in specimen at left and stout columella in sectioned specimen at right.
TRUNCOCOLUMELLA 753

flavor at the time of eating, a strong licorice-like flavor began to permeate his mouth an
hour or two later, and lingered for several hours!
COMMENTS: The yellow to buff, stumplike or branched columella plus the yellowish
to olive exterior make this a relatively easy species to identify. Other species: T. rubra has
a reddish or reddish-staining exterior, a dingy yellowish spore mass that stains blue when
cut, and larger spores. It was originally described from Washington but has been found in
the San Bernardino Mountains of southern California. For other blue-staining false
truffles, see comments under Chamonixia ambigua.

RHIZOPOGON
Small to medium-large underground or erumpent fungi found mainly under conifers. FRUITING
BODY round to oval or irregular (potato-like), variously colored. Peridium (skin) present, often
fibrillose, felty, or overlaid with rhizomorphs (mycelial strands). SPORE MASS (interior) sponge¬
like, i.e., composed of minute chambers;/im?, crisp, rubbery, or cartilaginous when young, some¬
times becoming soft or gelatinous in age; usually cinnamon-brown to dingy olive-brown or grayish
at maturity (but often white when young). STALK and columella typically absent. SPORES
hyaline (colorless) to yellowish or brownish under the microscope, typically smooth and elliptical
to cylindrical, sometimes amyloid. Basidia formingahymenium that linesthe walls of thechambers.
Capillitium absent.

THESE dingy, unattractive, potato-like fungi are the Russulas of the underworld—
unappreciated except by squirrels, but ubiquitous.* The 100+ known species are dif¬
ferentiated primarily on chemical and microscopic features such as whether or not the
spores are pronged and what color the hyphae of the peridium stain when mounted in
potassium hydroxide. However, the sameness and mundaneness of the Rhizopogons
make them relatively easy to recognize as a group. The fruiting body usually has a tough or
rubbery (“better bounced than trounced”) consistency and the interior is composed of tiny
chambers that give it a spongelike appearance (use a hand lens!). Also, the exterior is often
overlaid with mycelial strands (rhizomorphs), there is no stalk or columella (or rarely a
rudimentary columella) inside the spore mass, and the spores are typically smooth. Finally,
nearly all Rhizopogons are mycorrhizal with members of the pine family. (One unidenti¬
fied local species seems to grow only beneath cow patties or “meadow muffins,” but may
still be mycorrhizal.) In some species the spore mass becomes soft or gelatinous in old
age, but the chambers are never filled with a gel as in Alpova and Melanogaster, nor are
they separated by pallid veins, nor does the spore mass become powdery as in the puff¬
balls and earthballs.
Rhizopogons are not only the most ubiquitous of all the hypogeous (underground)
fungi, they are also among the most visible. Many are erumpent (i.e., they burst through
the surface of the ground at maturity); others are excavated by squirrels. A few species,
(e.g., R. occidentalis, R. smithii) are edible, but most have not been tested, and as already
pointed out, identification is very difficult. My own experience with them is limited. Not
only do I have an “allergy” to microscopes, but I just can’t seem to get excited about these
dingy, dumpy, dirty “small potatoes” of the mushroom microcosm when thereare so many
boletes to be picked and Russulas to be kicked. Over 200 species of Rhizopogon have
been described, most of them from western North America, but my carefully cultivated
ignorance of the subject permits only a very meager treatment here: a grand total of three
species are described and several others are keyed out. For a much more extensive treat¬
ment, see How to Know the Non-Gilled Mushrooms by Alexander Smith (who has “the
world’s largest collection of unidentified Rhizopogons”), Helen Smith, and Nancy Weber.
*Whereas the Russulas’ brittle flesh is irresistible to those who like to trounce things, the Rhizopogons’ rubbery
texture is a blessing to those who like to bounce things.
754 HYMENOGASTRALES & ALLIES

Key to Rhizopogon
1. Peridium (skin) yellow to ochre or tawny beneath a coating of brown to reddish-brown rhizo-
morphs (mycelial threads), but often developing reddish to brownish stains in age; common
under pine .R. ochraceorubens, p. 755
1. Not with above features; peridium often white or whitish when young.2
2. Fruiting body small (0.5-1.5 cm), bright yellow ... R. cokeri (see R. ochraceorubens, p. 755)
2. Not as above . 3
3. Peridium and/ or spore mass (interior) staining pinkish, reddish, or distinctly vinaceous when
cut or bruised (and often in age) .R. rubescens & many others, below
3. Not as above (but may stain purplish-gray, bluish, fuscous, or black) .4
4. Common on west coast under conifers other than pine(e.g., Sitka spruce, Douglas-fir); exterior
staining blue-black or purplish when bruised or in age . . . R. parksii (see R. e/lenae, p. 755)
4. Not as above . 5
5. Exterior usually staining or aging bluish .R. subcaerulescens (see R. ellenae, p. 755)
5. Not as above . 6
6. Peridium staining yellow when cut open ..../?. pinyonensis (see R. ochraceorubens, p. 755)
6. Not as above . 7
7. Fruiting body rather small (averaging 1-2 cm), vinaceous-tinged at least in age; spores not
amyloid; found under coastal pines.R. maculatus (see R. ochraceorubens, p. 755)
7. Not as above; fruiting body usually larger, often staining brown, gray, pinkish-gray, purple-
gray, or black with age or handling; spores amyloid or not R. ellenae & many others, p. 755

Rhizopogon rubescens (Blushing False Truffle)

FRUITING BODY 1-6 cm broad, round to oval or somewhat irregular (potato-like).
Outer surface somewhat cottony or fibrillose, often sparsely overlaid with mycelial strands
(rhizomorphs); white when young but staining pinkish to red or vinaceous when bruised
or cut, and becoming yellow to greenish-yellow or reddish-stained in age. SPORE MASS
(interior) minutely chambered, firm or rubbery to spongy or slightly gelatinous in age;
white when young, but staining pinkish when cut and becoming tawny-olive to olive-buff
to dark olive-brown or brown in age. STALK and columella absent. SPORES 5-10 ><3-4.5
microns, elliptical to oblong, smooth, usually tinged yellow under the microscope.
HABITAT: Scattered to gregarious or clustered in duff or soil under conifers (usually
buried or half-buried), especially pine; widespread and common. In our area this species
and/ or its numerous look-alikes (see comments) can be found whenever it is damp.
EDIBILITY: Delectable—if you’re a squirrel!
COMMENTS: The reddish-staining peridium (skin) is the outstanding feature of this
Rhizopogon, but there are many other “blushing” species that can only be separated
with certainty by using a microscope. Their ranks include: R. smithii, an aromatic species
that grows with Boletus edulis in pine forests and is eaten by Italian-Americans as a white
truffle substitute (it is whitish when young, reddens when bruised, and darkens consi¬
derably with age); R. vinicolor, a common associate of Douglas-fir that is white when
young and becomes vinaceous-red in age; R. roseolus, which becomes pinkish in age; R.
subaustralis, which develops black stains in age; R. succosus, with a strong rotten egg
odor in age; R. occidentalis, a common white to yellow, edible western species said to have
a sweetish taste and skin that stains orange to reddish-brown when rubbed and reddish
when sectioned; R. couchii, widespread, with few or no rhizomorphs and a pink-staining
spore mass when fresh; R. evadens, similar to R. smithii but with a disagreeable “metallic”
(Smith) odor in age, very common with ponderosa pine;/?, atroviolaceus, with strongly
amyloid spores and a white fruiting body that stains pinkish, then develops grayish-purple
to dark brownish tones in age; and/?, subsalmortius, a diffcult-to-identify species “com¬
plex” that is white to salmon-pink with salmon- or vinaceous-tinged rhizomorphs on its
exterior, especially common under mountain conifers (e.g., fir and spruce).
Rhizopogon ochraceorubens is one of many Rhizopogons with prominent mycelial threads (rhizo-
morphs) on its exterior (specimen at right). The interior (left) is white when young, darker in age. It is
also shown on p. 740. (Herb Saylor)

Rhizopogon ochraceorubens
FRUITING BODY 2-8 cm broad, round to oval or somewhat flattened or irregular
(potato-like). Outer surface yellow to tawny or ochraceous when young, but usually over¬
laid with conspicuous brown to reddish-brown rhizomorphs (mycelial strands), often
mottled with red or reddish-brown areas (especially in age), sometimes reddening some¬
what when bruised or discoloring brown to rusty-brown. SPORE MASS (interior) pallid
to grayish becoming olive or olive-brown in age, firm or spongy becoming tough as it dries;
minutely chambered. STALK and columella absent. SPORES 6-8 * 2-3 microns, oblong,
smooth, hyaline or tinged yellow or greenish under the microscope.
HABITAT: Scattered to gregarious or clustered (usually buried, sometimes erumpent)
under pine; widely distributed in western North America and very common. It is often
abundant in the Sierra Nevada as well as along the coast whenever it is damp enough.
EDIBILITY: Unknown.
COMMENTS: The conspicuous brown to reddish-brown rhizomorphs that cover the
exterior of the fruiting body are a distinctive though by no means unique feature of this
species. It doesn’t redden as much as R. rubescens, and is not white when young; there are
several similar species which can only be differentiated microscopically. Other species:
R. maculatus is a small (1-2 cm) vinaceous-tinged species with small rhizomorphs on its
outer surface and a grayish mature spore mass that becomes quite hard as it dries; it occurs
under coastal pines. R. cokeri (-R. truncatus) is an infrequent but widely distributed
species with a small (0.5-1.5 cm), bright yellow fruiting body that does not stain; it grows
under conifers also, particularly white pines. R. pinyonensis has a peridium that stains
yellowish when broken; it is one of several Rhizopogons associated with pinyon pine.

Rhizopogon ellenae
FRUITING BODY 1 -9 cm broad or long, oval to elongated to roundish, often somewhat
flattened. Outer surface white when young but developing dingy pinkish-gray to purplish-
lilac to fuscous (dark purple-gray or smoky) tones in age or after handling; smooth to
fibrillose or with a few rhizomorphs (mycelial strands). SPORE MASS (interior)
minutely chambered, firm and white when young, becoming gray to olive-gray in age.
STALK and columella absent. SPORES 7-9 * 3-4 microns, elliptical to oblong, smooth,
hyaline (colorless) or tinged yellow under the microscope, weakly amyloid at maturity.

755
Rhizopogon ellenae is very common under pine in many parts of the West. Note relatively large
size (for a false truffle) and dark stains.

HABITAT: Scattered to gregarious or clustered in duff or soil under conifers (particu¬

larly pine), often erumpent (breaking through the ground); common in western North
America. I have seen large fruitings under pine in the Sierra Nevada and along the coast.
EDIBILITY: Unknown.
COMMENTS: This is one of several medium-sized to rather large Rhizopogons that
stain dingy purplish- or pinkish-gray (not reddish as in R. rubescensl) to fuscous when
handled and/or in age. Other species: R. idahoensis also has amyloid spores (a rather
unusual feature for a Rhizopogon)', it stains brownish to grayish-lilac when bruised and
is especially common in the northern Rockies under Douglas-fir. R. amyloideus has
strongly amyloid spores, but is more prevalent in the coastal forests of northern Cali¬
fornia. R. colossus is a large species that is whitish when fresh but stains brown to blackish
when handled or dried; it has non-amyloid spores and, like the others, occurs under
western conifers. R. parksii is a very common coastal species, particularly under Sitka
spruce and Douglas-fir. It is pallid to brown or pinkish, but stains dingy purple to bluish-
black on its surface when bruised or with age. R. subcaerulescens tends to develop
bluish or greenish stains when handled or in age and is widely distributed. It is white at
first, dark vinaceous-brown in age. Also see the species listed under R. rubescens.

ALPOVA & MELANOGASTER

Small to medium-sized underground fungi. FRUITING BODY roundish to oval or knobby
(potato-like), variously colored but not normally white. SPORE MASS (interior) yellow to
greenish, brown, or black at maturity; composed of numerous chambers filled with spore-
containing gel, the chambers separated by paler, meandering veins. ST ALY^and columella typically
absent. SPORES elliptical to cylindrical and often truncate or cupped at one end, smooth, non¬
amyloid; thick-walled and brown to dark brown or purple-black, with a pore (Melanogaster),
thin-walled, pore-less, and hyaline (colorless) to medium brown under the microscope (Alpova).
Basidia and spores contained inside the chambers, not forming a hymenium. Capillitium absent.

THESE false truffles, which comprise the family Melanogastraceae, are easily told by
their chambered spore mass that is gelatinous and colored at maturity and marbled with
pallid or yellow veins. Microscopically they are also distinctive by virtue of their smooth,
lemon-shaped to cylindrical spores. In Alpova, the spores are colorless to brownish under
the microscope and the spore mass ranges in color from yellow to brown when mature. In
Melanogaster, on the other hand, the spores are darker and the mostcommon species have
a dark brown to black mature spore mass. The marbled interior can lead to confusion with

756
Left: Close-up of an Alpova (A. olivaceotinctus), showing the numerous gel-filled chambers that
comprise the spore mass. (Herb Saylor) Right: Melanogaster species are easily told by their blackish
interior marbled with white to yellow-orange veins. This is probably M. variegatus.

the true truffles (e.g., Tuber), which, however, have a harder, non-gelatinous interior and
bear their spores inside asci rather than on basidia. Hysterangium is also somewhat
similar, but usually has a translucent columella instead of veins, while Leucogaster and
Leucophleps have a whitish spore mass even when mature.
Both Alpova and Melanogaster are widely distributed. Neither is worth eating (unless
you’re a squirrel or wood rat!), although the latter has been used in Europe as a cheap
substitute for truffles. One representative of each genus is described here and several
others are keyed out.

Key to Alpova & Melanogaster

l. Peridium (skin) bright yellow when young (but often mottled with reddish, orange, or brown
in age); interior (spore mass) brownish-yellow to brownish-black but not truly black; asso¬
ciated with western conifers .A. trappei(see A. diplophloeus, below)
1. Not as above . 2
2. Interior (spore mass) typically black, at least at maturity; spores strongly brown to dark brown
under the microscope, usually with rather thick walls and a pore .
.Melanogaster euryspermus & others, p. 758
2. Not as above (spore mass sometimes brownish-black, but usually paler and spores paler under
the microscope) . 3
3. Typically associated with alder ... A. diplophloeus, below
3. Not as above .4
4. Found in western North America .A. alexsmithii& others (see A. diplophloeus, below)
4. Found in eastern North America . 5
5. Peridium (skin) darkening when handled or bruised; associated with conifers .
.A. olivaceoniger {see A. diplophloeus, below)
5. Not as above; found mostly with hardwoods .... A. nauseosus(see A. diplophloeus, below)

Alpova diplophloeus (Alder False Truffle; Red Gravel)

FRUITING BODY 0.5-3 (5) cm broad, oblong or oval to nearly round, or less commonly
irregularly lobed. Outer surface pallid to pinkish-yellowish-buff when young, soon be¬
coming yellow-brown to cinnamon, reddish-brown, or brown, usually staining dark
reddish-orange to reddish-brown where bruised; smooth or with only a very few rhizo-
morphs (mycelial threads); odor usually fruity at maturity. SPORE MASS (interior)
composed of small chambers separated by pale yellow or whitish veins; chambers 0.5-3
mm broad, filled with a gelatinous substance, pallid to pale yellowish or olive when young,
Alpova diplophloeus. Note somewhat gelatinous interior (specimen on left) and smooth exterior.

becoming orange-brown to reddish-brown to dark vinaceous-brown at maturity. STALK

and columella absent. SPORES 4-6 * 1.5-3 microns, elliptical to oblong, smooth, hyaline
(colorless) under the microscope.
HABITAT: Widely scattered to gregarious in humus or soil (usually buried) under or near
alder; widely distributed, but particularly common in western North America. I have
found it in the fall and winter in coastal California and in the late summer, fall, and spring
in the Sierra Nevada and Cascades.
EDIBILITY: I can find no information on it.
COMMENTS: Also knownas^4. cinnamomeusandRhizopogondiplophloeus, this false
truffle can be distinguished by its cinnamon-colored exterior (when mature), fondness
for alder (a tree not noted for its diversity of false truffles), and its spore mass composed
of small, gel-filled chambers and pallid veins. At maturity the interior (spore mass) is
reddish-brown and somewhat reminiscent of “red gravel.” It can even turn dark vinaceous-
brown, but never becomes black as in Melanogaster. Other species: A. trappei(-A. luteus)
is a brilliant yellow species that becomes dull yellow to orange- or reddish-brown (or
mottled with these colors) in age. It has a brownish-yellow to brownish-black spore mass
and a garlicky to slightly fruity odor when mature. It grows with fir, Douglas-fir, and
other western conifers; I have found it locally in the winter and spring. A. olivaceotinctus
is a yellowish-brown or olive-tinged western species that is apparently rare. A. alexsmithii
is yellowish-brown to dark brown and grows with mountain conifers (especially hem¬
lock). Melanogaster parksii of California is said to have a brown interior. Alpovas in
eastern North America include: A. olivaceoniger, a brownish-olive conifer-lover that
darkens when handled and has a fruity odor at maturity; A. nauseosus, brown to reddish-
brown, with a yellow-brown to blackish-brown interior and strong aroma of rotten fruit
at maturity, favoring hardwoods; and A. superdubius, whose stalklike rooting base causes
false truffle expert James Trappe to be “superdubious” as to whether it is really an Alpova.

Melanogaster euryspermus (Black Veined False Truffle)

FRUITING BODY 1.5^1 (6) cm broad, round to elongated or lobed (potato-like), often
with rhizomorphs (mycelial threads) at base. Outer surface slightly fibrillose to downy
or felty, rusty-brown to reddish-brown, warm brown, or dark brown (but often yellower
when young), often blackening where bruised. SPORE MASS (interior) composed of
large (up to 5 mm broad) blackish gel-filled chambers separated by whitish to yellow
meandering veins; odor usually strong in age (sometimes like sewer gas, at other times with
a pleasant citrus component). STALK and columella absent. SPORES 9-18 * 7-11
microns, lemon-shaped (but with two small prongs at one end), thick-walled, smooth;
dark brown under the microscope.
HABITAT: Solitary, widely scattered, or in small groups in duff or soil under both hard¬
woods and conifers; widely distributed and fairly common (for a false truffle), but rarely
fruiting in large numbers. In California this species and M. variegatus occur throughout
the mushroom season. I have found them under oak, tanoak, and madrone, and in the
Sierra Nevada, under conifers.
758
Melanogaster variegatus (see comments under M. euryspermus). Note brown exterior (right) and
gelatinous black interior that is divided into chambers by paler veins (left). (Herb Saylor)

EDIBILITY: According to European sources, M. variegatus (see comments) is edible

when young. I can find no information on American material.
COMMENTS: This species and its close relatives are easily distinguished from other
false truffles by their blackish gelatinous spore mass composed of large chambers separ¬
ated by pallid to yellow veins. They might be mistaken for true truffles because of the veins,
but they bear their spores on basidia and are much softer and stickier inside. The name
M. variegatus has traditionally been used for our common species. However, the
genus is being critically studied and species concepts may be revised. It is very similar to
M. euryspermus, but has elliptical spores that are usually less than 11 microns long. M.
intermedius is also similar, but has spores intermediate in size between those of M. eury¬
spermus and M. variegatus. M. ambiguus has large, elongated spores and a spongy, more
olive-colored exterior plus a somewhat metallic odor in age; I usually find it under Doug-
las-fir. M. macrocarpus is a large (up to 12 cm!) conifer-loving species with large spores.
M. parksii of California is said to have paler brown spores and a brown rather than black
spore mass. See also Alpova trappei and A. nauseosus (under A. diplophloeus).

LEUCOPHLEPS & LEUCOGASTER

Small to medium-sized underground woodland fungi. FRUITING BODY roundish to oval or
knobby (potato-like), usually pallid in Leucophleps and usually brightly colored in Leucogaster.
Peridium (skin) sometimes overlaid with rhizomorphs. SPORE MASS (interior) white or pallid
even when mature; composed of numerous chambers that are filled with a clear spore-containing
gel and/or exude a white latex when cut (unless old). STALK and columella absent or rudimen¬
tary. SPORES hyaline (colorless) under the microscope, round to elliptical, pitted-reticulate or
spiny within a clear gelatinous outer wall, not amyloid. Basidia and spores contained inside the
chambers, not forming a hymenium. Capillitium absent.

THESE potato-like fungi can be told by their white or pallid interior which is composed
of gel-filled chambers (which may, however, be very small). Fresh specimens often exude a
white latex when cut, but the spores are not amyloid as in Zelleromyces, and there is no
stalk or columella. The interior of most other false truffles is darker when mature and/ or
has a different texture.
In Leucophleps the exterior of the fruiting body is usually whitish, the chambers are
tiny, and the spores are spiny (within their outer wall), while in Leucogaster the exterior
is often brightly colored, the chambers are usually larger, and the spores are pitted or
reticulate. Their evolutionary affinities are unclear, and modern taxonomists isolate
them in their own order, the Leucogastrales. Both are fairly small genera, but relatively
common, especially under conifers. As is commonly the case with false truffles, many
of the species are poorly known and/or difficult to identify. One representative is
described here and several others are keyed out.

759
760 HYMENOGASTRALES & ALLIES

Key to Leucophleps & Leucogaster

1. Peridium (skin) white or at times pale yellowish, ochraceous, or grayish; spore mass chambers
minute; spores prickly inside a clear gelatinous outer wall . 2
1. Peridium usually darker or more brightly colored than above; spore-bearingchambers relatively
large (easily visible); spores reticulate or pitted within clear outer wall . 3
2. Peridium white or whitish (or grayish in age), usually covered with conspicuous mycelial threads,
at least near base .Leucophleps spinispora, below
2. Not as above .Leucophleps magnata (see L. spinispora, below)
3. Peridium pinkish to reddish or reddish-brown (often browner in age); found under western
conifers .Leucogaster rubescens (see Leucophleps spinispora, below)
3. Not as above .Leucogaster odoratus & others (see Leucophleps spinispora, below)

Leucophleps spinospora (White Jellied False Truffle)

FRUITING BODY 1 -3 cm broad, round to oval or irregularly lobed (potato-like). Outer
surface white or whitish, often becoming grayish in age, usually wrinkled and typically
covered with rhizomorphs (mycelial threads), but young specimens usually showing rhizo-
morphs only at base. SPORE MASS (interior) white, often exuding a white latex when
cut, composed of many tiny (less than 1 mm broad), clear, gel-filled chambers (unless old
or late to develop); sterile veins absent. STALK and columella absent. SPORES 10-13 *
10-11 microns, round or nearly so, with small spines inside a clear gelatinous outer wall.
HABITAT: Solitary to gregarious in soil or duff (usually buried) under western conifers.
In the Sierra Nevada it is fairly common from spring through fall. The duff of our coastal
forests is more apt to cough up species of Leucogaster (see below) when ruffled for truffles.
EDIBILITY: Unknown, but the texture is hardly appealing.
COMMENTS: This common whitish false truffle might be mistaken for a Rhizopogon
because of its rhizomorphs, but the white gel-filled chambers and/ or presence of a latex
distinguish it. Specimens which mature slowly, however, may lack a gel and latex or be
hard and Tuber-like. Other species of Leucophleps are bestdifferentiated microscopically.
One, L. magnata, can usually be told by its whitish to pale yellowish or ochraceous ex¬
terior with less conspicuous rhizomorphs, plus its larger spores with larger warts or spines.
Leucogaster is also fairly common. Its ranks include: Leucogaster rubescens, with large
gel-filled chambers and an exterior that becomes pinkish to brick-red to brownish as it
ages; L. odoratus, less common, with a yellowish-orange exterior, large chambers, and
a pungent odor in age; and L. carolinianus, one of several species in eastern N orth America.

RADIIGERA
Medium-sized woodland fungi usually found underground. FRUITING BODY typically round
or slightly flattened. Peridium (skin) thick, composed of two or three layers of tissue. SPORE
MASS (interior) with threads or plates that radiate from the columella, at first white and fleshy,
then becoming gooey and finally powdery; brown to black at maturity. STALK absent, but a
columella present; columella fairly thick, often rounded, white, unbranched, not percurrent.
SPORES warted or spiny, round, brown under the microscope. Capillitium present.

THIS small genus is easily recognized by its thick skin and rounded, unbranched columella
(see photo). The multi-layered peridium (skin), warted or spiny round spores, powdery
spore mass in old age, presence of capillitium, and rounded columella (or “pseudocolu¬
mella,” to be technically accurate) indicate a close relationship to the earthstar genus
Geastrum, and it is placed in the Lycoperdales by most taxonomists. However, the outer
skin never splits into rays as it does in Geastrum, and the underground growth habit is
also distinctive. It is not a particularly common genus. Three species are keyed below, but
undescribed species also occur, at least in California.
The interior of an immature Radiigera, showing the thick rounded columella and thick skin. This
species was never determined because the spores had not yet matured; it was growing under an oak.

Key to Radiigera
1. Outer peridium (skin) splitting into rays when mature (look around for older specimens); ex¬
terior usually smooth (but often dirty); sometimes growing just below the ground, sometimes
on top .(see Geastrum, Astraeus, & Myriostoma, p. 699)
1. Peridium not normally rupturing or splitting; exterior smooth or felty; usually underground 2
2. Mature spore mass blackish; outer layer of peridium rather heavy and felty R. atrogleba, below
2. Not as above; mature spore mass brown to dark brown . 3
3. Fruiting body typically 2-4 cm broad; spores warted .../?. taylorii(seQ R. atrogleba, below)
3. Fruiting body typically 3-8 cm broad; spores spiny . . R.fuscogleba(see R. atrogleba, below)

Radiigera atrogleba
FRUITING BODY 2.5-5 cm broad, round to slightly flattened or depressed, often im¬
bedded in white mycelial threads (rhizomorphs). Peridium (skin) three-layered. Outer
surface tough and felty or fibrous-cottony, rough, white to grayish, often developing buff,
pinkish, vinaceous, or brownish tints in age; usually separating readily from the two inner
layers. Inner layers closely adhering to each other, whitish to rosy or pinkish when fresh
(but may turn olive or buff when bruised), 3-5 mm thick, white when sectioned (or with a
greenish tinge). SPORE MASS (interior) composed of plate-like bundles of hyphae which
radiate from the columella to the peridium; white, fleshy, and rather softatfirst, becoming
gooey or inky and blackish, and eventually powdery (and blackish); odor often rather
unpleasant in the inky stage. STALK absent, but the basal portion of peridium usually
thickened slightly and giving rise to a prominent columella which is nearly round, cushion¬
shaped, or narrowed at the base and enlarged (rounded) at the apex; columella usually
0.8 cm thick or more at the widest portion, usually penetrating the spore mass at least half
way (never percurrent); white or tinged gray, sometimes disintegrating in old age.

Radiigera atrogleba, maturing specimen. Interior (shown at right) is black and gooey in this stage,
but eventually turns powdery. The top of the columella can be seen at the center, but its base has been
obscured by the gooey spore mass. Note how thick the skin is. (Herb Saylor)
Radiigera fuscogleba, immature specimens. Note how lines radiate from the columella in sliced
specimen. See comments below for more details.

SPORES 5.5 -6.5 microns, round, minutely warted, deep brown under the microscope.
HABITAT: Solitary or more often in groups or large, mycelium-incrusted clusters in soil
or duff in woods and along old roads through the woods, usually buried or only partially
exposed; not uncommon in the Pacific Northwest and California, but probably more
widespread. Radiigeras occur mainly with conifers in the summer and fall, but I have found
R. taylorii (see comments) locally under oak and pine in the winter.
EDIBILITY: Unknown; the gooey black spore“mess” of older specimens isn’t appetizing!
COMMENTS: The thick, often rounded or cushion-shaped columella plus the thick skin
and blackish spore mass at maturity make this a most distinctive fungus. Scleroderma
species also have a thick skin and dark spore mass, but lack a columella. The spore mass
passes through a gooey stage before becoming powdery. In this stage (see photo) it looks
like the “ink” from a shaggy mane and stains everything it comes into contact with. Other
species: R. fuscogleba is quite similar, but has an olive-brown to brown mature spore mass
and spiny spores (see photo above). R. taylorii, on the other hand, is smaller, with a
smoother brownish to drab exterior, paler brown spore mass, and warted spores. Both of
these species occur in California and the Pacific Northwest but may be more widespread.

HYSTERANGIUM & Allies

Small to medium-sized, mostly woodland fungi usually found underground. FRUITING BODY
typically round to irreguarly lobed (potato-like), often with one or more mycelial threads or cords
at base. SPORE MASS (interior) composed of small chambers, usually greenish to olive-brown to
olive-gray (but not always); typically firm and tough when young but usually gelatinous or muci¬
laginous in old age; odor sometimes fetid at maturity. STALK typically absent, but a columella
usually present; columella whitish or translucent, branched or unbranched, rarely percurrent.
SPORES spindle-shaped to oblong or elliptical, smooth or encased in a wrinkled utricle (outer sac);
hyaline (colorless) to greenish-brown under the microscope. Capillitium absent.

HYSTERANGIUM is closely related to the stinkhorns (Phallales), but is treated here

with the false truffles because of its hypogeous (underground) tuberlike fruiting body.
Like the stinkhorns, most Hysterangiums have a greenish to olive-brown spore mass that
becomes slimy and stinky in old age. Those species with mycelial cords at the base are
reminiscent of stinkhorn “eggs,” but never hatch, i.e., a specialized spore-bearing structure
never emerges. Hysterangium is a fairly sizable genus. Most of its 30+ species occur in the
woods and are best differentiated microscopically. One very common and farflung species
is described here. A related genus, Phallogaster, is included in the key.

Key to Hysterangium & Allies

1. Fruiting body with a stalk or distinctly narrowed and elongated base, pear-shaped or developing
lobes on top; spore mass divided into distinct chambers; found on rotten wood, mainly in
eastern North America .Phallogaster saccatus (see Hysterangium separabile, below)
1. Not as above . 2

762
Hysterangium separabile, immature specimens. In this stage the spore mass (shown in specimens on
left) is odorless and quite firm or cartilaginous. Note presence of a branched, translucent columella.

2. Fruiting body showing a thick gelatinous layerjust under the peridium(skin) when sliced open,
eventually “hatching” if old enough (i.e., a specialized spore-bearing structure emerging from
it); one or more thick mycelial cords usually emanating from the base; often found in lawns,
gardens, and other urban areas (but also in woods) .(see Phallales, p. 764)
2. Not with above features; usually found in woods . 3
3. Columella typically percurrent (extending all the way through spore mass); often fruiting
above the ground.H. darkeri(see H. separabile, below)
3. Columella not typically percurrent; usually underground or only partially exposed.4
4. Associated with eucalyptus .H.fuscum(see H. separabile, below)
4. Not as above . 5
5. Spore mass (interior) tough and pale pinkish when young; found under oak . . H. occidentale
5. Not as above .6
6. Fruiting bodies enmeshed in a copious mass of white mycelial threads; usually growing
in rocky or gravelly soil .H. crassum(see H. separabile, below)
6. Not as above ..H. separabile & many others, below

Hysterangium separabile
FRUITING BODY round to somewhat flattened or lobed, 0.5-3 cm broad, sometimes
with mycelial fibers at the base. Outer surface white to pinkish or sometimes buff, often
becoming pinkish where bruised or handled. Peridium (skin) easily cracking or peeling
away from the spore mass, at least at maturity. SPORE MASS (interior) olive-brown to
greenish, composed of small chambers; at first firm and tough (cartilaginous), but be¬
coming slimy and stinky (putrid) at maturity or in old age. STALK absent, but a columella
present; columella thin, translucent or whitish, arising from a rudimentary sterile base,
often branched and typically extending about half way into the spore mass. SPORES
12-19 x 6-8 microns, more or less spindle-shaped, smooth within a wrinkled utricle (sac),
hyaline (colorless) to pale greenish-brown under the microscope.
HABIT AT: Solitary to gregarious in humus or soil (usually buried) under both hardwoods
and conifers; very widely distributed, but especially common in western North America.
In our area it is abundant under oak and other trees practically year-round. In the Sierra
Nevada and Cascades it’s common in the late spring, summer, and fall.
EDIBILITY: Edible and choice—if you’re a squirrel or chipmunk.
COMMENTS: Once called H. clathroides and also known as H. coriaceum, this subter¬
ranean fungus look like a stinkhorn “egg,” but never hatches. The greenish spore mass and
translucent, branched columella distinguish it from most other false truffles, and the ten¬
dency of the peridium to crack and peel easily from the spore mass is also distinctive.
There are a number of similar species that differ microscopically. A few can be recog¬
nized in the field, including: H. setchellii and H. crassirhachis, with thick whitish septa
(partitions) emanating from a scarcely branched columella(the latter especially common);
H. aureum, with a golden-brown exterior; H. crassum, whose fruiting bodies are en¬
meshed in a copious mass of white mycelial threads, fairly common under western hard¬
woods and conifers, particularly in rocky or gravelly soil; H. darkeri, a slightly larger (1 -5
Hysterangium separabile. Spore mass is putrid and gelatinous in old age (specimen on right), but rub¬
bery or tough when young, and skin is easily separable. Note mycelial cords emanating from base.

cm) species that has a percurrent columella and often grows above the ground; H.fuscum
(-H. fischeri), associated with eucalyptus; and H. stoloniferum, fruiting body anchored
by a thick mycelial cord or “stolon” and spore mass sometimes bluish-tinged.
Another stinkhorn relati\e, Phallogaster saccatus, is quite different. It usually grows on
decayed wood and has a more or less pear-shaped fruiting body with a tapered, stalklike
sterile base. The peridium is white to pinkish or lilac, and typically develops lobes and
shallow depressions at the top which rupture irregularly at maturity in order to expose
the spores. The spore mass is mucilaginous, olive-colored, and fetid at maturity, and
separated into several large chambers within the fruiting body. It is fairly common in
eastern North America but absent in our area. Also see Rhopalogaster(in key on p. 724).

Stinkhorns
spores

PHALLALES
STINKHORNS are among the most fascinating and highly specialized of the fleshy
fungi. They differ from other Gasteromycetes in having a slimy or sticky, putrid spore
mass (gleba) that is borne aloft at maturity. However, they begin as puffball-like “eggs”
completely encased in a skin (peridium), and usually anchored by a thick mycelial cord(s).
A sectioned stinkhorn “egg” reveals a differentiated interior with a gelatinous substance
beneath the outer skin, a compressed stalk (and/or arms), and a brown or olive-colored
spore mass (see Color Plates 191-198). The superficial resemblance to a puffball is soon
rudely—and forever—dispelled, however, as the enclosed stalk or other spore-bearing
structure elongates rapidly, bursting out of its “shell” and into the world.*
In the common stinkhorns (Phallaceae), the mature fruiting body is unbranched and
explicitly phallic, with the spore slime coating the apex of the stalk or“head.” In the ornate
stinkhorns (Clathraceae), the fruiting body is branched or latticed and the spore mass
generally coats the inside surfaces of the branches or latticework. In both families the rup¬
tured peridium forms a sack (volva) at the base of the fruiting body, much as in Amanita. * *
*This rapid elongation process, which can take as little as one hour, is possible because all of the stinkhorn’s parts
are fully formed (though greatly compressed) within the “egg.” Its emergence is mainly an act of expansion
(elongation of cells) rather than of growth or development. As a result, stinkhorn “eggs” can be “hatched” at
home by keeping them in a humid environment.
**Though their developmental stages are superficially similar, the stinkhorns differ from the Amanitas in several
fundamental respects. Stinkhorns lack gills, and like other Gasteromycetes, do not forcibly discharge their spores.
The spores, although borne aloft at maturity, are produced internally (within the “egg”). Amanitas, on the other
hand, do not form their spores until the cap has expanded. Their spores are produced on gills and are forcibly
discharged when ripe.
764
PHALLALES 765

The stinkhorns’ most outlandish feature, however, is the unpleasant or provocative

odor of the mature spore slime, which has been variously characterized as “foul,” “fetid,”
“evil,” “odious,” “obnoxious,” “cadaverous,” “putrid,” “maddening,” “aggravating,”
“compelling,” “intolerable,” “filthy,” “vile,” “disgusting,” “distressing,” “disconcerting,”
“spermatic,” “garbageous,” “nauseating,” “like rotting carrion,” “like spent incense,”
“like the damp earthy smell we meet with in some of our churches on Sundays,” “enough
to cause one to think that all the bad smells in the world had been turned loose,” and most
apt and understated of all: “indiscreet.” Lured from afar by the stench, flies and carrion
beetles come to feast on the slime, and if the day ishot,rollaroundinit. With their eventual
reluctant departure, spore dissemination is accomplished (some spores stick to their feet,
others are presumably passed through their digestive tracts). All in all, it is a rather
ingenious method of spore dispersal more typical of the so-called “higher” plants than
of the “lowly” fungi.
Opinions on the edibility of stinkhorns range from ill-disguised disgust to idle specu¬
lation to passionate praise. The “intolerable” odor of mature specimens would seem
to be enough to discourage even the most ardent and confirmed toadstool-tester from
sampling them. However, the odorless stinkhorn “eggs” are considered a delicacy in
parts of China and Europe, where they are pickled raw and even sold in the markets
(sometimes under the name “devil’s eggs”). Captain Charles Mcllvaine, of course, pro¬
nounces them delicious (see comments on edibility of Phallus impudicus), suggesting
they be sliced and fried like a Wiener schnitzel. My own experience with them has left
me nonplussed (see comments on the edibility of Clathrus archeri). None are known
to be dangerously poisonous, but it is reported that a young person of twenty-two, “having
eaten a morsel, was seized with violent convulsions, lost the use of his speech, and ulti¬
mately fell into a stupor which lasted forty-eight hours; prompt attention was given to
him, but it appears to have been some months before he was perfectly cured.” (Note: one
source described the victim as a “young English girl.”)
Stinkhorns are notoriously spontaneous and unpredictable in their habits—they are
liable to pop up at almost anytime, anywhere, providing conditions are to their liking.
They occur throughout the world but attain their maximum diversity in the tropics.
Several of our species were probably introduced accidentally—surely no one would do it
on purpose!—along with soil or plant material from exotic lands. They are not common
in comparison to other fleshy fungi, but they command a degree of attention far dispro¬
portionate to their numbers because of their fantastic shapes and repugnant odors. For
this reason I have keyed out the common North American species, plus several “aliens”
that may turn up in greenhouses, botanical gardens, the backyards of stinkhorn
specialists, or other hospitable places. They are, after all, among the most beautiful of all
mushrooms—providing you hold your nose!

Key to the Phallales

1. Spore slime exposed or borne aloft at maturity; fruiting body variously shaped, emerging from
an “egg” whose peridium (skin) forms a sack or volva . 2
1. Not as above; peridium either remaining intact or rupturing to form irregular holes at the top;
volva absent .(seeHysterangium& Allies, p. 762)
2. Fruiting body unbranched (but sometimes with a drooping lacy skirt or “veil”); spore slime
coating the outer surface of the apex or “head” (unless washed off) .Phallaceae, p. 766
2. Fruiting body branched to form several arms (which may or may not be fused at their tips),
columns, or a lattice-like framework; spore slime borne on the inner surfaces of the arms
or latticework (but if arms unfold, the inner surfaces become the upper surfaces).
.Clathraceae, p. 772
766 PHALLALES

PHALLACEAE (Common Stinkhorns)

Medium-sized, foul-smelling (at maturity) fungi found on ground or rotten wood. FRUITING
BODY emerging from a round to oval "egg” that has an inner gelatinous layer; roughly cylindrical
and unbranched when fully expanded, with or without a swollen head. STALK present asacolumn
by which the spore slime is elevated; hollow, spongy, usually perforated at the tip. VEIL absent,
or present as a drooping, netlike “skirt” (indusium) below the “head.” VOLVApresent as a mem¬
branous sack or pouch at base of stalk. SPORE MASS usually greenish to dark olive-brown,
mucilaginous or slimy at maturity, coating the exterior of the “head” or top of the stalk. Spores
more or less oblong (bacillus-like), smooth. Capillitium absent.

THE common stinkhorns are unmistakable by virtue of their explicitly phallic fruiting
body which is simple (that is, unbranched), but may terminate in an enlarged “head” on
which the foul-smelling spore slime resides. Three genera are common in North America:
Phallus, which has a well-defined cap or “head”; Dictyophora, which resembles Phallus
but boasts a netlike veil or indusium that hangs from the lower edge of the “head”; and
Mutinus (the so-called “dog stinkhorns”), which lacks a differentiated “head.” (Dictyo¬
phora is incorporated into Phallus by some stinkhorn specialists, and Ithyphallus is an
outdated synonym for Phallus.)
Whether or not stinkhorns are handsome or repulsive has been the subject of con¬
siderable debate. The verdict seems to rest largely on personal prejudice and one’s ability
to overlook their obnoxious odor and the swarms of blowflies that come to wallowin their
spore slime. They are undeniably phallic, however, and as might be expected, their sug¬
gestiveness has given rise to a veritable “mother lode” of stinkhorn lore. For instance,
German hunters believed they grew where stags rutted, and it is said that their putrid
carcasses are burned outside houses in Thailand to discourage unwanted guests (a rather
drastic practice that might discourage wanted guests as well!). They’ve been used in count¬
less ointments and potions, e.g., as a cure for gout, epilepsy, and gangrenous ulcers.
They’ve been blamed for cancer and prescribed as a sure-fire remedy for it. And of course,
they’ve been employed as aphrodisiacs, and are supposedly still given to cattle for that
purpose in some parts of Europe. Alexander Smith tells of a remarkable encounter with
an elderly gentleman who carried a mammoth specimen under his hat in the certain
belief that it would cure his rheumatism! And, in an otherwise tedious book of Victorian
reminiscences by Gwen Raverat, there is this astonishing passage:
In our native woods there grows a kind of toadstool, called in the vernacular The
Stinkhorn, though in Latin it bears a grosser name. This name is justified, for the
fungus can be hunted by the scent alone; and this was Aunt Etty’s greatest invention:
armed with a basket and a pointed stick, and wearing special hunting cloak and gloves,
she would sniff her way round the wood, pausing here and there, her nostrils twitching,
when she caught a whiff of her prey; then at last, with a deadly pounce, she would fall
upon her victim, and then poke his putrid carcass into her basket. At the end of the
day’s sport, the catch was brought back and burnt in the deepest secrecy on the drawing¬
room fire, with the door locked, because of the morals of the maids.

One wonders how such an innocuous, splendid, upright organism could be so ruthlessly
and unjustly maligned, and I submit that this particularly perverse brand of fungophobia
be christened phallophobia. (“Aunt Etty,” incidentally, was Charles Darwin’s daughter!)
Attitudes change, however, and in the twentieth century the stinkhorns’ impudence
and imprudence have begun to be appreciated. Mycological literature is replete with
wonderfully provocative accounts of close encounters (of the casual kind) with stinkhorns
-as riddled with them, in fact, as an old Suilluspungens is with maggots. When stinkhorns
are discussed, the language makes a startling and unprecedented qualitative leap, from
monotonous minutiae to half-baked hyperbole, as if the authors were suddenly taking
Fruiting body development in the tropical basket stinkhorn, Dictyophora indusiata (p. 770). This
sequence was photographed within a time period of thirty minutes! Note how the ridges on the cap
become more visible in age (as the spore slime is carried away by flies and/ or rain). In old specimens
completely bereft of spore slime the cap is whitish. (Keith Muscutt)

an interest in what they were saying. They are lavish in their praise as they tread the fine
line between double-entendre and forthright fungal fact: “The mischief-maker is a hand¬
some specimen, as its plate shows”... “This is a highly specialized type of fungous fruiting
body” . . . “It is one of the seven unnatural wonders of the natural world”... “Never have
I seen such intricate lacework as on a Phallus” ... “It undoubtedly emerges from the depths
for a single noble and grand purpose—that of disseminating its spores. All of its parts
have been developed to accomplish this function in the most effectual manner possible”...
“It is inconspicuous while encased in its skin, and unlikely to be noticed by anyone who
is not looking for it. When fully elongated, however, it is pointedly, inescapably promi¬
nent” . . . “By extraordinary growth and expansion it carries the banquet of spores into
the light—a natural Jack-in-the-box” ... “When its expansion is complete, it begins imme¬
diately to become limp, bent, sinks down, and undergoes putrefaction” . . . “Once while
collecting fungi with other students we smelled a phalloid and tried to trace it down in
order to learn the species, but it seemed to keep moving. Finallly, we noticed that an old
man was also in the woods collecting fungi, and as I worked over toward him I realized
that a phalloid of some kind was very close” . . . “One is curious to learn the mechanism
by which so much is accomplished in apparently so short a time, and find in this instance,
as in all others where great things are accomplished with ease, that many forces have been
slowly at work to insure everything being in readiness for the success of the final flourish”
.. . “The banquet is prepared underground and the table, with its viands ready, is pushed
into the light while the invitation to guests is wafted swiftly on the breeze” . . . “It is a
glorious fungus—an admirable specimen of herculean proportions pokes up periodically
in my front yard.”
The common stinkhorns are more prevalent in temperate regions than their ornate
cousins, the Clathraceae, but still attain their greatest diversity in the tropics. Species
of Phallus are especially fond of populated areas, where they lurk in lawns, gardens,
flower beds, along roads, in ditches, under bushes, hedges, porches, and houses, in the
vicinities of churches and lumberyards, trash heaps, and old sawdust piles. Mutinus and
Dictyophora species, on the other hand, are more frequent in forests and are largely
restricted to eastern North America and the tropics. All of these fruit whenever
conditions are conducive to development—that is, warm and moist—and none are
known to be poisonous. One species from each genus is described here, and several others
are mentioned and/or keyed out.

767
768 PHALLALES

Key to the Phallaceae

1. Stalk with a latticed head, the spore slime coating the inside of the latticework .
.(see Clathraceae, p. 772)
1. Not as above; “head” absent or if present, then spore slime coating its exterior.2
2. Lacelike or netlike skirt (indusium) present and usually prominent (hanging from the lower
margin of cap) . 3
2. Skirt (indusium) absent or rudimentary . 5
3. Fruiting body, or at least the skirt, yellow to orange-yellow or even orange-red; primarily
tropical . Dictyophora multicolor (see D. indusiata, p. 770)
3. Fruiting body typically white (excluding the spore slime and volva); temperate or tropical . 4
4. Skirt (indusium) large, often reaching the ground; tropical . . . Dictyophora indusiata, p. 770
4. Skirt smaller (3-6 cm long); fairly common in forests of eastern North America .
. Dictyophora duplicata (see D. indusiata, p. 770)
5. Spore slime borne on a ring or “collar” beneath the tip of the fruiting body; fruiting body white
except for the slime; tropical . Staheliomyces cinctus
5. Not as above; spore slime not borne on a sharply defined ring or collar .6
6. Spore slime borne on the upper portion of the stalk, i.e., a sharply defined “head” not present;
stalk typically slender (up to 1.5 cm thick) . 7
6. Spore slime borne on a differentiated (sharply defined), often swollen “head”; stalk sometimes
slender, but usually at least 1.5 cm thick .9
7. Fruiting body entirely or partially red to pink or orange . 8
7. Fruiting body white except for spore slime Mutinus caninus var. albus(see M. caninus, p. 771)
8. Fruiting body 9-17 cm long, usually slightly thicker in the middle and tapered gradually toward
the tip .Mutinus elegans (see M. caninus, p. 771)
8. Fruiting body 5-10 cm long, usually more or less equal except for the very tip, which is bluntly
and rather abruptly narrowed .Mutinus caninus, p. 771
9. Fruiting body red to scarlet (at least in part), usually quite slender .
.Phallus rubicundus (see P. impudicus, below)
9. Fruiting body white to pale pinkish(except for spore slime), but volva may be pink to purple 10
10. Cap or “head” reticulate (pitted and ridged) beneath the spore slime; widely distributed .
.Phallus impudicus & others, below
10. Cap or “head” not reticulate, pitted, or ridged, but often granular; found mainly in eastern
North America .Phallus ravenelii(see P. impudicus, below)

Phallus impudicus (Stinkhorn) Color Plates 193,194

FRUITING BODY beginning as an “egg” up to 6 cm high. PERIDIUM (skin) white to
yellowish-white in one variety, lurid pinkish to purple in another (see comments), with a
gelatinous layer beneath; rupturing to form a volva as the stalk elongates and thrusts the
slimy, swollen “head” upward. CAP (“head”) 1.5-4 cm broad, coated with the putrid,
copious spore slime which eventually drips off or is carried or washed away, revealing
the whitish reticulate (pitted and ridged) surface beneath; top with a hole which is some¬
times covered by a clinging piece of the peridium (volva) and/ or by numerous flies.
STALK 1.5 -3 cm thick, equal or tapered at both ends, entirely white or sometimes pinkish
below; minutely honeycombed (spongelike), hollow, fragile. INDUSIUM (“veil”) absent
or rudimentary. VOLVA present at base of stalk as a white to pinkish or purple, loose,
lobed sack formed by the ruptured peridium; base usually with a thick, similarly-colored
mycelial cord(s). SPORE MASS slimy or mucilaginous, olive-green to olive-brown,
with an obnoxious odor at maturity (see p. 765). Spores 3-5 * 1.5-2.5 microns, elliptical
or oblong, smooth.

HABITAT: Solitary or in groups or clusters in lawns, gardens, sandy or cultivated soil,

under trees or shrubs, in rich humus, etc.; widely distributed and especially common in
the West, fruiting whenever conditions are favorable. Mcllvaine states that, “its favorite
This purple-egged variety of Phallus impudicus is considered a distinct species, P. hadriani, by some
stinkhorn specialists. Left: A round “egg” with telltale mycelial cord. Right: An “egg” just prior to
hatching, and a malodorous full-grown (but rather small) specimen. Note how the spore slime coats
the reticulate “head”; after it has been dispersed the “head” is whitish. See color plates for another
full-grown specimen and a beautiful sectioned “egg.”

abode is in kitchen yards and under wooden steps where, when mature, it will compel the
household to seek it out in self-defense,” and his contemporary Nina Marshall says, “the
distracted housewife searches in vain for a solution to the difficulty and the odor disappears
as mysteriously as it came. If she is one of the initiated, however, she will search until she
finds the haunt of the offender andthen destroy it on the spot to avoid further repitition of
the nuisance.” The largest fruitings I have seen were on the lawn of a school in Los Angeles,
and in front of an old cathedral in Santa Fe, New Mexico. Stinkhorn specialist William
Burk recalls seeing massive clusters that apparently sprung from the foundation of
a house in Salt Lake City, Utah. If the “eggs” are carried home and transplanted in
cool, damp earth, more often than not they will continue to develop so that the fasci¬
nating elongation process can be observed firsthand.
EDIBILITY: Not poisonous (see comments on p. 765), but as Alexander Smith says,
“who would want to eat even the eggs?” Captain Charles Mcllvaine (the plenipotentiary
extraordinaire of turn-of-the-century toadstool testers), for one. He says, “(the eggs) are
semigelatinous, tenacious, and elastic, like bubbles of some thick substance. In this
condition, they demand to be eaten . . . cut in slices and fried or stewed, they make a most
tender, agreeable food.”
COMMENTS: This is one “wild” mushroom that is truly unmistakable. In shape it resem¬
bles nothing so much as a slimy cigar, leaky pipe, malodorous thumb, putrid horn, hollow
baton, spongy candle, or putrescent pencil. But the most distinctive thing about the stink¬
horn is its utter spontaneity. In the words of one founding member of the New York
Mycological Society, “you never know when one is going to pop up right in front of you,
so fast you can actually see it growing. And there is no way to predict or control them. It’s
the one thing in the world you can’t push or hurry.” The stinkhorn also hasa highly refined
sense of poetic justice. One “going strong” is said to have appeared in the concrete floor
of the newly built house greeting a newly married couple. To accomplish this instructive
feat, the mycelial cords had to force their way into the foundation from an old stump in the
garden!
The stinkhorn is sometimes mistaken inexplicably for an old morel, perhaps because
770 PHALLALES

of the pitted head and somewhat similar shape. However, the “indiscreet” odor and slimy
spore mass (both of which may be gone in old age) and presence of a volva(and frequently,
a squadron of flies) leaves no doubt as to its identity. The common variety in California
has a pinkish to purple peridium (see Color Plate 194) and is considered a distinct species,
P. hadriani (-P. imperialis, P. iosmus) but some authorities on the subject. Other species:
P. ravenelii is very similar, but has a smooth to granular rather than reticulate “head.” It is
quite common in eastern North America, especially in lignin-rich humus and on old
sawdust piles. The tropical “Devil’s Stinkhorn,”/*. rubicundus, is a rather long, slim, red
to scarlet species. It might be mistaken at first glance for a dog stinkhorn (Mutinus), but
has a clearly differentiated, detachable, more or less conical or bell-shaped “head.” It
occurs in southern and eastern North America (at least as far west as New Mexico), but
is not common. See also Dictyophora duplicata (under D. indusiata).

Dictyophora indusiata (Basket Stinkhorn) Color Plate 191

FRUITING BODY beginning as an “egg” up to 6 cm high. PERIDIUM (skin) white or
sometimes tinged buff, gray, or reddish-brown, with an inner gelatinous layer; rupturing
to form a volva as the stalk elongates. Mature fruiting body 7-25 cm tall, unbranched,
consisting of a more or less conical to bell-shaped cap on a stalk, with a large lacelike veil
or “skirt” flaring out from beneath the cap. CAP (“head”) 1.5-4 cm broad, coated at first
with the putrid olive-green to brown spore slime, which eventually drips off or is carried
away by insects, revealing a white or yellowish, ridged and pitted (reticulate) surface
beneath; top with a hole in it. STALK 1.5-3 cm thick, equal or tapered at either end, white,
hollow, fragile, sometimes curved, minutely chambered and porous (spongelike). INDU-
SIUM (“veil”) white, initially tucked under the cap margin but soon unfurling; skirtlike
or basketlike when fully expanded and touching or nearly touching the ground (6 cm or
more high); attached to the top of the stalk or just under the margin of the cap; composed
of white strands that form an intricate chainlike or lacelike net whose “holes” are large
and polygonal. VOLVA present at base of stalk as a white or pallid, loose, often lobed
sack formed by the ruptured peridium; base usually with a thick mycelialcord(s) attached.
SPORE MASS slimy or mucilaginous, olive-green to olive-brown or brown, with a fetid
odor at maturity that attracts flies, beetles, and other insects (but odor not always strong,
and in one form somewhat sweetish). Spores 3.5-4.5 x 1.5-2 microns, elliptical to oblong,
smooth.
HABITAT: Solitary or in groups in humus or on rotten wood in tropical forests and at
their edges; fruiting in wet weather, widely distributed. It is quite common in Central
America and South America, as well as in Australia, the South Pacific, Africa, India,
and Japan.
EDIBILITY: Edible in the egg stage, but probably not choice (see comments on edibility
of Clathrus archeri)—in other words, it is better eyed than fried. In many parts of the
world (e.g., New Guinea) it is worshipped for its beauty and/ or used as an aphrodisiac.
COMMENTS: What’s an indisputably tropical fungus doing in a reputedly topical field
guide like this? Well, I could justify its presence on the flimy pretext that it just might
show up in somebody’s hot house, sauna, or backyard bamboo forest. Or I could cite the
popular but shamefully arrogant political notion that Central America is our “backyard,”
and that any stinkhorn found in one’s backyard merits mention. However, the truth is
that I’ve included it because I couldn’t resist including it. In my opinion, such a piece
de resistance of nature deserves a full-fledged description no matter where it grows (and I
offer the color plate as proof). The lacy or netlike “veil” which is responsible for the name
“basket stinkhorn” is initially hidden (greatly compressed), but swells out soon after the
DICTYOPHORA 771

stalk elongates (see photographs on p. 767). In its prime the veil touches—or nearly
touches—the ground, and may be globelike or broadly skirtlike. Later, however, it shrinks
or collpases somewhat and begins to droop. Aside from its spectacular skirt, D. indusiata
resembles our common stinkhorn (Phallus impudicus) in shape, size, color, and texture
(including the pitted head), and some phallologists consider it “a good Phallus.” Other
species: D. duplicata (the “Veiled Stinkhorn”) is a robust temperate zone version that is
fairly common in the hardwood forests of eastern North America. It resembles D. in¬
dusiata, but is not so spectacular, the skirt being smaller-meshed and only 3-6 cm long
(it looks somewhat like the second photo on p. 767). D. multicolor is a colorful (yellow
to orange-red) tropical species. D. rubrovolvata of China has a reddish volva.

Mutinus caninus (Dog Stinkhorn)

FRUITING BODY beginning as an “egg” up to 2.5 (4) cm high. PERIDIUM (skin) white
or occasionally with a faint pinkish or yellowish tinge, with an inner gelatinous layer;
rupturing to form a volva at base of stalk. Mature fruiting body 5-10 cm high and 0.5-1.2
cm thick, unbranched, slender, erect or curved slightly, roughly cylindrical (equal) or
thicker near top; lacking a differentiated cap, but with a blunt, rounded or abruptly nar¬
rowed, often perforated tip. FERTILE PORTION covering the upper 2-3 cm of fruiting
body (except the very tip), bright orange-red to red to pink, but covered at first with olive
to olive-brown spore slime. STALK more or less equal, colored like cap (orange-red to
orange or pink) or often paler or even white toward the base; hollow, fragile, spongy
(minutely chambered). INDUSIUM (“veil”) absent. VOLVA present at base of stalk
as a white, lobed sack or pouch formed by the ruptured peridium; base usually with one
or more white mycelial cords attached. SPORE MASS mucilaginous, olive to deep olive-
brown, coating the upper portion of stalk; odor fetid at maturity (but not as malodorous
as some stinkhorns). Spores 3-7 * 1.5-2.5 microns, elliptical or oblong, smooth.
HABITAT: Solitary to gregarious or clustered on ground and rotten wood in gardens,
roadsides, woods, etc.; widely distributed and fairly common in eastern North America
in the late summer and fall (along with M. elegans—see comments). It is apparently absent
in the West, but a white variety (see comments) has been found in Oregon, and it seems
only a matter of time before the typical form and its longer look-alike, M. elegans, show up
on the west coast, as several other erotic exotics have.

EDIBILITY: Nonpoisonous. Mature specimens, of course, are hardly tempting, but Bill
Roody of Elkins, West Virginia, says the “eggs” are excellent “peeled and rolled in flour
seasoned with garlic salt and pepper, dipped into beaten egg and then once again in the
flour mix before frying in butter or oil. They’re great as is or served with crackers and
cream cheese.”
COMMENTS: The slim, cylindrical, pink to orange-red fruiting body capped with olive-
brown spore slime is a unique and memorable—if suggestive—sight, and it is easy to see
how this “phalloid” got its common name (see color plates of M. elegans). The odor of
mature specimens can be rather feeble compared to that of Phallus impudicus or Clathrus
ruber, but is obnoxious nevertheless. The stalk varies from orange or pink to white, and
a variety that is entirely white (except for the spore slime) has been found in Oregon and
Michigan. (It has been called M. caninus var. albus, but may very well be a distinct species.)
Another dog stinkhorn, M. elegans (-M. bovinus, M. curtisii) (COLOR PLATES 195,
196) is sometimes called “Devil’s Dipstick.” It is also common in eastern North America,
and is frequently mistaken for M. caninus. It can be distinguished, however, by its longer
(9-18 cm) fruiting body that is usually thickest in the middle and tapered gradually toward
the tip. Also, a larger area of the fruiting body (up to 6 cm) is smeared with spore slime. In
my experience it favors the same habitats as M. caninus, but is more common.
772 PHALLALES

CLATHRACEAE (Ornate Stinkhorns)

Medium-sized, foul-smelling (at maturity) fungi found on ground or rotten wood. FRUITING
BODY emrging from a round to oval or flattened “egg" that has an inner gelatinous layer; usually
branched to form arms, columns, tentacles, or a latticedframework; often brightly colored. ST ALK
absent or if present then hollow, spongy, fragile, and often perforated or open at the top. VEIL
(indusium) absent. VOLVA present as a membranous sack or pouch at base of fruiting body.
SPORE MASS usually greenish to dark olive-brown (or drying blackish), mucilaginous or slimy
at maturity, coating the inner surfaces of the arms, tentacles, or latticed framework. Spores more
or less oblong (bacillus-like), smooth. Capillitium absent.

THESE stinkhorns are more fantastic than—but not nearly as phallic as—the common
or “true” stinkhorns (Phallaceae). Some have long “tentacles” that make them look like
starfish, others have stubby arms, still others boast two or more thick columns or an
elaborate, lovely lattice-like framework. None look like “horns,” but all of them stink
(to a greater or lesser degree), and flies seem to find them just as desirable as members of
the Phallaceae. Thus the label “stinkhorn,” though not completely accurate, is amply
deserved. In most cases the spore slime coats the inward-facing surfaces of the latticework,
columns, branches, or tentacles (or the upper surfaces of the tentacles if they unfurl). As in
the Phallaceae, the stalk (if present) is tubular, spongy, and fragile, and emerges from a
membranous peridium (volva) which has an inner gelatinous layer.
Most of the stinkhorns in this family are tropical, but several have established them¬
selves as “resident aliens” in the U nited States. The three most common genera are: Clath-
rus, whose fruiting body is composed of columns, a latticed ball, or four or more tentacles
(with or without a short stalk); Pseudocolus, with three or four tentacles and a short stalk;
and Lysurus, which has several thick arms ora latticed “ head” atop a long stalk or column.
Most of the ornate stinkhorns are edible in the egg stage, but in the words of Alexander
Smith, “are unlikely to become popular as food” (for obvious reasons!). One species,
Clathrus ruber, is said to be poisonous raw, but this reputation may rest on a single bizarre
incident (see p. 765). Five species are described here and several others are keyed out.
Key to the Clathraceae
1. Stalk typically much longer than the arms or fertile “head”; arms (if present) typically short
(less than 3 cm long) and thick . 2
1. Stalk absent (but two or more fused columns may be present), or if stalk present then relatively
long (2.5 cm or more) and slender arms or “tentacles” also present; stalk when present usually
short but sometimes fairly long .4
2. Fruiting body consisting of a rounded to somewhat flattened latticed “head” or netlike frame¬
work mounted on a stalk . Lysurus periphragmoides, p. 776
2. Not as above; stalk with several short arms that may or may not be fused at their tips .3
3. Arms usually bright red and fused at their tips to form a “spire” (but sometimes breaking free
from each other in age); stalk usually fluted or several-sided .Lysurus mokusin, p. 776
3. Arms red to pinkish, flesh-colored, brownish, or white, their tips sometimes touching at first
but not fused into a “spire,” and usually separating in age; stalk more or less cylindrical (i.e.,
round in cross-section) .Lysurus cruciatus, p. Ill
4. Fruiting body with three or more slender arms or “tentacles” which arise from stalk (but stalk
may be very short and hidden by the volva, or occasionally practically absent). 5
4. Fruiting body either a latticed ball or netlike framework or comprised of two or more thick
columns which are usually fused at their summit; stalk absent or rudimentary .7
5. Fruiting body typically with 3-4 “tentacles” that often remain joined at their tips .
.Pseudocolusfusiformis(see Clathrus archeri, p. 774)
5. Fruiting body typically with 4 or more “tentacles” that may initially be joined at their tips but
which usually break free and sometimes bend back in age .6
6. Arms or “tentacles” diverging from a flat disclike expansion of the stalk apex, often appearing
divided or paired; spore slime mostly in center of fruiting body or coating bases of the arms;
tropical .Aseroe rubra (see Clathrus archeri, p. 774)
6. Not as above; spore slime borne on inner or upper surfaces of arms . Clathrus archeri, p.774
CLATHRACEAE 773

7. Fruiting body lantern-like, i.e., the spore slime borne in a specialized structure slung between
2-4 red to orange or pink columns that merge at the top to form an arch; tropical (reported
from Miami, Florida) . Laternea triscapa
7. Not as above; fruiting body with columns or a latticed framework, but lacking a separate spore¬
bearing structure . 8
8. Fruiting body consisting of a latticed network of branches and “windows” (somewhat like a
“whiffle ball”—see photo on p. 774) .Clathrus ruber & others, below
8. Fruiting body composed of2-5 thick columns joined at their summit(the columns occasionally
with one or two transverse branches, but not enough to form a lattice) .9
9. Fruiting body composed of two creamy to yellow to orangish columns at first pressed closely
together but then bowing at the center; rare (native to Japan) .Clathrus bicolumnatus
9. Fruiting body typically with (2) 3-5 red to pink or orange columns; widely distributed and not
uncommon (especially in eastern North America) Clathrus columnatus {see C. ruber, below)

Clathrus ruber (Latticed Stinkhorn)

FRUITING BODY beginning as a round to somewhat flattened or knobby “egg” up to 6
cm broad. PERIDIUM (skin) white and membranous, smooth becoming wrinkled and
grooved as it gets larger, with an inner gelatinous layer; rupturing to form a volva. Mature
fruiting body 5-14 cm high, consisting of a round to oval latticed ball or netlike framework
with large polygonal or elongated “windows.” BRANCHES of the framework bright pink
to red to orange or pale orange, and often paler toward the base; flattened, hollow, very
fragile, minutely chambered (like a sponge) and transversely ribbed or wrinkled on the
outer surfaces; inner surfaces covered with the sticky, putrid spore slime. STALK absent
or rudimentary. VOLVA present at base of fruiting body as a thick, loose, white sack or
pouch, usually with thick mycelial cord(s) attached. SPORE MASS coating the inside
surfaces of the latticework; mucilaginous, olive to olive-brown or drying blackish, with an
extremely obnoxious stench at maturity. Spores 5-6 * 1.5-2.5 microns, oblong, smooth.
HABITAT: Solitary to densely gregarious or clustered in soil, wood chips, rich humus,
etc.; widely but erratically distributed, apparently fruiting most any time (providing con¬
ditions are favorable). It is said to be a native of southern Europe (its likeness appears on
more than one postage stamp from that region), but has turned up in many localities in
North America (Florida, Virginia, North Carolina, etc.), usually in landscaped areas or
other places where exotic plants have been introduced. It is a common sight in the parks of
San Francisco (especially in the late fall), and I have also seen it in Santa Clara County.

EDIBILITY: Nonpoisonous according to some sources, harmful (at least raw) according
to others (see the quote on stinkhorn poisoning on p. 765).
COMMENTS: Also known as C. cancellatus, this “wild” mushroom looks like a red or
orange “whiffle ball” (see photograph). The white volva from which it emerges and the
intolerable stench that develops at maturity are also distinctive. The color of the branches
varies considerably—from bright red to very pale orange—apparently depending on the
temperature and humidity. The putrid perfume attracts flies in large numbers and is,
in my humble fungal opinion, the vilest of any stinkhorn. It must be smelled to be believed!
(According to Ramsbottom, the smell is “so fetid that more than one artist has related
that it was impossible to paint it without discomfort. . . when the color photograph was
taken it would have been pleasing to have shown one or two flies at work, but they settled
in such swarms that they had to be driven off.”). The “eggs,” on the other hand, are quite
odorless and very knobby or furrowed just prior to bursting. In this stage they are easily
“hatched” at home. C. crispus is a similar but even more spectacular neotropical species
whose “windows” are circumscribed by “coronas”; it has been found in Florida. A species
of the Old W orld tropics, C. crispatus, is also quite similar, but the upper part of its lattice-
work regularly breaks up into fragments in age. C. columnatus (-Laternea columnata)
has 2-5 thick, orange to red columns which arise independently but are fused at the top and
Clathrus ruber is easily recognized by its bright red to pink to pale orange latticework. Left Mature
specimens; note absence of a stalk. Right: Note how lumpy this “egg” is!

may branch horizontally (but not enough to form a latticework). It is widely distributed
and has been found in many localities in eastern North America (particularly the South),
plus Hawaii. C. preussii and Ileodictyon cibarium (-C. cibarius) are but two of many
southern and/ or tropical stinkhorns with a delicate white latticework and large “win¬
dows.” The first is a native of Africa; the latter is common in the southern hemisphere.

Clathrus archeri (Octopus Stinkhorn) Color Plates 197,198

FRUITING BODY beginning as a round to somewhat flattened “egg” up to 6 cm in dia¬
meter. PERIDIUM (skin) membranous, white to dingy buff, pinkish, or tinged purple,
smooth or scurfy, with a gelatinous inner layer; rupturing to form a volva at base of stalk.
Mature fruiting body 5-12 cm high, consisting of (4) 5-7 (8) arms or “tentacles” arising
from a common stalk or “tube,” or sometimes with up to 12 arms arising from two fused
stalks. ARMS long and slender, 3-9(12) cm long, tapered toward their tips, at first upright
and roughly parallel and joined at their tips (often in pairs), soon opening outward like
the petals of a flower and eventually curling back and under so that the tips often touch
the ground (but sometimes one or more arms branched, and at other times the tips
remaining joined to others). Inner (or upper) surfaces of arms bright red to pinkish-red,
usually paler toward base, ribbed or reticulate and coated with spore slime. Outer sur¬
faces (undersides) pale pink, longitudinally grooved. STALK 1-3 (5) cm long, typically
short and sometimes hidden by the volva (but a long-stalked form is also known); hollow
and tubular (open at the top), detaching easily from the volva, white or pallid below,
pinkish above. VOLVA present at base of stalk as a loose sack formed by the ruptured
peridium; base usually with one or more mycelial cords (rhizomorphs). SPORE MASS
coating the inner (upper) surfaces of the “tentacles,” mucilaginous, olive to dark olive-
brown when fresh, but usually blackening as it dries; odor obnoxious at maturity (some¬
what reminiscent of rotting crab). Spores 4-5.5 (7) * 2-2.5 microns, elliptical, smooth.

HABITAT: Solitary to gregarious or clustered on ground (especially sandy soil) or rotten

wood in various habitats; widely distributed (Tasmania, Russia, etc.), but rare in North
America. It is fairly common in the riparian woodlands along the San Lorenzo River in
Santa Cruz County, California (especially in the spring, but fruiting most any time). 1
have also found it under a rose bush growing on a mixture of compost and rice hulls.
EDIBILITY: N onpoisonous, and like most stinkhorns, edible in the egg stage. One spring
evening a former friend and I decided to sample two “eggs,” following a recipe for French-
fried stinkhorn eggs (see comments on edibility of Mutinus caninus). The flavor wasn’t
bad, but we neglected to strip away the gelatinous outer layer. It proved to be so slippery
that the “eggs” slid down our throats before we could savor them, leaving behind only the
sticky spore mucilage (see Color Plate 198), which clung to our throats and tongues so
tenaciously that we were still trying to wash it away several hours later!

774
Clathrus archeri is aptly called the “Octopus Stinkhorn” or“Stinkopus” because of its long, tentacle¬
like arms. They are intially joined at their tips (central specimen), but then separate and recurve(spe-
cimen at left). Note cords emanating from the “egg” in foreground. Also see the color plates.

COMMENTS: The octopus stinkhorn—or “stinkopus,” as I am fond of calling it—is

one of my five favorite fleshy fungal fructifications. It is a most attractive fungus when
fresh, easily told by its long red “tentacles” which are initially joined at their tips but then
unfold to form a slender-armed “star.” It definitely looks more like a sea creature than a
mushroom, but its obnoxious aroma—which attracts flies and jaded fungophiles from
afar—identifies it as a member of the stinkhorn tribe. The arms are longer than those in
Lysurus, and the stalk is proportionately shorter (or at times practically absent). It was
formerly placed in the genus Anthurus (as A. archeri or A. aseroeformis), but since the
“tentacles” are sometimes branched to form a rudimentary latticework (when young), it
is now classified as a Clathrus. A common tropical stinkhorn, Aseroe rubra (see photo
below) is sometimes confused with C. archeri, but looks more like a starfish or sea anenome
than an octopus. It has five to ten or more “tentacles” which are deeply divided lengthwise
(so that they may seem to occur in pairs) and arise from a flattened, disclike expansion of
the stalk apex. The spore slime rests in the center of the disc or coats only the very bases

Top view of the tropical stinkhorn Aseroe rubra (see comments above). Note how arms are deeply
divided and spore slime is concentrated at center. Stalks are not visible in this photo. (M ichael Fogden)
776 PHALLALES

of the “tentacles.” Although strictly tropical, it may turn up, like so many other exotic
fungi, in a greenhouse or similarly sultry environment. Also similar to C. archeri is the
“Stinky Squid ” Pseudocolus fusiformis(-P. schellenbergiae, P. javanicus). It is common
in scattered localities in eastern North America (especially Chapel Hill, North Carolina,
the home of stinkhorn specialist William Burk), as well as in Europe, Australia, Asia, and
Tierra del Fuego. It has a white to brown volva and only three to four “tentacles” which
are red-orange to orange or pinkish, arise from a common stalk, and are usually fused at
their tips (see photo on p. 777) but sometimes break free in age.

Lysurusperiphragmoides (Stalked Lattice Stinkhorn)

FRUITING BODY beginning as a round to oval “egg” up to 5 cm broad. PERIDIUM
(skin) white to buff, with a gelatinous inner layer, rupturing to form a volva at base of stalk.
Mature fruiting body 6-16 cm high, composed of a small rounded latticed “head” on a long
stalk. CAP or “head” 1.5-3.5 cm broad, round to somewhat flattened, comprised of a
latticework of red to orange (or sometimes yellowish or white) branches which form rather
small meshes or “windows”; outer edges of branches keeled. STALK 5-13 cm long, 0.8-3
cm thick, hollow, fragile, rather spongy, equal or tapered, usually red above and paler
below, but yellow or white in some forms. VOLVA present at base of stalk as a loose, lobed
sack or pouch formed by the ruptured peridium, usually with whitish mycelial cords at¬
tached to base. SPORE MASS coating the inside surfaces of the latticed “head” and some¬
times spilling out, mucilaginous, dark olive to olive-brown or drying blackish; fetid at
maturity. Spores 3.5-4.5 * 1.5-2.5 microns, elliptical to oblong, smooth.
HABITAT: Solitary to gregarious (occasionally two arising from the same volva) in rich
soil, lawns, gardens, open woods, on rotten wood, etc.; widely distributed. It is not uncom¬
mon in mild wet weather in the southern United States and Midwest (North Carolina,
Texas, Nebraska, New Mexico, even New York), but has yet to be found in California.
EDIBILITY: Presumably edible in the egg stage, but who wants to eat it?
COMMENTS: Also known as Simblum sphaerocephalum and 5. texense, this
southern stinkhorn looks like a miniature Clathrus ruber mounted on a long stalk. In
some specimens the transverse (horizontal) branches of the network are reduced or even
absent, making its relationship to other species of Lysurus more obvious. White and
yellow variants are known, but the red one is the most common in North America.

Lysurus mokusin (Lantern Stinkhorn) Color Plate 192

FRUITING BODY beginning as a round to oval “egg” up to 6 cm high. PERIDIUM (skin)
white and membranous, with an inner gelatinous layer; rupturing to form a volva at base
of stalk and sometimes also leaving a piece of tissue stuck to the top of fruiting body.
Mature fruiting body (3) 5-12 (16) cm high, consisting of a stalk that is branched above
into 4-6 (usually 5, rarely 7) arms which normally remain joined at their tips to form a
“spire,” but which sometimes break free from each other. ARMS 0.8-3 (4) cm long, short
and thick, usually bright red, erect and only sightly separated, or bowed to form a lantern¬
like structure; 3-sided; outer surfaces roughened, with a central longitudinal ridge; sides
at first coated with spore slime; “spire” long or short (1-20 mm long), erect or bent down¬
ward. STALK 6-13 cm long, 0.5-2 cm thick, equal or more often tapered downward, flesh-
pink to pink or reddish-pink above, paler below and usually white at base; hollow, tubular,
fragile, minutely chambered and marked by longitudinal rows of V-shaped depressions;
cross-section usually showing 4-6 (7) angles (same number as arms). VOLVA present
at base of stalk as a loose, lobed, white sack or pouch formed by the ruptured peridium,
usually with one or more white mycelial cords attached to base. SPORE MASS borne in
LYSURUS 111

the vertical slits between the arms and coating their sides, mucilaginous, light brown to
olive-brown becoming darker (blackish) as it dries, with an unpleasant odor at maturity.
Spores 3.5-4.5 * 1.5-2 microns, oblong, smooth.
HABITAT: Solitary to densely gregarious or clustered in lawns, gardens, hard-packed
soil, etc.; common in southern California and fruiting there year-round, but partial to
warm weather. It is probably native to Asia, but is well established in California at least as
far north as Fresno. It has also been found in Texas and Washington, D.C.
EDIBILITY: Edible in the egg stage, but in my opinion, no better than Clathrus archeri
(I have fried it). However, it is considered a great delicacy in China.
COMMENTS: This gaudy stinkhorn is easily recognized by its red to pink color and
distinctive shape. In the words of Paul Rea (who wrote an extensive analysis of the deve¬
lopmental stages of this species), “the form of the expanded plant may be likened to a
moat (the volval cup) surrounding the base of a tower (the polygonal stipe) bearing a
belfry or lantern (the arms) surmounted by a spire”—a very apt description, except that
the “belfry” is usually besieged by flies (see color plate)! Specimens growing in hard
ground are apt to be larger than those growing in rich or loose soil, perhaps because they
take longer to develop. The arms sometimes break free from each other, but the several¬
sided stalk and often brighter color distinguish it from L. cruciatus.

Lysurus cruciatus (Lizard’s Claw Stinkhorn)

FRUITING BODY beginning as a round to oval “egg” up to 6 cm high. PERIDIUM
(skin) white and membranous, with a gelatinous inner layer; rupturing to form a volva at
base of stalk. Mature fruiting body 6-12(16) cm high, composed of a stalk that is branched
at the top to form4-7 (usually 5) stubby arms. ARMS initially incurved and touching(but
not permanently fused) at their tips, then separating at least slightly and remaining more
or less erect (i.e., not unfolding); short and thick (1-2.5 cm long), hollow, 3-sided; outer
surfaces pallid to brownish, flesh-colored, pinkish, orange, or red, with a longitudinal
groove. Inner surfaces wrinkled and irregularly roughened or knobby, at first covered
with spore slime. STALK 6-10 cm long, 1-2 cm thick, usually tapered downward, hollow,
fragile, minutely chambered and faintly striate longitudinally; entirely white or tinged
yellowish above and white below. VOLVA present at base of stalk as a thick, loose, lobed,
white sack formed by the ruptured peridium; usually with one or more mycelial cords
attached to base. SPORE MASS supported within the arms and coating their inner sur¬
faces, mucilaginous, olive to olive-brown becoming blackish as it dries, with an unpleasant
(fetid) odor at maturity. Spores 3-4 * 1-2 microns, elliptical to oblong, smooth.
Left: Pseudocolus fusiformis (see comments at top of p. 776) has three or four long arms which are
normally fused at their tips but may separate in age; stalk is not visible in this picture. (Joan Zeller)
Right: Lysurus cruciatus (- Anthurus borealis) has several short, thick arms at apex. (Bob Tally)
778 PHALLALES

HABITAT: Solitary or in groups or clusters in lawns, gardens, under trees, in rich soil,
on rotten wood, etc.; apparently native to Australia and New Zealand, but now widely
distributed and well established in various parts of the U nited States. It is not uncommon
in southern California, but is not nearly as numerous as L. mokusin. It favors warm
weather but can be found most anytime. I have not seen it north of Santa Barbara.
EDIBILITY: Presumably edible, but see comments on the edibility of Clathrus archeri.
COMMENTS: Better known asL. (-Artthurus) borealis, this stinkhorn is easily told by its
short, thick arms which separate slightly but do not unfold a la Clathrus archeri. It can be
dull or brightly colored, and the tips ofits arms are neverfusedintoa spire asinL. mokusin.
It has often been confused with/,, gardneri, a tropical species whose arms have sterile bases.

Bird’s Nest Fungi

spores

NIDULARIALES
Tiny fungi found on soil, wood, dung, and vegetable debris. FRUITING BODY usually rather
tough, at first round to cylindrical or cushion-shaped, usually becoming cup- to mug-shaped(nest¬
like) at maturity, with several “eggs” enclosed (only one “egg” in Sphaerobolus). PERIDIUM
(wall of “nest”) conposed of 1 -4 layers. PERIDIOLES (“eggs”) usually flattened and lens- or
lentil-shaped, white to gray, brown or black, sometimes with minute cords attached. STALK
absent (but base of “nest” may be narrowed). SPORES borne inside the peridioles, typically
smooth and hyaline. Capillitium absent.

THESE minute Gasteromycetes look like miniature bird’s nests. The fruiting body
typically consists of a tiny vase or “nest” (peridium) furnished with several spore capsules
or “eggs” (peridioles), and in most cases a protective membrane or “lid” (epiphragm)
that initially covers the top of the nest. As such there is very little that the bird’s nest fungi
can be confused with, though when very young they might be mistaken for minute puff¬
balls, and when old and bereft of eggs they look like tiny, tough cup fungi.
The spores are dispersed with the aid of raindrops and animals. The force of a single
raindrop will splash the eggs out of the nest and as much as seven feet away (hence they
are sometimes called “splash cups”). The outer wall of the egg then decays or is eaten away
by insects and the spores within are exposed. Whereas most fungi produce millions of
spores, the bird’s nest fungi need only a few—each egg contains the correct mating strains,
so a fertile secondary (dikaryotic) mycelium develops directly, obviating the need for
large numbers of spores.
There are four common genera of bird’s nest fungi. In Cyathus and Crucibulum, each
egg is anchored to the nest by a minute cord or “stalk” (funiculus), while in Nidula and
Nidularia the eggs are imbedded in a sticky gel. Both the cords and the sticky mucilage help
the splashed eggs to adhere to whatever they land on. Also treated in this chapter is
Sphaerobolus, a dynamic relative of the bird’s nest fungi that is sometimes called the
“cannon fungus.” It features only one egg that is shot out with terrific force, accompanied
by an audible “pop.” (The Latin name literally means “sphere-thrower.”)
Bird’s nest fungi are found on various types of organic or vegetable matter—rotting
wood and sticks, herbaceous stems, humus, dung, and manure (the spores of dung-
inhabiting individuals presumably pass through—and out of—grazing animals). They
are gregarious creatures, but as they rarely attain heights of more than 15 mm, they are
difficult to see and worthless as food. Most of the common North American species are
keyed out here, but additional exotic types may turn up in greenhouses and flower pots.
NIDULARIALES 779

Key to the Nidulariales

1. Fruiting body containing only one peridiole (“egg”), round at first, then splitting open to form
4-9 orange rays around the central “egg” .Sphaerobolus stellatus, p. 781
1. N ot as above; fruiting body cylindrical to mug- or cup-shaped when mature. Containing more
than one egg (unless all but one have been expelled) . 2
2. Fruiting body lacking a “lid” when young, at first round to cushion-shaped and more or less
covered with a felty or powdery material that ruptures irregularly; nest soon disintegrating;
eggs imbedded in a sticky mucilage which eventually dries out .3
2. Not as above; fruiting body typically witha“lid” when very young, the nest usually well formed
and persistent; eggs may or may not be imbedded in a mucilage .4
3. Eggs brown to reddish, round and flattened (lens-shaped); not common .... Nidularia farcta
3. Eggs grayish-brown and often irregularly-shaped; fairly common . Nidulariapulvinata
4. Eggs gray to brown or dark brown, imbedded in a sticky mucilage or jellylike substance which
eventually dries out; eggs not attached to the nest by a small cord or stalk; sides of nest usually
vertical (i.e., fruiting body more or less mug-shaped) . 5
4. Eggs white to gray, brown, or black, often (but not always) attached to side of nest by a minute
cord or short stalk, not imbedded in a mucilage; sides of nest vertical to tapered .6
5. Exterior of nest white and shaggy when fresh Nidula niveotomentosa (see N. Candida, p. 780)
5. Exterior of nest scurfy or hairy, brown to cinnamon or at times grayish A1 idula Candida, p. 780
6. Interior of nest longitudinally striate (with distinct radial grooves).
.Cyathus striatus & others (see C. stercoreus, p. 780)
6. Interior of nest smooth or at least not striate or grooved . 7
7. Eggs typically white to buff; interior of nest not black ... Crucibulum laeve &. others, below
7. Eggs typically gray to brown or black; interior of nest variously colored (including black) . 8
8. Fruiting body typically less than5 mm high; exterior of nest smooth to fibrillose but not shaggy;
found on sticks, dung, etc., in arid regions . . . Cyathuspygmaeus (see C. stercoreus, p. 780)
8. Fruiting body typically 5-15 mm high; exterior smooth or shaggy; cosmopolitan.9
9. Exterior of nest shaggy (with long hairs), at least when young; fruiting body averaging5-10 mm
high, the rim more or less circular at maturity .Cyathus stercoreus, p. 780
9. Exterior of nest fibrillose to smooth even when young (without long hairs); fruiting body
averaging 10-15 mm high, the rim often wavy in age Cyathus o//a(see C. stercoreus, p. 780)

Crucibulum laeve (Common Bird’s Nest Fungus)

FRUITING BODY tiny, at first nearly round, becoming cylindrical and then deeply
cup-shaped (i.e., with a wide flaring mouth); 5-12 mm high and broad (at the top) when
mature, the rim more or less circular and covered at first by a hairy lid. Peridium (wall
of nest) one-layered, tough, persistent. Exterior velvety or shaggy, yellowish or tawny to
cinnamon-brown, becoming nearly smooth in age and often darker or whiter. Interior of
nest smooth, somewhat shiny, white to silvery, gray, or pale cinnamon. PERIDIOLES
(eggs) 1-2 mm in diameter, several, whitish to buff or with a very slight brownish tinge,
circular but flattened (lens- or disclike), usually attached to nest by long thin cords. Spores
(4)7-10 x 3-6 microns, elliptical, thick-walled, smooth, hyaline.
HABITAT: Scattered to densely gregarious on sticks, wood chips, nut shells, vegetable
debris, humus, and manure; widely distributed. It is not as common locally as some of the
other bird’s nest fungi, but occurs year-round, sometimes in the company of Cyathus.
EDIBILITY: Much too miniscule to merit being munched on.
COMMENTS: Also known as C. levis and C. vulgare, this fetching and farflung little
fungus is easily told by its whitish eggs that are initially attached to the nest by long, thin
cords. It lacks the sticky mucilage of Nidula, and the eggs are never lead-colored or black
as in Cyathus. Other species: Crucibulum parvulum is a similar but even more minute
(2-4 mm high) species with a white to grayish or buff exterior. It grows on dead juniper
and other organic matter in arid habitats.
Left Cyathus stercoreus growing on dung. The camera’s strobe has made the “eggs” appear paler
than they actually are. Note how rim of nest is circular. Right: Cyathus olla growing on wood. Note
how rim of nest is often wavy. (Nancy Burnett)

Cyathus stercoreus (Dung-Loving Bird’s Nest Fungus)

FRUITING BODY tiny, goblet- to vase-shaped or resembling an inverted cone, 5-10(15)
mm high and 4-10 mm broad at the top, the rim more or less circular when mature and
covered at first with a thin pallid or whitish lid that soon disappears. Peridium (wall of
nest) tough, persistent, 3-layered (but the layers not always distinguishable). Exterior
tan to golden-brown, brown, grayish-brown, or reddish-brown and shaggy (but often
smooth and blackish in old age); base often with a pad of brown to reddish-brown
mycelium. Interior of the nest smooth, pale gray becoming dark gray or lead-colored and
often blackish in age. PERIDIOLES (eggs) several, 1-2 mm in diameter, dark gray to
black, flattened or lentil-like, hard, smooth, often with a short cord or “stalk” attached
(especially the lower eggs in the nest). Spores large: 22-40 * 18-30 microns, variable in
size and shape but mostly round to oval, thick-walled, smooth, hyaline.
HABITAT: Densely gregarious on dung, manure, and other organic debris; widely dis¬
tributed and common, but seldom noticed. In our area it can be found most any time. The
largest fruiting I’ve seen was on a well-manured lawn in March.
EDIBILITY: A meager morsel, much too puny to be of value.
COMMENTS: The dark gray to black eggs, shaggy exterior (at least when young), and
smooth interior are the principal fieldmarks of this cosmopolitan bird’s nest fungus. The
cords by which the eggs are attached are not always evident, at least in my experience.
Other species: C. olla is a similar but slightly larger species. Its nest is gray to brown with
an often wavy (rather than circular) rim and a smooth to finely hairy (fibrillose) but not
shaggy exterior. Its eggs are also larger (2-3 mm) and gray to brown or blackish, and its
spores are much smaller (8-15 microns). It is widely distributed and common, but not as
numerous in our area as C. stercoreus. C. pygmaeus is a miniature version of C. olla
(fruiting body only 4-5 mm high) that commonly grows on sticks, dung, etc., in the drier
parts of the West. Two other species are worth mentioning because of their beautifully
pleated or grooved (radially striate) interiors: C. slrialus, widely distributed and quite
common, with a shaggy cinnamon-brown to grayish-brown or dark brown exterior plus
slightly triangular eggs; and C. helenae, found mostly in arid or alpine habitats, with
a grayer, thicker-walled nest that has tufted hairs on its exterior. The latter two species
can grow in manure but are more common on sticks and other vegetable matter.

Nidula Candida (Jellied Bird’s Nest Fungus)

FRUITING BODY tiny, cylindrical or cushion-shaped becoming mug- or flower-pot¬
shaped at maturity (with vertical sides and a flaring mouth); 5-15 (20) mm high and 3-8
mm broad at the top when mature, the rim more or less circular and covered at first by a

780
Nidula niveotomentosaisee comments under N. Candida), young specimens in which the nest is still
covered by the epiphragm(“lid”). Note white exterior and mug-shaped fruiting body. (Herb Saylor)

lid. Peridium(wall of the nest) tough, persistent. Exterior whitish beneath a gray to brown
or dull cinnamon scurfy or shaggy layer that covers at least the basal portion and the lid.
Interior of the nest smooth, white to yellowish-brown or brown. PERIDIOLES (eggs)
several, 1-2 mm indiameter, pallid to gray or brown (but often darker on underside),
flattened, without cords, instead imbedded in a sticky mucilage or gel which eventually
dries out. Spores 6-10 * 4-8 microns, elliptical to nearly round, smooth, hyaline.
HABITAT: In groups on rotting wood, berry canes, and herbaceous debris in gardens,
woods, along streams, etc.; widely distributed. It is common in our area in the late fall
and winter or even spring, but the empty nests persist for months without decaying and
sometimes give rise to new ones.
EDIBILITY: Academic; you’d have to be slightly looney to bother with something so puny.

COMMENTS: A common but frequently overlooked little fungus, easily told by the
brownish to cinnamon scurfy exterior and the sticky gel in which the eggs are imbedded,
plus the presence of a covering or lid when young. The species name, Candida, which
means “shining white,” is a misnomer since the fruiting body is neither shiny nor white.
However, a similar species witha shaggy white exterior (and numerous small browneggs),
N. niveotomentosa, is common in California and the Pacific Northwest, usually on sticks
or in moss. See also the genus Nidularia (in the key) and Cyathus stercoreus.

Sphaerobolus stellatus (Cannon Fungus; Sphere Thrower)

FRUITING BODY minute, 1-3 mm broad, at first more or less round and white to dull
yellow-orange or ochraceous, the outer wall (peridium) then splitting into 4-9 bright
orange starlike rays or “teeth,” exposing the single spore-containing peridiole (“egg”),
which is then shot out like a cannonball as the entire structure turns inside out (leaving
behind a translucent whitish ball perched on the rays). PERIDIOLE (“egg”) chestnut-
brown to olive-black, sticky or slippery, smooth, more or less round. Spores 7-10 * 3.5-5
microns, oblong, smooth, hyaline.
HABITAT: Gregarious on rotting wood, sawdust, plant debris, and dung or manure;
widely distributed, but easily overlooked. I have seen it several times in nursery flats; it
fruits whenever moisture is sufficient.
EDIBILITY: Unknown. Several hundred would be needed for a mouthful!
COMMENTS: This cousin of the bird’s nest fungi is easily recognized by its diminutive
dimensions, bright orange star-shaped “catapult,” and single central “egg” or spore ball.
It makes a marvelous—albeit liliputian—laboratory pet: a small pop is said to accom¬
pany the ejection of the spore ball, which sticks to whatever it lands on. Its flight path
has been measured at 14 ft. high and 17 ft. long—or more than 1000 times the size of the
fruiting body! To equal such a prodigious feat, we puny humans would have to throw a
discus over one mile high and far! It is also interesting to note that the “sphere thrower”
does not “do its thing” in complete darkness—light is apparently needed to trigger the
“cannon,” as well as a sufficient supply of moisture.

781
782

Ascomycetes

ASCOMYCOTINA
THE Ascomycetes are the largest subdivision of the true fungi. With over 15,000 known
species, they pose a challenge, in the words of one noted mycologist, to “an army”
of students. They are also an exceedingly diverse group, ranging from the most econo¬
mically important of all the fungi, the single-celled yeasts, to powdery mildews to bread
molds like Penicillium (the original source of penicillin) to prized edible mushrooms such
as the morels and truffles. Their common bond is fundamental but microscopic: the
sexually-produced spores are formed inside saclike mother cells called asci (singular:
ascus). The ascus is roughly analagous to the basidium of a Basidiomycete, although
fusion of the parent nuclei typically occurs in a separate cell, the ascogonium. Each ascus
contains one to several thousand spores depending on the species. The most frequent
number is eight.
Only a small fraction of the Ascomycetes have fruiting bodies large enough to merit
mention in this book. In the field they can usually be distinguished from Basidiomycetes
by a process of elimination, i.e., if your fleshy fungal fructification does not fit one of the
common categories of Basidiomycetes pictured on pp. 52-54 (agarics, boletes, puffballs,
etc.), then chances are it’s an Ascomycete. The fleshy Ascomycetes treated in this book
fall into two broad categories, keyed below.

Key to the Ascomycetes

1. Fruiting body more or less round (spherical) to oval or knobby (potato-like), growing under¬
ground or inside very rotten wood . Discomycetes, p. 783
1. Not as above; growing on wood or on ground, on insects, other mushrooms, plants, etc. . . 2
2. Growing on wood (but wood sometimes buried) . 3
2. Growing on ground or on insects, herbaceous plants, or other mushrooms .4
3. Fruiting body usually black or very dark brown (but often covered with white or grayish pow¬
der), rounded to irregularly knobby and charcoal-like or fingerlike to clublike or antlerlike
and very tough or hard; asci borne in flasklike nests (perithecia) which often give the fertile
area of fruiting body a minutely pimpled appearance .Pyrenomycetes, p. 878
3. Fruiting body cuplike or variously shaped but not as above, or if colored as above then texture
usually different (fragile, fleshy, rubbery, gelatinous, etc.); asci typically borne in a palisade
(hymenium), not in perithecia . Discomycetes, p. 783
4. Growing on insects (adults, pupae, larvae), spiders, or on other mushrooms (but hosts are often
buried, so dig up carefully!); asci typically borne in flasklike “nests” (perithecia) .
.Pyrenomycetes, p. 878
4. Growing on ground or plant material, but not on insects or other mushrooms; asci typically
borne in a palisade (hymenium), not in perithecia . Discomycetes, p. 783

Left: Fruiting body of a cup fungus (a

common type of Discomycete), showing
how the paraphyses (sterile cells) and asci
are arranged in a palisade on the upper
(inner) surface. In reality there are thou¬
sands more asci than shown here. Right:
An ascus containing eight spores (the most
common number).
 783

DISCOMYCETES
DISCOMYCETES are not Ascomycetes that like to dance, nor are they necessarily disc¬
shaped. Discomycetes are Ascomycetes in which a palisade of asci line an exposed surface
of the fruiting body much as a palisade of basidia line the gills of an agaric. This palisade
of asci (see drawing at bottom of p. 782) is called the hymenium, and the fruiting body of
a Discomycete is called an apothecium (as opposed to the perithecium of a Pyreno-
mycete). All but a handful of the Ascomycetes treated in this book are Discomycetes.
Examples are the morels, false morels, elfin saddles, cup fungi, and earth tongues. Truffles
are also Discomycetes, but they have specialized underground fruiting bodies in which
the hymenium is infolded and internalized, or in some cases, non-existent (see p. 841 for
more details). The Discomycetes have been divided here into three orders (large groups),
keyed below. To facilitate identification, the key is based largely on the shape and size
of the fruiting body rather than on more critical(i.e., microscopic) characteristics. Excep¬
tions to the generalizations put forth in the key are then keyed out individually under
the various orders, families, and genera.

Key to the Discomycetes

1. Fruiting body nearly always underground (or inside very rotten wood), round to oval or knobby
(potato-like), i.e., stalk absent or extremely rudimentary; spores nearly always borne inside
the fruiting body, the interior usually (but not always) with channels, veins, or one or more
cavities .Tuberales, p. 841
1. Not as above; fruiting body occasionally buried but usually above the ground at maturity or on
wood, moss, etc.; spore-bearing surface exposed (external) at maturity . 2
2. Fruiting body cup- to ear-shaped, spoon-shaped, disclike (flat), cushion-like, top-shaped, or
sometimes contorted; stalk absent or present only as a narrowed base (but fruiting body some¬
times growing erect like a rabbit’s ear); asci operculate (i.e., with “lid” at tip) Pezizales, below
2. Fruiting body erect, with a stalk and often a cap, cup, or “head” . 3
3. Fruiting body with a well-defined cap or splitting into rays at maturity; cap cuplike, disclike,
wrinkled, brainlike, saddle-shaped, pitted, honeycombed, or thimble-like (i.e., usually with
a sterile underside or sterile hollow interior) . 4
3. Not as above; fruiting body clublike or with an enlarged or flattened “head,” but lacking a
distinct cap with a sterile underside; “head” not cuplike, brainlike, saddle-shaped, disclike,
or honeycombed, and not splitting into rays; fruiting body usually small; asci inoperculate
(i.e., tip usually thickened, with a pore but no “lid”) . Helotiales, p. 865
4. Flesh gelatinous or semi-gelatinous (slice open fruiting body lengthwise); surface of fruiting
body often viscid; cap rounded or wrinkled, often brightly colored but not dark brown to
black; asci inoperculate . Helotiales, p. 865
4. Not as above; asci operculate . Pezizales, below

Morels, Elfin Saddles, and Cup Fungi

PEZIZALES
THIS large order includes most of the familiar fleshy Ascomycetes and nearly all of the
prominent ones: the fabulous morels, grotesque false morels, elegant elfin saddles, and
lowly cup fungi. All of these fungi typically have an exposed spore-bearing surface (hy¬
menium) and their asci have “lids” which open when the spores are forcibly expelled.
The asci usually discharge their spores simultaneously. The discharge can easily be
triggered by tapping, breathing on, or otherwise disturbing the fruiting body, with the
result that it will sometimes “smoke” (spew out clouds of spores) when picked!
784 PEZIZALES

The fertile surface ranges from concave or flat in the cup fungi to convex or convoluted
in the false morels and elfin saddles to deeply pitted in the true morels. A stalk is present
in the latter forms but often absent in the cup fungi.
Members of the Pezizales can be found almost anywhere at any time, but are most
numerous and diverse in forests, in the spring. This means that in regions with mild winters
they usually peak after most of the Basidiomycetes have fruited, while in regions with cold
winters they are among the first fungi to appear after the snow has melted. Excluding the
truffles (which are treated separately in this book), there are seven major families in the
Pezizales, five of which are cup fungi. These are keyed below.
Key to the Pezizales
1. Fruiting body an irregularly cabbage-like, cauliflower-like, contorted, brainlike, or pitted mass
of tissue, with or without a stalk; flesh not gelatinous . 2
1. Not as above .. 3
2. Stalk absent or rudimentary, or if stalk present then stalk long (15-30 cm!) and solid and brown
and often buried and restricted to eastern North America; fertile portion of fruiting body
whitish to yellowish-brown, beige, pinkish, or lilac-tinged; rare . Pezizaceae & Allies, p. 817
2. Not as above; stalk present, long or short, usually hollow or partially hollow or complexly
folded or chambered in cross-section; fertile portion of fruiting body colored as above or
often darker; very common and widely distributed . 5
3. Fruiting body cup-shaped (concave) to disclike (flat), cushion-shaped, or sometimes top¬
shaped or splitting into rays; stalk present or absent; flesh gelatinous, fragile, or tough ... 4
3. Fruiting body with a cap and stalk; cap convex to conical to bell-shaped, round, lobed, brainlike,
saddle-shaped, or pitted but not cuplike (or cuplike only when very young); flesh fragile or
tough but not gelatinous. 5
4. Stalk absent, or if present then often (but not always!) short or merely a narrowed, downward
extension of the cup; stalk when present usually lacking distinct ribs; fruiting body fleshy,
fragile, rubbery, or gelatinous, sometimes brightly colored, large to minute; tips of asci amy¬
loid or not amyloid .Pezizaceae & Allies, p. 817
4. Stalk present, clearly differentiated from cap; stalk usually longer than width of cup or if shorter
then usually ribbed or fluted; fruiting body fleshy or fragile but not gelatinous and not brightly
colored, large to fairly small but not minute; tips of asci not amyloid . . Helvellaceae, p. 796
5. Cap honeycombed with ridges and pits, the pits usually fairly deep, or if not then the ridges
with a strong vertical orientation; cap intergrown with stalk (or in one case, only upper part
of cap intergrown with stalk) .Morchellaceae, below
5. Not as above . 6
6. Flesh gelatinous; fruiting body often viscid and brightly colored .... (see Helotiales, p. 865)
6. Not as above; flesh not gelatinous . 7
7. Cap roughly conical to bell-shaped, like a thimble on a finger (i.e., attached only to very top of
stalk, the sides hanging down freely like a skirt) .Morchellaceae, below
7. Not as above; cap lobed, brainlike, saddle-shaped, etc. and/or attached to stalk differently
.. Helvellaceae, p. 796

Morels and Allies

spores
MORCHELLACEAE
THIS is a small but famous family with only three genera: Morchella, Verpa, and Disciotis.
Morchella (the true morels or “sponge mushrooms”) has a pitted cap that is intergrown
with the stalk (see p. 785 for more details). Verpa also has a well-developed stalk, but
differs in having a smooth to wrinkled or somewhat pitted, thimble-like cap. Disciotis
lacks an obvious stalk and is likely to be mistaken for a veined Peziza; consequently it is
keyed out under the cup fungi. All three genera have non-amyloid asci and smooth ellip-
MORCHELLACEAE 785

tical spores. The spores lack large oil droplets but typically have “crowns” of minute
droplets at both ends.
Key to the Morchellaceae
1. Fruiting body cuplike or spreading (flat); stalk very short or absent .Disciotis, p. 796
1. Not as above; fruiting body with a cap or “head” and well-developed stalk . 2
2. Cap with pits and ridges; at least the upper part of cap intergrown with stalk Morchella, below
2. Cap smooth to wrinkled or shallowly pitted, sitting on the stalk like a thimble (i.e., attached only
to very top of stalk, the sides hanging free like a skirt) .Verpa, p. 793

MORCHELLA (Morels)
Medium-sized to large, mostly terrestrial fungi. FRUITING BODY with a cap and stalk; interior
hollow. CAP honeycombed with ridges and pits (chambers), attached to the stalk for all of its length
(or in one case, for 1 /3-2/3 of its length). STALK well-developed, smooth to scurfy or wrinkled
below. SPORES typically elliptical, smooth, without large oil droplets. Asci lining inside surfaces
of pits; 8-spored, operculate, not amyloid.

THESE prized delicacies are among the best-known wild mushrooms in North America,
and they are so esteemed in Europe that people used to set fire to their own forests in hopes
of eliciting a bountiful morel crop the next spring! Luckily, morels are among the most
unmistakable of all fungi by virtue of their hollow, pitted or honeycombed “heads” (they
are also known as “sponge mushrooms”). False morels (Gyromitra) are vaguely similar,
but have a wrinkled or convoluted (not pitted) cap, while thimble morels (Verpa) have a
smooth to wrinkled or shallowly pitted cap with free (skirtlike) sides. Morels have also
been confused inexplicably with stinkhorns (Phallus), perhaps because of their similar
shape. The head of a fresh Phallus, however, is coated with a sticky, stinky spore slime and
there is a sack or volva at the base of the stalk.*
Much has been said about where and when to find morels, but no self-respecting morel
hunter will divulge any truly crucial information unless he or she is planning to per¬
manently leave the country (and then only for a stiff price!). Morel hunters are so
protective of their favorite “patches,” in fact, that they regularly disseminate misleading
—if not downright erroneous—information, and they practice a presidential evasiveness
when asked: “Where did all those morels come from?” Thus any “tips” or “secrets” you
manage to squeeze out of morel hunters should be taken with a grain (better make that a
bucket!) of salt. Having said this, I will now tell you where and when to find morels . ..
Morels usually grow outdoors: in forests (under both hardwoods and conifers) and open
ground, in abandoned orchards, gardens, landscaped areas, under hedges, on roadcuts
and driveways, near melting snow, in gravel, around wood piles or tree trunks, and in
sandy soil along streams. In other words, morels grow wherever they please! They will
even fruit in barbecue pits and on scorched ground in the wake of forest fires—sometimes
in awesome quantities, but only if conditions are favorable!
Morels are almost universally associated with the spring (“May is morel month in
Michigan”), but occasionally appear in the summer, fall, and winter. In the Midwest,
where they are particularly abundant, the various species appear in a definite succession
from late April through early June, and the annual morel festivals are a major tourist
attraction. The state of Minnesota, in fact, recently declared the morel its official “state*
mushroom.”' Morels seem to respond to a warming trend following a cold spell. This
explains why they are particularly abundant in regions with cold winters—which is only
right, because the people who survive those winters deserve a delicious spring! Alas, in
tepid coastal California morels are not nearly as common as one would wish. In our area,
in fact, finding them is largely a matter of luck, i.e., being in the right place at the right

* Don’t feel too embarrassed should you manage to confuse them—Morchella esculenta was originally classified
as a Phallus by Linnaeus!
Springtime bounty: A basket of black morels (Morchella elata group, p. 790).

time. In the Sierra Nevada and Cascades, however, they can actually be hunted—and
sometimes harvested in large quantities, particularly if there are spring rains. Although
timing is of critical importance because of competition from other morel hunters, it has
been shown that morels develop and age more slowly than most mushrooms, over a period
of three to four weeks! They have resisted all attempts to raise them commercially, but one
ambitious entrepeneur tells me that success isjustaround the corner. The spores germinate
readily in culture and the mycelium flourishes, but getting it to fruit has been the major
stumbling block (perhaps it needs to be chilled to simulate a cold winter?).
Although the genus Morchella is unerringly distinct, the disposition of species is largely
a matter of opinion. Dozens have been described based on differences in color, shape, and
orientation of the pits and ridges. However, each “species” seems to intergrade with the
next, leading some morelizers to recognize only three or four species. To most people the
“true” identity of a morel is academic anyway—what counts is that it is edible, and
incredible!
Raw morels often cause digestive upsets, so to stay on their good side, always cook them.
They are delicious sauteed (providing you don’t drown them in butter and herbs), and you
can serve them over toast or in soup. Because they are hollow, morels are easy to stuff,
but it is even easier to stuff yourself with morels! Before cooking, always split them length¬
wise to check for millipedes, slugs, and other critters that like to hide inside. Transfer all
such tenants to the compost pile or some other comfortable spot (just because you’re
evicting them from their home doesn’t mean you have to kill them!) and carefully remove
all grit or sand, using water if necessary. If you are lucky enough to stumble on a large
batch of morels and are unwilling to share the surplus with me, you can preserve them by
canning or sauteeing and freezing. Or you can string them into beautiful necklaces and
hang them up to dry (I will accept them in any form).
Four “species” of Morchella are described here, and several others are discussed. As
I am meticulously mapping their variation and distribution, I request you, generous
reader, to deliver any and all morels you find to my doorstep. Then, while I am savoring
the superb flavor of croutes aux morilles a la normande (having first, of course, studied
them thoroughly), you can bask in the altruistic satisfaction that comes from contributing
to science.*
*The least you can do is invite me over for dinner.

786
MORCHELLA 787

Key to Morchella
1. Volva (sack) present at base of stalk (dig up carefully!); odor often fetid (see Phallales, p. 764)
1. Volva absent . 2
2. Cap usually brainlike and irregularly lobed (as well as pitted), brown to reddish-brown; stalk
complex (i.e., cross-section half way up stalk revealing numerous internal folds), often massive
(up to 20 cm thick!) .(see Gyromitra, p. 799)
2. Not as above; cap not conspicuously lobed; upper stalk or mid-portion more or less simple in
cross-section (but base often folded or complex) . 3
3. Lower 1 / 3-2/3 of cap free from stalk (only the upper part intergrown) M. semilibera, p. 791
3. All or nearly all of cap intergrown with stalk (lower edge may be creased).4
4. Ridges of cap dark (dark gray to olive-brown to black) at maturity and sometimes dark when
young . M. data group, p. 790
4. Not as above (but ridges may blacken when they dry out and shrivel up) .5
5. Ridges white or markedly paler than the pits when young, but often becoming same color as
pits in age; pits usually large and often elongated vertically; fruiting body small to medium¬
sized (not large); common in suburbia, orchards, etc., less often in woods M. delidosa, p. 789
5. Not with above features (but may have some of them) .6
6. Fruiting body pale (whitish to buff), not darkening in age; pits usually vertically elongated;
mostly found in woods (especially montane) M. sp. (unidentified) (see M. deliciosa, p. 789)
6. Not as above; if pale then pits roundish to irregular . 7
7. Fruiting body medium-sized to large (11-30 cm tall or more), not reddish-tinged; stalk often
swollen at base and sometimes massive in age; base usually wrinkled, folded, or buttressed
(like a tree trunk); found mainly with hardwoods . . . M. crassipes(see M. esculenta, below)
7. Not as above; fruiting body usually small to medium-sized or reddish-tinged; stalk often
somewhat wrinkled at base but not normally buttressed; found in many habitats.8
8. Fruiting body reddish to reddish-brown or with a reddish tinge and/ or pits arranged in definite
rows . M. data group, p. 790
8. Fruiting body not reddish-tinged; pits usually rounded or irregular .... M. esculenta, below

Morchella esculenta (Morel; Yellow Morel) Color Plate 203

CAP 3-11 cm high, 2-6 cm broad, round to oval to bluntly conical or irregular; margin
attached to the stalk (but often with a crease at point of attachment); overall color tan to
yellow-brown to warm buff or even buff. Pits roundish to irregular in shape and not nor¬
mally arranged in well-defined rows; often quite small but sometimes large, yellowish
to brown or tan. Ridges typically meandering rather than in lines, usually quite narrow,
same color as pits or paler (or occasionally slightly darker). Interior hollow, whitish,
roughened. Flesh often rather thin. STALK 1-5 (10) cm long, 1-2.5 (3.5) cm thick, usually
minutely granular or scurfy, equal or enlarged at the base, usually relatively short and
narrower than cap, the base often somewhat wrinkled or pitted; white to buff, sometimes
with brownish or cinnamon stains; typically hollow in cross-section. SPORES 16-25
x 9-14 microns, elliptical, smooth, without oil droplets.
HABITAT: Solitary to widely scattered, gregarious, or clustered in a variety of habitats
(woods, streamsides, old orchards, cultivated or disturbed ground, burned areas, etc.);
very widely distributed, but especially common in eastern North America and the Mid¬
west. Like other morels, it fruits in the spring but will occasionally turn up at other times.
In eastern North America it is most often found under oak, maple, beech, hickory, elm,
ash, fruit trees, and other hardwoods, especially in May (or in the words of Ingrid Bartelli,
“when the oak leaves are as big as squirrel’s ears’’). Large crops can also be found around
the bases of dying (but not quite dead) elms attacked by Dutch elm disease. In the West I
look for it under deciduous oaks and in sandy soil or riverbottoms where there is willow,
cottonwood, or alder, from February to May or June depending on the elevation
and climate. It also occurs in burned areas, but not as commonly as the black morels, and
it is not as frequent as the latter at higher elevations.
Left: Morchella crassipes (see comments below). This morel varies considerably in shape but is
always large and has a wrinkled, buttressed stem. A sliced specimen is shown at top of p. 792. Right:
Morchella deliciosa (p. 789), young specimens from eastern North America with very white ridges.
The form in coastal California is not usually as white (see photo on p. 789). (Alan Bessette)

EDIBILITY: Edible and one of the most avidly hunted of all wild mushrooms (see
comments on pp. 785-786).
COMMENTS: This, the common morel or “sponge mushroom,” is one of the most readily
recognized of all edible fungi. It can be told from other morels by its modest size, warm
brown to tan or yellowish color, and irregularly-arranged pits (see color plate), and from
the poisonous false morels (Gyromitra species) by its pitted or honeycombed rather than
brainlike or lobed cap. It could also be confused with the stinkhorn (Phallus impudicus),
but does not have a volva and lacks the foul-smelling spore slime of that species. It is typi¬
cally a denizen of low hardwood forests, riparian woodlands, and fruit orchards, whereas
the black morels are more common in northern latitudes or at higher elevations under
conifers. The ranges of the two overlap, however, and they can sometimes be found
growing in the same area. Like other morels, M. esculenta varies considerably in shape
and appearance. In most cases the pits are quite irregular-looking, but in some forms
the transverse ridges are not as well-developed as the vertical ones, giving the pits an
elongated, slotlike appearance. The ridges never blacken as in the M. data group, but
conical forms intergrade with M. deliciosa (see comments under that species). Pale
forms can also be confused with M. deliciosa, but usually have smaller, paler pits that
are not as vertically elongated. A giant morel with irregularly arranged pits occurs in
the same habitats as M. esculenta, but usually a little later. This is M. crassipes (or
M. esculenta var. crassipes). It measures 6-20 or more cm high and usually has a massive
stalk with an enlarged, wrinkled and folded or buttressed base (hence its nicknames,
“Thick-Footed Morel,” “Big-Foot,” and “Tree Trunk Morel”). It also tends to have
grayer pits when young with paler or even whitish ridges, but like M. esculenta, becomes
tan in age. Many morel connoisseurs in eastern North America prize it above all others.
In the West, unfortunately, it is relatively rare. I have found it under cottonwood in
Oregon, among forget-me-nots in an old orchard in California (see photo above), and
in landscaped areas.

788
MORCHELLA 789

Morchella deliciosa (White Morel) Color Plate 201

CAP 1.5 -6 cm high, 1-3.5 cm broad, round to oval or conical, the margin attached to the
stalk but sometimes creased at its juncture. Pits usually vertically elongated and quite
large at maturity, but not necessarily arranged in rows; dark gray to dark brown to grayish-
tan, brown, or tan (usually darker when young and paler in age). Ridges usually widely
spaced and mostly vertical (the horizontal ones often few or not as prominent), white
to creamy (lighter than pits) when young, usually becoming tan or same color as pits
in age but not blackening unless shrivelled up. Interior hollow, the surface pallid or
whitish, roughened. Flesh rather thin. STALK 1.5-6 cm long, 0.5-2 (3.5)cm thick, equal or
thicker below, the base often somewhat wrinkled, pitted, or irregular; white to creamy or
buff, usually scurfy or minutely warted; typically hollow in cross-section. SPORES
18-25 * 10-15 microns, elliptical, smooth, without oil droplets.
HABITAT: Solitary to gregarious in gardens and other suburban habitats, under fruit
trees, in old orchards, in woods and at their edges; etc.; widely distributed, fruiting mainly
in the spring. In eastern North America its appearance marks the beginning of the end of
the morel season. In coastal California it is not uncommon in the spring, especially in
sandy soil, but occurs practically year-round. One couple I know gets a small crop under
their plum tree every Christmas! Another “white morel” occurs in the Sierra Nevada and
Cascades (see comments).
EDIBILITY: Delectably delicious, as the species epithet implies. Like all morels, it
should be cooked (see comments on pp. 785-786 for more details).
COMMENTS: As already pointed out, morels are perplexingly polymorphic and resist
our obtrusive attempts to categorize them. The name M. deliciosa has been applied to
more than one kind of morel, and the above description has been broadened to include a
number of confusing and intergrading, small to medium-sized morels with white or pallid
ridges at least when young and pits which are usually large and often vertically elongated.
In age the pits and ridges often become the same color(yellowish-tan to brown) as shown
in the color plate, but the shape and size of the pits helps distinguish it from M. esculenta.
The name M. conica has also been used for this species, but is more properly applied to
morels with blackening ridges (see the M. elata group). Other species: Another “white
morel” occurs in the S ierra Nevada and Cascades about the same time as the black morels.
It has a whitish to buff cap that does not darken appreciably in age. The pits are usually
elongated vertically but the ridges often form vertical lines as in the black morels. I do
not know the identity of this morel, but 1 do know that it is delicious!

The coastal Californian form of Morchella deliciosa (or what I have identified as that species). Note
how the ridges are whitish when very young but become colored like the pits in age. Also note how
large and elongated the pits are, and how the ridges on dried-up specimen at left have darkened. Two
mature specimens are shown in the color plate.
Black morels (Morchella elata group). This narrow-headed variety has also been called M. angusti-
ceps and M. conica(see comments on p. 791). Note the conical head and strong vertical orientation of
the ridges. They were found in a mountain meadow—a favorite haunt of the narrow-headed variety.

Morchella elata group (Black Morel) Color Plates 199, 202

CAP 2-6 (10) cm or more broad and 2-10 (18) cm or more high; usually conical to oval or
somewhat irregular in shape; margin joined to the stalk but often with a crease at its point
of attachment; overall color usually quite dark, especially in age. Pits usually vertically
elongated and/ or arranged in vertical rows, but in some forms meandering (see com¬
ments); yellow-brown to brown, grayish, olive-brown, or even reddish-brown, sometimes
becoming blackish in age. Ridges vertically aligned in some forms, colored like the pits
at first, usually darker (olive-brown to smoky-brown to black) before or by maturity
(and sometimes dark from the beginning). Interior hollow, the surface pallid or
whitish or tinged cap color, roughened. Flesh fragile in some forms. STALK 1.5-10 (20)
cm or more long, (0.5) \A cm or more thick, slender in some forms, as thick as the cap in
others (often proportionately longer and thicker in age); equal or enlarged at either end,
often grooved and/ or wrinkled, especially toward base; sometimes nearly smooth, more
often entirely or partially scurfy from a coating of small granules or warts; white to creamy,
buff, pinkish-tan, or sometimes with a reddish tinge; typically hollow in cross-section.
SPORES (18) * 20-25 (30) * 11-15 microns, elliptical, smooth, without oil droplets.
HABITAT: Solitary, scattered, or in groups, clusters, or troops on ground in woods and
at their edges (especially under conifers and aspen) and in burned areas, less commonly
in urban and suburban settings; very widely distributed (Japan to the Himalayas, Europe,
etc.), but especially common in northern and western North America. Like other morels,
its season is the spring(or early summer at higher altitudes), but“freak” fruitings can occur
in the fall, particularly if a warm wet spell follows cold weather. In coastal California,
where the seasons are not strongly defined, black morels occur year-round, but seldom in
quantity. In colder regions they are often abundant, usually two to five weeks after the
snow melts. Look for them in the Sierra Nevada when the snow plants (bright red sapro¬
phytic plants) have fully emerged, and in the Cascades when the calypso orchids are in
bloom (see Color Plate 199). Don’t expect to find them everywhere, however. For one
thing, you face fierce competition from other collectors; for another, they are devilishly
difficult to see, because they look just like fallen pine or fir cones (see Color Plate 202).
Also, they show a definite preference for semi-disturbed areas, e.g., campgrounds, along
roads, and in logged and burned areas. Bushels can be harvested one to two years after a
forest fire, providing the spring weather is favorable.
EDIBILITY: Edible and delectable (see comments on pp. 785-786). However, some
people are apparently “allergic” to it. Like all morels, it should never be eaten raw.

790
mm

Black morels (Morchella elata group). Left: This fat-headed specimen has the radially-arranged pits
of the narrow-headed variety (see comments below). Right: Typical examples of the fat-headed form.

COMMENTS: The above description covers a number of black morels (the so-called
“M. elata-M. angusticeps-M. conica” complex). Many of the morels in this category
are black or at least dark in age, but some are brown, reddish, or pinkish-tinged and/or
have pits that are usually more or less radially arranged (that is, arranged in rows).
Others are quite dark and may have pits that are not arranged in obvious rows. Excluded
are those morels with a “half-free” cap (see M. semilibera), brown or tan to yellowish-
capped morels with irregularly arranged pits (see M. esculenta), and morels with
dark pits and pale ridges when young (see M. deliciosa). There are at least two common
kinds of black morels in North America. The first variety, sometimes called the “Fat-
Headed Black Morel,” is probably the “true” M. elata, though it has gone under the name
M. angusticeps and has also been called M. conica. It has a roundish to oval or conical
“head” that is relatively large in comparison to the stalk, pits which are often elongated
but not necessarily radially arranged, and ridges that are gray to black before maturity
(and are often dark while very young). This variety is shown above and in the color plates.
The second variety, sometimes called the “Narrow-Headed Black Morel,” is shown in
the photo on p. 790. Its cap is often more conical or elongated and often spongier or
more fragile than the first variety. Its pits are usually arranged in rows and its ridges usually
blacken, but not as quickly or as consistently as the fat-headed variety. Its stalk is often
long and relatively thick (often as thick as the base of the cap), and is typically more
furrowed or wrinkled toward the base than the fat-headed variety, and also wartier or
scurfier. According to “morelizers,” this second variety, if distinct from the first, is
probably the “true” M. angusticeps, but it has also been called M. conica. A giant form of
this variety is shown on the back cover of the book.
These two black morels grow in the same regions, but the fat-headed one seems to be
the commoner of the two. Sometimes they seem distinct, at other times they intergrade or
“hybridize.” A third variety, often called the “Red Morel,” is shown on p. 31. It resembles
the previous two (especially the narrow-headed one), but a distinct reddish or pinkish
tinge pervades its stalk and/or cap. This variety, which may simply be a growth form,
has a particularly fine flavor. Still another variety has a whitish to buff cap. It may be an
albino form, or a separate species entirely (see comments under M. deliciosa). Whew!

Morchella semilibera (Half-Free M orel)

CAP l .5 -5 cm high and broad, bluntly conical to round or oval when young, usually conical
in age; margin (lower edge) free from stalk for about one half (1 / 3-2/3) the distance to the
apex of the cap, often flared outward away from the stalk in age. Pits large and elongated,
yellowish-brown to brown or grayish-brown. Ridges vertically oriented (with few if

791
Left: In most species of Morchella the cap is completely intergrown with the stalk, as shown in this
sliced M. crassipes. Right: In Morchella semi/ibera, however, the lower part of the cap hangs free
from the stalk. These are mature specimens.

any transverse ribs), yellowish-brown to brown or olive-brown (usually slightly darker

than the pits), often becoming blackish in old age or as they dry out. Underside (interior)
whitish or pallid, roughened. Flesh rather thin and fragile. STALK 3-10 cm long, 1-2.5
cm thick at apex, equal or thicker below in age, fragile, white to yellowish; surface usually
noticeably rough or with scurfy granules that may form ribs (as in Verpa)', usually hollow
in cross-section. SPORES 22-34 * 15-21 microns, elliptical, smooth, without oil droplets.

HABITAT: Solitary to gregarious on ground in woods and under trees (mainly hard¬
woods); widely distributed, fruiting in the spring. In our area it occurs in sandy soil along
streams with cottonwood and alder, but is not as common as Verpa bohemica, which
favors the same habitats. Farther north (e.g., in Idaho) it is more frequent.
EDIBILITY: Edible, but more fragile and not quite as flavorful as other Morchellas.

COMMENTS: Also known as M. hybrida and Mitrophora semilibera, this is a dis¬

tinctive species because of its “half-free” cap that is intermediate between Verpa bohemica
(whose cap is attached to the stalk only at its apex) and other Morchellas (whose cap is
completely intergrown with the stalk). It is also intermediate in its fruiting behavior, for
it typically appears shortly after the Verpas and shortly before the other Morchellas.
The stalk may be quite short when young and the “head” relatively large, but in age the
“head” shrinks and the stalk lengthens considerably, giving it the aspect of a Verpa.

Morchella semilibera, prime specimens. Note “half-free” cap and roughened stalk. Compare the cap
to the completely free, thimble-like caps of the Verpas (pp. 793-795).
MORCHELLA 793

The roughened stalk surface is, in the words of mycologist Orson K. Miller, “the texture
of a cow’s tongue.” Typical M. semilibera is unlikely to be confused with other morels,
but it sometimes intergrades with the black morel (M. elata group).

VERPA (Thimble Morels)

Medium-sized to fairly large terrestrial fungi. FRUITING BODY with a cap and stalk. CAP bell¬
shaped to conical and thimble-like, i.e.,free from the stalk except at the very top; surface smooth
to deeply wrinkled or sometimes pitted. Flesh rather fragile and thin. STALK well-developed,
smooth or roughened by granules but not fluted; usually stuffed with a pith. SPORES elliptical,
smooth, without large oil droplets. Asci lining outer surface of cap; asci 8-spored or 2-spored,
operculate, not amyloid.

A VERPA looks like a thimble stuck on a finger, i.e., its smooth to wrinkled or pitted cap
is attached only to the very top of the stalk so that its sides hang free like a skirt. The true
morels (Morchella), in contrast, featured a pitted cap that is entirely or partially inter-
grown with the stalk, while the false morels and elfin saddles (Gyromitra and Helvetia)
have lobed, brainlike, or saddle-shaped caps.
Verpas are edible, but are more fragile than the true morels and not as flavorful. At
least one species, V. bohemica, is poisonous to some people, so it is advisable to eat Verpas
sparingly if at all. You can find them in woods and under trees and shrubs, particularly in
well-drained soil along rivers and creeks. They are most abundant in the spring about a
week to a month before the true morels (they are sometimes called “early morels”), but in
coastal California they also occur in the winter. Two farflung species are depicted here.
Key to Verpa
1. Upper portion of cap intergrown with stalk; cap pitted . . . (see Morchella semilibera, p. 792)
1. Not as above; cap free from stalk except at very top (like a thimble on a finger) . 2
2. Cap strongly wrinkled, the wrinkles usually oriented vertically and sometimes branched to
form pits; asci 2-spored... V. bohemica, below
2. Cap smooth or at times irregularly wrinkled; asci 8-spored . 3
3. Stalk often rather spongy and usually stuffed with a cottony pith (but may be hollow in age);
spores without oil droplets . V. conica, p. 794
3. Not as above; spores usually with one oil droplet at maturity .(seeHelvetia, p. 805)

Verpa bohemica Color Plate 205

(Early Morel; Wrinkled Thimble Morel)
CAP 1 -5 cm broad and 2-5 cm high(but sometimes much larger—see comments), bluntly
conical to somewhat bell-shaped, squarish, or irregular, attached to the stalk only at
the apex, the sides hanging down freely like a skirt; margin often touching(but not joined
to!) stalk when young, sometimes flaring or upturned in old age. Fertile surface pale
to dark yellow-brown or tan, often becoming darker brown in age; deeply wrinkled by
branching folds or ribs which are often vertically oriented and sometimes branch to form
pits. Underside whitish to brownish. Flesh rather thin and fragile. STALK 6-15 cm long,
0.8-3 thick (or sometimes much larger), equal or tapered slightly in either direction;
whitish to creamy or becoming tan or ochre in age, smooth or often roughened by small,
orangish to brownish granules which may form transverse belts or “ribs”; more or less
round in cross-section, usually stuffed loosely with a cottony white pith. SPORES huge;
54-80 * 15-18 microns, elliptical-elongated, smooth, without oil droplets. Asci 2-spored.
HABITAT: Widely scattered to gregarious in woods, thickets, and forest edges, etc.,
especially in sandy or well-drained soil in stream valleys and ravines; usually fruiting in
the early spring, widely distributed. It typically appears one to three weeks before the true
 Verpa bohemica. Note how the cap is strongly wrinkled vertically, and may even be pitted (as shown
in old specimen at center). Note also how cap is attached only to very apex of stalk, and how the stalk
is stuffed with a cottony white pith, at least when young.

morels (Morchella species), sometimes in the same places. It is abundant in parts of the
Pacific Northwest. In our area I have seen large fruitings under cottonwood and alder.
EDIBILITY: Edible with caution. Although eaten by many people, it can cause severe
stomach cramps and loss of muscular coordination, particularly when consumed in large
amounts or on several successive days. The flavor is strong but not on a par with the true
morels. Always cook it, and beware: some of the “morels” sold in markets are Verpas!

COMMENTS: Also known as Ptychoverpa bohemica because of its two-spored asci,

the early morel is easily told by its conspicuously wrinkled or pitted, thimble-like cap and
smooth to roughened but not fluted stem. The cap is not as lobed as that of Gyromitra and
Helvetia, and the sides of the cap are not intergrown with the stalk as in Morchella. Med¬
ium-sized specimens are the norm, but as in some of true morels, a giant form also occurs.
Typical V. bohemica is most likely to be confused with Morchella semilibera, whose cap
is “half-free” (intergrown with the stalk over its upper portion), and with V. conica, which
has a smoother or more irregularly wrinkled cap and 8-spored asci. When in doubt, you
can quickly resolve the issue by examining the spores. Those of V. bohemica are the
largest of any stalked Discomycete!

Verpa conica (Thimble Morel; Bell Morel)

CAP 1 -4 cm broad and high, usually more or less thimble-shaped (broadly conical to bell¬
shaped), but sometimes lobed (see comments) and sometimes developing a depression at
the top in age; attached to stalk only at its apex, the sides free like a skirt; margin inrolled
or incurved at first and sometimes touching the stalk (but not joined to it!), often lobed,
often flaring out or turning up in old age. Surface smooth to slightly wrinkled, or in one
form irregularly wrinkled (see comments), ochre-brown to brown or dark brown. Under-
side pallid. Flesh thin and rather brittle or fragile. STALK (2.5)4-12 cm long, (0.4) 0.5-1.5
cm thick, equal or tapered upward (or occasionally downward), white to yellowish, tan,
or tinged ochre-orange; smooth or with granular or minutely scaly transverse bands or
ribs (granules often browner or oranger than background); often rather spongy and
usually stuffed with a loose cottony pith (but often becoming hollow in age); more or less
round in cross-section. SPORES (20) 22-30 (34) * 12-17 (19) microns, elliptical, smooth,
without oil droplets. Asci 8-spored.

794
 Verpa conica, typical specimens. Cap is relatively smooth and thimble-like, i.e., it is attached only to
the apex of the stalk and it pops off easily.

HABITAT: Solitary, widely scattered, or gregarious in soil or humus in forests, riparian

woodlands, under shrubbery and fruit trees, etc., usually in the spring; widely distributed
but not particularly common. In our area I have found it as early as February and as late
as May, but it usually peaks a week or two after V. bohemica. I have seen fairly large
fruitings under redwood, oak, and cottonwood; in southern California it sometimes fruits
in great quantity under chaparral shrubs. I have also seen it in Yosemite.
EDIBILITY: Edible when cooked, but rather fragile and not often occurring in enough
quantity to warrant collecting.
COMMENTS: The thimble-like cap with a free margin is the principal fieldmark of this
farflung fungus. A rather strongly wrinkled, irregularly lobed form sometimes occurs,
however (see photo below). The shape of the typical form is somewhat reminiscent of a
stinkhorn, but it lacks a volva and its cap is never coated with putrid spore slime. Since
the cap is attached to the stalk only at the very top, it pops off easily, and the entire mush¬
room is rather fragile. The strongly wrinkled form does not have the vertical ridges
characteristic of V. bohemica, and is actually reminiscent of a small Gyromitra. The
spongy stalk with a cottony pith inside plus the free cap margin that is inrolled when young
are sufficient to distinguish it. Morchella semilibera is also somewhat similar, but has a
pitted cap that is partially intergrown with the stalk.
Verpa conica, a gyrose (lobed and wrinkled) form (see comments above). The cap is reminiscent of a
small Gyromitra, but is attached only to the apex of the stalk and has a strongly inrolled margin at first.
Note how stalk is stuffed with a cottony pith. This form is quite different from the typical form shown
at top of this page, but is microscopically identical.
796 MORCHELLACEAE

DISCIOTIS
THIS genus includes one farflung species, described below, with a brown, more or less
cuplike fruiting body. Consequently, it is keyed out under the cup fungi. Microscopic
features, however, relate it to the morels.

Disciotis venosa (Veined Brown Cup Fungus)

FRUITING BODY (3) 5-20 cm broad, at first more or less cup-shaped, but usually
flattening out in age. Fertile (upper or inner) surface reddish-brown to brown, dark brown,
or at times ochre-brown, sometimes smooth when young but usually becoming radially
wrinkled or veined to reticulate, corrugated, or pebbled by maturity (at least toward the
center); margin often wavy and/ or splitting. Exterior scurfy, roughened, or minutely
warty, whitish to buff or tinged brown, often fluted or wrinkled at base (i.e., appearing
“gathered”) to form a short stalk. Flesh fairly thick and brittle. STALK when present
short, thick, usually buried. SPORES 19-25 (30) * 12-15 (17) microns, elliptical, smooth,
without large oil droplets; asci lining upper surface of fruiting body, not amyloid.
HABITAT: Solitary, scattered, or in small groups in damp soil or humus under or near
trees or occasionally in the open; widely distributed, fruiting mainly in the spring. I usually
find it on well-drained soil in riparian woodlands (cottonwood, willow, etc.), but in my
experience it is not very common.
EDIBILITY: Edible, but not recommended. Although the flavor is said to be like that of
morels, people without a microscope can easily confuse it with other brown cup fungi
(Peziza, Discina, etc.) which may or may not be edible. One source lists it as poisonous
unless cooked.
COMMENTS: U ntil it is seen several times, this prominent brown cup fungus can be dif¬
ficult to identify in the field. It is often quite large and conspicuously veined or reticulate,
but several other brown cup fungi can mimic it. Microscopically it is quite distinct, how¬
ever, for the smooth, elliptical spores lack oil droplets and the asci do not blue in iodine.
It used to be called Peziza venosa, but because of these microscopic differences it has
been rewarded with its own genus and placed in the morel family.

False Morels and Elfin Saddles spores

HELVELLACEAE
MOST of the fungi in this family have a well-developed cap and stalk, but a few are cup¬
like or disclike. In some species of Helvetia (the largest genus in the family), the cap is
concave (cup-shaped), with the hymenium (fertile tissue) lining the upper or inner surface
of the cup. In other Helvellas the cup turns inside-out as it matures and becomes lobed
or saddle-shaped, while in still others it is saddle-shaped or irregularly lobed from the
beginning. In Gyromitra the cap is often brainlike, but not deeply pitted as in the morels.
Microscopic features that unite the family include the non-amyloid asci and smooth to
roughened spores that typically contain one to three large oil droplets.
Several species of Gyromitra contain a deadly poisonous rocket fuel called MMH
(see p. 893). Although these species are usually harmless when cooked or dried, it is safer
to avoid them and to treat all members of the family with extreme caution (i.e., always
cook them, don’t eat large amounts, and be certain of your identification).
HELVELLACEAE 797

Key to the Helvellaceae

1. Fruiting body flattened and spreading (often undulating), attached to substrate by numerous
“roots” (rhizomes); stalk absent; fertile (upper) surface dark reddish-brown to brown or even
blackish, the margin often paler; growing under conifers, especially in recently burned areas;
widespread but not very common . Rhizina undulata (-R. inf lata)
1. Not as above; fruiting body differently shaped, or if similar then not attached by “roots” . . 2
2. Fruiting body shallowly cup-shaped to flattened, disclike, umbilicate, or with a slightly down-
curved margin; stalk absent, or if present then short and rather thick; fertile (upper) surface
smooth or wrinkled but not brainlike, yellowish to tan, brown, or reddish-brown but not nor¬
mally gray, grayish-brown, or black; underside lacking prominent hairs or ribs (but stalk may
be ribbed or base of fruiting body may appear “gathered”); found mainly in spring and early
summer when or soon after snow melts; spores often apiculate at maturity . . Discina, below
2. Not with above features (but may have some of them) . 3
3. Fruiting body an erect club or column that lacks a distinct cap, up to 12 cm tall; interior of
fruiting body with large chambers or compartments; exterior white to creamy or brownish,
usually fluted or furrowed lengthwise; found in eastern North America and Midwest, usually
under hardwoods; not common. U nderwoodia columnaris
3. Not as above; fruiting body with a cap (or cup) and stalk.4
4. Fruiting body small (cap up to 3 cm broad but usually less than 2 cm); cap usually round or con¬
vex, but sometimes compressed or broadly saddle-shaped, the margin usually tucked in toward
stalk; usually found in groups or clusters; spores often needle-like . . (see Helotiales, p. 865)
4. Not with above features; usually larger or differently shaped; spores not needle-like . 5
5. Stalk very thin (less than 3 mm); cap hollow and bladderlike, whitish to yellowish; found in eas¬
tern North America, usually clustered; spores borne on basidia (see Aphyllophorales, p. 548)
5. Not as above; spores borne on asci; widespread . 6
6. Cap (or “cup”) gray to black or white . 9
6. Cap (or “cup”) some shade of brown or tan . 7
7. Cap brainlike to irregularly convoluted or wrinkled (at least at maturity), or if saddle-shaped
then spores with two oil droplets .Gyromitra, p. 799
7. Cap saddle-shaped, lobed, cup-shaped, plane, etc., but the surface only slightly wrinkled if at
all; spores typically with one oil droplet . 8
8. Stalk arising from a swollen tuber like structure (sclerotium); cap cup-shaped to disclike; fruiting
body small .(see Helotiales, p. 865)
8. Not as above; stalk not normally arising from a sclerotium . 9
9. Fruiting body minute; cap usually cup-shaped or disclike and less than 1.5 cm broad; stalk very
thin (usually 1 -2 mm thick); growing mostly on plant tissues (living or dead); asci not operculate
(i.e., without “lids”) .(see Helotiales, p. 865)
9. Not as above; fruiting body small to fairly large, not minute; cap variously shaped (including
cuplike or disclike); growing in soil, humus, or rotten wood; asci operculate Helvetia, p. 805

DISCINA (Pig’s Ears)

Medium-sized fungi found on ground or rotten wood, usually in the spring soon after the snow
melts. FRUITING BODY shallowly cup-shaped to disc-shaped (flat), umbilicate, or slightly
convex, with or without a short stalk. Fertile (upper) surface yellowish to tan or brown{oi less
commonly reddish-brown), smooth to somewhat wrinkled; underside not normally ribbed.
STALK when present short and thick, usually ribbed orappearing“gathered.” SPORES elliptical
to spindle-shaped, smooth to roughened or reticulate, the ends typically pointed or with short
projections (apiculate). Asci lining upper surface of fruiting body, typically 8-spored, operculate,
not amyloid.

THESE are springtime cup- or disc-shaped fungi with a yellowish to brown fertile sur¬
face and a thick, short stalk or none at all. As such they are likely to be mistaken for cup
fungi (and are keyed out under that group), but microscopic features such as the non¬
amyloid asci and apiculate spores relate them closely to Gyromitra of the Helvellaceae.
Discina perlata. This common “snowbank” mushroom looks like a cup fungus, but usually has a
short stalk and is often umbilicate (i.e., with a “navel,” as shown here). The fertile surface can be
smooth or quite wrinkled. Microscopic features relate it to Gyromitra.

Pig’s ears (a common name also applied to Gomphus clavatus, a member of the chan¬
terelle family) are a common feature of our western springtime or“snowbank” mushroom
flora. They are said to be edible, but because of their close relationship to the poisonous
Gyromitras, it is best to eat them in moderation (if at all) and always cook them. The dozen
or so species of Discina are distinguished primarily on microscopic features such as the
shape and ornamentation of the spores. As only one is described here, a key hardly seems
necessary.

Discina perlata (Pig s Ears)

FRUITING BODY (2)3-10 (20) cm broad, at first cup-shaped but soon becoming saucer¬
like, then wavy or flattened (disclike) to very broadly convex (with the margin turned
down) or umbonate, or umbilicate (with a central depression); outline round to irregular
or somewhat angular in old age. Fertile (inner or upper) surface some shade of brown:
tan to yellow-brown to brown, cinnamon-brown, or dark brown(usually paler or yellower
when growing in or near snow); usually wrinkled, veined, orconvoluted, especially toward
the center, but sometimes smooth. Exterior (underside) more or less smooth, paler or
whitish or somewhat translucent when moist. Flesh rather thick but brittle. ST ALK some¬
times absent but more often present as a short (up to 1 cm long), thick (1-3 cm), narrowed
base; white or tinged tan to brown, usually appearing “gathered” (i.e., with broad ribs
and/ or pits). SPORES 25-35 *(8) 11-16 microns, spindle-shaped with an apiculus(knob
or short projection) at each end, smooth or becoming minutely roughened at maturity,
with one large central oil droplet and two or more smaller ones at the ends.
HABIT AT: S olitary to gregarious or clustered on ground or around old stumps or occa¬
sionally on rotten wood in forests; widely distributed and common, fruiting mainly in the
spring or early summer under conifers. It is one of the characteristic “snowbank” fungi of
western mountains and often occurs with Gyromitra gigas, G. esculenta, and morels.
Sometimes it begins developing while still under the snow!
EDIBILITY: Not recommended. Some people eat it, but care must be taken to cook it
thoroughly and identify it correctly.
DISCINA 799

COMMENTS: This common springtime fungus is best told by its saucerlike to flattened,
umbilicate, or slightly downturned, yellowish to brown fruiting body that frequently has
a short, thick stalk. Like many of the larger Discomycetes, it develops quite slowly, and
since spores are not produced until the fruiting body is fully grown, “sterile” specimens are
often encountered. D. ancilis is apparently a synonym. There are several similar species of
Discina that are best differentiated microscopically. These include: D. apiculatula andD.
macrospora (the latter often with a reddish-brown fruiting body when mature); D.
olympiana, a smaller species found in the Pacific Northwest; andD. leucoxantha, a yel¬
lowish, sometimes stalkless, truncate-spored species found under both hardwoods and
conifers, particularly in eastern North America. Also very similar is Gyromitra mela-
leucoides (-Peziza melaleucoides, Paxina recurvum), with a somewhat waxy-looking
underside when moist, a sometimes slightly longer and narrower stalk, and much shorter
(10-14 microns), non-apiculate spores. See also Disciotis venosa and Peziza species.

GYROMITRA (False Morels)

Medium-sized to large fungi found on ground or rotten wood. FRUITING BODY with a cap and
stalk. CAP usually contorted, lobed, wrinkled, or brainlike (butsometimes saddle-shaped); margin
free from stalk or attached. Fertile surface some shade of brown. Flesh often rather brittle. STALK
smooth or ribbed, simple (round) or complex (folded or chambered) in cross-section. SPORES
typically elliptical or elongated, smooth or roughened, usually with two oil droplets. Asci lining
outer surface of cap; typically 8-spored, operculate, non-amyloid.

THIS is a small but prominent and infamous genus whose grotesque fruiting bodies have
inspired a number of fanciful nicknames: brain mushrooms, elephant ears, lorchels, beef¬
steak morels, and of course, false morels. Despite the latter name, Gyromitras are unlikely
to be mistaken for morels {Morchella and Verpa) because their cap is neither deeply pitted
nor conical or thimble-like. They are easily confused with Helvellas, however, and several
species have been shuffled back and forth between Gyromitra and Helvetia because of
their “in-betweenness.” As currently defined, forms with a brown brainlike cap belong
to Gyromitra, while those with a differently colored and/or saddle-shaped cap (with
the exception of G. infula and its look-alikes) are placed in Helvetia.
Gyromitras are particularly common under northern and mountain conifers in the
spring, often at the same time as morels. G. infula, however, fruits in the summer and fall
(or the winter in coastal California). Several species, notably G. esculenta, G. infula, and
G. ambigua, are dangerously poisonous (even deadly!) raw, yet are eaten without ill effect
by many people. This apparent contradiction can be explained by two facts: (l)The toxin,
M M H (monomethylhydrazine), is extremely volatile, i.e., it is usually removed by cooking
or drying, and (2) there is a very narrow threshold between the amount of M M H the human
body can “safely” absorb and the amount that will cause acute poisoning and even death.
It seems foolhardy to risk eating the MMH-containing species, yet many people do and
dried G. esculenta is sold in many European markets. If you must try them, then
always either dry them out (then rehydrate and cook them) or parboil them first and
throw out the water, being sure not to inhale the cooking vapors (which contain MMH),
then saute them. Never eat them raw and never eat a large amount. MMH, incidentally,
is also carcinogenic and is used as a rocket fuel. See p. 893 for more details on its effects.
Gyromitra is a much smaller genus than Helvetia, but the fruiting bodies are usually
larger. Four species are fully described here and several others are discussed.

Key to Gyromitra
1. Cap shallowly cup-shaped to flattened, disclike, or with the edges turned down slightly; under¬
side of cap not ribbed .G. melaleucoides (see Discina perlata, p. 798)
1. Not as above . 2
800 HELVELLACEAE

2. Stalk smooth or indistinctly grooved; cross-section of mid-stalk round to somewhat flattened

(i.e., with one round hollow or two flattened ones) . 3
2. Stalk with very distinct vertical ribs and I or complex (i.e., folded, chambered, or convoluted
in cross-section—but make the cross-section above the base); stalk usually thick . 6
3. Cap usually saddle-shaped or “winged” (with 2-3 lobes), the surface usually wrinkled in central
portion (directly above stalk), but usually smooth toward edges; lower edge of cap usually free
from stalk; stalk stout, thick; typically found under hardwoods in eastern North America,
mostly in the spring . G. brunnea (see G. infula, p. 802)
3. Not as above; either more conspicuously brainlike at maturity or found elsewhere or surface
not wrinkled; stalk often relatively slender; common and widespread .4
4. Stalk whitish or pale-colored, often rather spongy and usually stuffed with a cottony pith (but
may be hollow in age); cap joined to stalk only at very top, the margin usually inrolled when
young; spores without oil droplets .(see Verpa, p. 793)
4. Not with above features . 5
5. Cap lobed and brainlike (intricately folded and wrinkled) at maturity, but surface usually
smoother when young; found in spring or early summer .G. esculenta, p. 801
5. Cap lobed (saddle-shaped, hoodlike, etc.) but not brainlike, the surface smooth or only slightly
wrinkled, even in age; found mostly in late summer, fall, and winter G. infula & others, p. 802
6. Stalk with deep, widely-spaced ribs that continue onto underside of cap; cap thin and spreading
(umbrella-like) or appearing “puffed up”; margin usually free from stalk . 7
6. Not as above . 8
7. Stalk often reddish- or purplish-tinged near base; spores elliptical; found in western North
America . G. californica, p. 804
7. Not as above; spores round; eastern .G. sphaerospora (see G. californica, p. 804)
8. Cap usually yellow-brown to ochre to tan or occasionally darker brown; stalk massive (nearly
as thick as cap), often short; entire fruiting body with a chunky, almost cubical stature; found
in spring and early summer when or soon after the snow melts ... G. gigas & others, below
8. Not as above; cap usually brown to reddish-brown. 9
9. Cap brainlike to irregularly wrinkled or even pitted at maturity, not strongly lobed .
.G. caroliniana{see G. esculenta, p. 801)
9. Cap often saddle-shaped or “winged” (with 2-3 large lobes), surface not wrinkled or wrinkled
mainly at center (in area directly above stalk) .G. brunnea (see G. infula, p, 802)

Gyromitra gigas
(Snow Mushroom; Snowbank False Morel; Bull Nose; Walnut)
CAP 3-10 (25) cm broad, 3-6 (15) cm high, brainlike or strongly and deeply convoluted
and wrinkled, but typically compact (i.e., without strongly projecting lobes); fertile
surface typically yellow-brown to butterscotch-brown or tan when fresh, but at times
darker brown or even reddish-brown (especially in age); attached to the stalk at or near
the margin. Interior chambered; underside (sterile surface) usually whitish. Flesh thin and
fairly brittle. STALK massive (usually as thick or almost as thick as the cap) and usually
short (sometimes completely hidden by the cap margin!), 2-10 cm long and thick, white or
whitish, often rather irregular in shape or thicker at base; ribbed or wrinkled and grooved;
strongly channelled or folded in cross-section. SPORES 24-36 * 10-15 microns, elliptical,
the ends lacking projections or with only very short, blunt ones, smooth or finely rough¬
ened, typically with one large central oil droplet and smaller ones at the ends.
HABITAT: Solitary, scattered, or in groups on ground or rotten wood in coniferous
forests (occasionally under hardwoods); common throughout the mountains and colder
parts of western North America in the spring and early summer, typically near melting
snow or soon after the snow disappears. In our area, where the winters are mild, it seems
to be absent. However, I have seen enormous fruitings in the Sierra Nevada and Cascades.
EDIBILITY: Edible and popular. Apparently it can be eaten safely without parboiling
(e.g., sauteed like a morel), but should never be eaten raw. Some people prefer it to the
true morels—but be sure you identify it correctly!
Gyromitra gigas. This edible “snowbank” species does not have the bad reputation of other Gyro-
mitras. Note the chunky, almost cubical stature and thick stalk that is folded or chambered within.

COMMENTS: The chunky or almost cubical stature (see photo), strongly wrinkled cap,
and internally folded stalk are the telltale traits of this common “snowbank” fungus.
G. montana and Neogyromitra gigas are synonyms. It sometimes grows with the dan¬
gerous G. esculenta, but can be distinguished by its yellower cap color, chunkier stature,
and complex stalk. G. californica has a ribbed stalk, but usually has a strongly spreading
(umbrella-like) or puffed-up cap. G. gigas shows considerable variation in size. Specimens
weighing more than two pounds have been reported, but the normal size range is walnut-
or softball-sized. G. korfii (called G. fasdgiata in some books) is a very similar vernal
species. It is more numerous in eastern North America than in the West (under both
hardwoods and conifers) and has spindle-shaped spores with prominent projections at
both ends. For other complex-stalked species, see G. caroliniana (under G. esculenta)
and G. brunnea (under G. infula).

Gyromitra esculenta (False Morel; Brain Mushroom) Color Plate 206

CAP 3-12 cm high and broad or occasionally larger, sometimes cup-shaped when very
young but soon becoming lobed or even saddle-shaped and at maturity intricately wrinkled
and folded (brainlike) but not pitted. Fertile surface nearly smooth when very young but
becoming more wrinkled as it matures, deep reddish-brown or purplish-brown to bay-
brown, dark brown, or brown, or in some forms yellowish-brown; typically attached to
the stalk at several points. Underside usually paler; interior hollow or chambered. Flesh
rather thin and brittle. STALK 2-10 (15) cm long, 1-2.5 (4) cm thick, white to tan, flesh-
colored, reddish, or sometimes colored like the cap (but often paler); equal or sometimes
thicker at either end, smooth or grooved vertically but not ribbed; stuffed or hollow (or
with two narrow hollows) in cross-section. SPORES 17-28 * (7)9-13 (16) microns, ellip¬
tical, smooth, usually with two oil droplets.
HABITAT: Solitary, scattered, or in groups under both hardwoods and conifers; fruiting
in the spring (or early summer in colder climates) and very widely distributed, but espe¬
cially common in northern and montane coniferous forests. It is not very common in our
area (perhaps the winters aren’t cold enough), but is often abundant in the Sierra
Nevada, Cascades, and other mountain ranges of the West. It is often encountered while
morel hunting. The fruiting bodies persist for several weeks before decaying.
801
Gyromitra esculenta. Note how younger specimens at center have a relatively smooth cap, and older
(but smaller) one at right is intricately convoluted or brainlike.

EDIBILITY: Dangerously poisonous, at least raw! Although eaten without ill effect by
many people, this species has caused numerous deaths in Europe (see p. 799 for more
details). Far fewer cases of poisoning have been reported from North America, suggesting
that the American version is safer than the European one, or that certain ecological
variants are less toxic. However, the paucity of poisonings may also indicate that fewer
Americans eat it, or that those who do treat it with more caution. I certainly don’t think it’s
worth the risk. In the words of mycophagist Charles Mcllvaine, who was famous for his
willingness to eat almost anything'. “It is not probable that in our great food-giving country
anyone will be narrowed to G. esculenta for a meal. Until such emergency arrives, the
species would be better left alone.”
COMMENTS: The brown cerebral cap and smooth to slightly grooved stalk plus its
occurrence in the spring are the hallmarks of this infamous fungus. Despite the moniker
“false morel,” it can scarcely be confused with the true morels, for the brainlike cap is
never deeply pitted or honeycombed. California specimens are more often reddish-brown
than yellowish-brown, but regional variation can be expected. The stalk is often slightly
grooved or compressed, but does not show the intricate folds in cross-section that are
typical of G. gigas. Other species: G. caroliniana is a springtime species that favors low
hardwood forests in eastern North America, but has also been reported from scattered
localities in the West. It has a brainlike to irregularly wrinkled or even pitted, brown to
reddish-brown cap, but has a massive stalk (up to 20 cm thick!) that is deeply furrowed
and complex (folded) in cross-section. It can attain weights of several pounds each, and is
said to be edible if prepared properly (parboiled, etc.). Like G. esculenta, however, it is
best avoided, particularly if there are morels out and about!

Gyromitra infula (Hooded False Morel)

CAP 3-12 (15) cm broad and high, sometimes cup-shaped when very young but soon
developing projecting lobes; at maturity with 2-3 (rarely 4) lobes, saddle-shaped to hood¬
shaped to irregularly lobed but not intricately wrinkled or brainlike; fertile surface usually
reddish-brown to dark brown, but in some forms yellow-brown, smooth to uneven but not
intricately folded; cap typically attached to stalk at several points, the margin usually
incurved (toward stalk). Underside (sterile surface) paler, minutely velvety; interior
hollow or chambered. Flesh rather thin and brittle. STALK 1-8 (12) cm long, 0.8-3 cm
thick, equal or tapering upward, smooth or minutely velvety, sometimes indistinctly
grooved but not ribbed; colored like the cap or paler(in some forms whitish); hollow(or
with two narrow hollows) in cross-section. SPORES( 15) 17-23 (26) ><(6)7-10(12) microns,
oblong-elliptical, smooth, typically with two large oil droplets.

802
Gyromitra infula, rather old specimens. Note how stalk darkens with age and how the shape of the
cap is quite variable but never brainlike. See next page for a distinctly saddle-shaped specimen.

HABITAT: Solitary to gregarious under both hardwoods and conifers, usually on rotten
wood but also on soil, humus, along roads, in burned areas, etc.; widely distributed. In
most regions it fruits in the late summer and fall—a feature that helps distinguish it from
G. esculenta and other vernal species. In coastal California, however, it fruits in the winter
and early spring and is the most common Gyromitra (especially under oak and pine).

EDIBILITY: Poisonous! It contains MMH and should never be eaten raw! For more
details, see comments on p. 799 and p. 893.
COMMENTS: The saddle-shaped to irregularly lobed cap which is usually reddish-brown
to dark brown plus the more or less smooth (non-fluted) stem are the distinguishing
features of this false morel, which is also known as Helvetia infula. Its penchant for grow¬
ing on rotten wood is also distinctive, but by no means consistent or definitive. Saddle-
shaped specimens are apt to be confused W\ih Helvetia, and it is keyed out under that genus.
The color is quite distinctive, however, and the stalk is thicker than that of most non-
fluted Helvellas. The surface of the cap can be smooth to uneven or slightly wrinkled,
but is not pitted as in the true morels nor intricately folded or brainlike as in G. esculenta.
Smooth specimens of the latter species can usually be distinguished by their growth in the
spring soon after the snow melts. Other species: G. ambigua is a “dead ringer ” for G.
infula. Although it sometimes has a slight lilac or purplish tinge to the fruiting body, it
can only be differentiated with certainty by its spores, whichare longer(22-33 microns) and
have blunt projections at both ends. It has a northern distribution, fruits at the same time,
and is also poisonous. G. brunnea{-G. under woodii and now called G.fastigiata, a name
which has also been applied to G. korfii) is a springtime species found principally under
hardwoods in eastern North America. It has a pale to dark reddish-brown cap that is
usually saddle-shaped or “winged” (divided into 2-3 lobes—hence its common name,
“Elephant Ears”). The cap surface is often wrinkled above the stalk but relatively smooth
near the margin, and the undersides of the lobes are free from the stalk but often fused to

Gyromitra infula. Note the relatively smooth spore-bearing surface. It can be confused with young
specimens of G. esculenta, but fruits at a different time, often on rotten wood.
Left: A dark, saddle-shaped example of Gyromitra infula. Right: Gyromitra californica. Although
dried out, this specimen shows the widely spaced ribs on the stalk which fan out on underside of cap.

each other. The stalk is usually white and fairly thick (2-5 cm) and ranges from hollow to
complex (folded) in cross-section. It has finely warted spores and is edible with caution.

Gyromitra californica (U mbrella False M orel)

CAP 4-12 (25) cm broad, convex to umbrella-like or undulating or occasionally broadly
saddle-shaped (usually strongly spreading and often appearing “puffed-up”); fertile
surface uneven to somewhat convoluted but not brainlike, tan to brown, olive-brown, or
light to dark grayish-brown; margin typically free from the stalk and incurved toward it.
Underside (sterile surface) whitish or creamy, ribbed, minutely hairy to smooth. Flesh very
thin and fragile, almost papery when dried. STALK 3-8 (12) cm long, 2-5 (6) cm thick,
equal or narrowed at base, often rather short, deeply and irregularly fluted or with several
widely spaced ribs that extend onto the underside of the cap, where they fan out and con¬
tinue to or nearly to the margin; white or sometimes aging grayish, buff, or yellowish, the
base (or sometimes the entire stalk) usually with a pinkish, rosy, vinaceous, or purplish
tinge; typically not internally chambered in cross-section. SPORES 13-19 * 7-10 microns,
elliptical, smooth; ends often apiculate at maturity, with small oil droplets.
HABITAT: Solitary, scattered, or gregarious in woods and at their edges, along streams,
or often in somewhat disturbed soil(e.g., along old skid roads); widespread in western
North America, fruiting mainly in the late spring and summer, but sometimes in the fall
or even winter. I have not found it in our area, but it occurs in the Sierra Nevada. As the
species epithet implies, it was originally collected in California, but seems to be more
common in the Pacific Northwest and Rocky Mountains. Alexander Smith says he could
have collected “several bushel baskets” of it on the Olympic Peninsula one year.
EDIBILITY: To be avoided. It is edible according to some reports, poisonous according
to others. The presence of MMH (see p. 799) would account for this contradiction.

COMMENTS: Also known as Helvetia californica, this species can be told from other
Gyromitras by its thin, strongly spreading or umbrella-like cap and distinctly ribbed stalk,
and from Helvetia maculata by its pinkish- or vinaceous-tinged stalk (a fairly reliable but
not foolproof feature). The cap color is rather variable, but is not typically as yellow as that
of G. gigas, nor as red as that of G. esculenta. Other species: G. sphaerospora is its eastern
North American counterpart. It is quite similar but has round spores, and occurs at least
as far west as Montana.

804
 805

HELVELLA (Elfin Saddles)

Small to medium-large fungi found on ground or rotten wood. FRUITING BODY with a cap and
nearly always a stalk. CAP cuplike to plane, saddle-shaped, or irregularly lobed but not usually
brainlike; margin usually free from stalk(except H. lacunosa). Fertile surface white to brown, gray,
or black. Flesh often rather brittle and thin. STALK usually well-developed{occasionally practi¬
cally absent), smooth or ribbed or deeply Jluted and pitted; simple or complex (folded) in cross-
section, fragile or tough. SPORES typically elliptical and smooth with one large central oil drop¬
let. Asci lining exterior of saddle or interior of cup, typically 8-spored, operculate, not amyloid.

HELVELLA is a common, distinctive, and attractive genus with a wide range of shapes
and sizes. Some species are cuplike (and are more aptly called “elfin cups’’); others look
like miniature saddles (hence the popular name “elfin saddles”); still others are irregularly
lobed. The cap, however, is not brown and brainlike as in Gyromitra, nor thimble-like as
in Verpa, nor prominently pitted as in Morchella.
Helvella( spelled Elvela in some older books) splits nicely into four groups: thosespecies
with a cuplike cap and a ribbed (or practically absent) stalk (e.g., H. acetabulum)', those
with a more or less saddle-shaped cap and smooth or non-ribbedstalk(e.g.,//. compressa)',
those with a cup-shaped cap and smooth, well-developed stalk (e.g., H. macropus); and
those with a saddle-shaped to irregularly lobed cap and a deeply ridged or fluted stalk
(e.g., H. lacunosa). Some authorites elevate some of these groups to genus rank (e.g.,
Leptopodia, Cyathipodia, Macropodia, Macroscyphus, Paxina), but their characters
intergrade to some extent and microscopically they are very similar.
The cup-shaped Helvellas are apt to be mistaken for cup fungi, but tend to have more
markedly ribbed and/ or longer stems. The convoluted species, on the other hand, can
be confused with Gyromitra, but are usually smaller and/ or differently colored.
Helvellas do not have the sinister reputation of the Gyromitras, but should be treated
with extreme caution. H. lacunosa is definitely edible, but many of the species have not
been adequately tested. Others are too small or rare to be of value. If you really must eat
Helvellas, always cook them thoroughly (in case they contain MMH) or dry them out.
Helvellas, like other Ascomycetes, are most numerous in the spring, but a few species
fruit in the summer and fall, and in mild climates such as ours they can be abundant in the
winter. They grow under trees and shrubs, along streams, and in the woods, or less com¬
monly in the open. Over 30 species have been reported from North America. Most of these
are keyed out or mentioned in this chapter; eleven are fully described.

Key to Helvella
1. Cap typically concave (cup-shaped) to plane, occasionally becoming convex or saddle-shaped
in old age (examine several specimens if possible) . 2
1. Cap typically convex (umbrella-like) to miter-shaped, hoodlike, saddle-shaped, irregularly
lobed, or brainlike, but sometimes with an upturned or inrolled margin when young that makes
it appear somewhat cup-shaped (examine several specimens if possible) . 14
2. Fertile (upper or inner) surface of cup black or blackish; stalk and base of cup not whitish . 3
2. Fertile surface not black, or if so then stalk or base of cup white or significantly paler than fertile
surface . 5
3. Stalk and exterior (underside) of cup or cap black or very dark, sometimes with ribs; fertile
(upper) surface not fading appreciably . 4
3. Stalk usually grayish (paler than fresh fertile surface); fertile surface often fading to grayish-
brown in age .H. villosa(see H. macropus, p. 810)
4. Stalk well-developed (1 -4 cm long, 2-5 (7) mm thick); fertile (upper) surface black or sometimes
narrowly margined with white; cup fleshy or brittle; spores elliptical, with one large oil droplet;
found in many habitats but not with melting snow ...//. corium (see H. macropus, p. 810)
4. Not with above features; stalk merely a narrowed base or if well-developed then fruiting body
tough or at least not breaking easily and/or growing in or near melting snow .
.(seeSarcosoma&c Allies, p. 826)
806 HELVELLACEAE

5. Stalk well-developed and slender, without prominent ribs, usually less than 6 mm thick (or if
thicker than fertile surface yellow-brown to olive-brown and underside blackish or fertile
surface more or less pinkish-brown with a scalloped or lobed margin) . 6
5. Not as above; stalk thicker and stouter than above and I or with prominent iongitudinal ribs 9
6. U pper stalk and exterior or underside of cup usually hairy or densely scurfy; stalk well-defined,
usually 1 cm or more long; fertile surface of cup grayish-tan to gray or grayish-brown or dark
brown; exterior same color or paler (not black) . 7
6. Not with above features . (see Pezizaceae & Allies, p. 817)
7. Underside of cup (sterile surface) often whitish, at least toward base; stalk often whitish also;
rather rare .H. cupuliformis (see H. macropus, p. 810)
7. Not as above; upper stalk and underside of cup usually grayish or colored like fertile surface
(but base of stalk may be whitish); fairly common . 8
8. Cap often becoming plane or even convex in age, the margin sometimes splitting to form lobes;
upper surface quite dark (grayish-brown); spores elliptical H. villosa(see H. macropus, p. 810)
8. Cap usually remaining cup-shaped (not often plane in age); upper or inner surface gray to
grayish-brown to grayish-tan or paler; spores often elongated H. macropus & others, p. 810
9. Cap often spreading or flattened in age and/ or center often wrinkled; fertile (upper) surface
yellowish to tan or brown (not normally gray or black); stalk usually short; usually found in
spring shortly after snow melts . (see Discina, p. 797)
9. Not with above features . 10
10. Ribs on stalk extending onto underside of cup, often nearly to the margin . 11
10. Ribs on stalk terminating at or near base of cup . 12
11. Cap gray to grayish-brown.H. griseoalba & H. costifera(see H. acetabulum, p. 807)
11. Cap light to dark brown or occasionally tinged violet. H. acetabulum, p. 807
12. Stalk (1) 3-7 cm or more long, usually well-developed, markedly ribbed; cap shallowly cup¬
shaped with opposite sides rolled up when young, often nearly plane in age H. queletii, p. 809
12. Stalk short (usually less than 2.5 cm long) or poorly developed; cup usually deeper or more
regularly shaped than above . 13
13. Interior of cup usually tan to pale brown to yellow-brown; exterior with brown hairs; texture
rather tough . (see Pezizaceae & Allies, p. 817)
13. Not as above .H. leucomelaena & others, p. 808
14. Stalk distinctly fluted, ridged, or ribbed longitudinally, often showing folds or chambers in
cross-section . 15
14. Stalk not as above, but may show a few indistinct grooves, especially near base; stalk simple and
round to slightly compressed in cross-section . 21
15. Cap typically spreading, undulating, or appearing puffed-up or umbrella-like, tan to brown or
grayish-brown, dark brown, or brownish-black (but not truly gray or black); margin of cap
usually free from stalk; stalk with widely spaced ribs that fan out on underside of cap and often
extend nearly to margin; stalk at least 1 cm thick (usually 2 cm or more), sometimes pinkish-
or reddish-stained at base .(see Gyromitra, p. 799)
15. Not with above features (but may have some of them); base of stalk not normally reddish- or
pinkish-stained; cap usually differently shaped or colored or stalk thinner . 16
16. Cap white to buff, creamy, or pale tan .H. crispa & others, p. 816
16. Cap darker (dark tan to brown, gray, or black) or sometimes pale gray or covered with a whitish
fungal parasite . 17
17. Cap typically gray to black (occasionally dark grayish-brown) . 18
17. Cap typically some shade of tan, brown, or grayish-brown . 20
18. Cap usually convex or umbrella-like; stalk slender, usually lacking external holes or pits ....
. H. phlebophora(see H. lacunosa, p. 815)
18. Cap usually saddle-shaped to irregularly lobed or brainlike; stalk often with external holes 19
19. Cap irregularly lobed or brainlike to saddle-shaped, the surface often wrinkled; stalk with
external pits and/or chambered in cross-section; very common, especially in western North
America .H. lacunosa, p. 815
19. Cap usually more or less saddle-shaped, surface not wrinkled; stalk ribbed but usually lacking
pits; more common in eastern North America than West H. sulcata^see H. lacunosa, p. 815)
HELVELLA 807

20. Cap brown to pale or dark reddish-brown or chocolate-brown; spores finely warted at maturity;
found mainly under hardwoods in eastern North America .(see Gyromitra, p. 799)
20. Cap brown to grayish-brown but not reddish-brown; spores smooth; found in western North
America .H. maculata &. others, p. 814
21. Stalk white to creamy or buff (significantly paler than cap unless cap is also pale-colored) 22
21. Stalk darker (brown, dark gray, or blackish), at least above (may be whitish toward base) 27
22. Stalk typically 0.8-2.5 cm thick; cap reddish-brown to dark brown or yellow-brown but not
grayish-brown or black; found on ground or rotten wood; spores with two oil droplets ....
.(see Gyromitra, p. 799)
22. Stalk typically less than 1 cm thick, or if thicker then cap grayish-brown to blackish but not
yellow-brown or reddish-brown; spores with one oil droplet; usually on ground . 23
23. Sterile surface (underside of cap) without hairs (use hand lens if unsure!) . 24
23. Sterile surface minutely hairy, at least when fresh . 26
24. Fertile surface of cap not usually very dark (i.e., yellow-brown to tan to brown or sometimes
grayish-brown or violet-tinged) .H. elastica, p. 813
24. Fertile surface of cap darker than above (grayish-brown to very dark brown to black) ... 25
25. Cap 3-7 cm broad and stalk 0.5-2 cm thick; margin of cap not significantly upturned when
young .H. leucopus, p. 812
25. Cap 1-2.5 cm broad and stalk 2-6 mm thick; margin of cap often upturned when very young
. H. albella{see H. compressa, p. 811)
26. Cap buff to tan, light brown, or cinnamon-buff .H. stevensii (see H. compressa, p. 811)
26. Cap typically darker than above, at least when fresh .H. compressa & others, p. 811
27. Cap irregularly lobed, or if saddle-shaped then stalk typically at least 8 mm thick; cap 3-15 cm
broad, yellow-brown to reddish-brown or dark brown but not gray-brown or black; spores
typically with two oil droplets .(see Gyromitra, p. 799)
27. Not as above; cap usually rather small, 1.5-4 (5) cm broad, usually more or less saddle-shaped,
gray-brown to dark brown to black; stalk typically 2-7 mm thick (occasionally thicker) . 28
28. Underside of cap (sterile surface) minutely hairy, at least when young and fresh. 29
28. Underside of cap hairless (use hand lens if unsure!) . 30
29. Cap grayish . H. ephippium (see H. atra, p. 813)
29. Cap dark brown to grayish-brown to blackish.H. pezizoides {see H. atra, p. 813)
30. Cap dark sooty-gray to black.H. atra, p. 813
30. Cap usually paler (brown to gray but not black) .H. subglabra (see H. atra, p. 813)

Helvetia acetabulum (Brown Ribbed Elfin Cup)

CAP 2-8 cm broad and up to 4 cm deep, cup- or bowl-shaped. Fertile surface (interior)
light to dark brown (or in one form tinged violet), smooth; margin even or irregularly split
in age. Exterior (underside) brown above, paler (white or creamy) at base, conspicuously
ribbed, the ribs blunt or sharp-edged, branching, usually creamy and typically extending
at least half way up the cup and often nearly to the margin; smooth or minutely hairy.
Flesh thin, rather brittle. ST ALK sometimes absent but usually present as a stout, deeply
ribbed base, 1-5 (9) cm long and 0.5-3 cm thick; equal or thicker below, white or creamy
(or sometimes brown if very short); convoluted or chambered in cross-section. SPORES
16-20 x 11-14 microns, elliptical, smooth, with a central oil droplet.
HABITAT: Solitary to gregarious on ground in woods and at their edges; widspread and
fairly common. It is usually listed as a spring and early summer species, but in our area it
fruits in the winter and early spring. I find it once or twice a year, usually under oak.
EDIBILITY: I can find no information on it.
COMMENTS: Formerly known as Paxina acetabulum, this species looks like a cross
between a cup fungus and an elfin saddle. (If the cup were inverted, you’d have an elfin
saddle!) The well-developed ribs that extend far up the underside of the cup help to
separate it from H. leucomelaena and H. queletii. Other species with ribs extending up the
Helvetia acetabulum. Note how prominent whitish ribs on stalk extend onto underside of cup (left),
and how stalk is complex in cross-section (bottom); upper surface of cup is brown or tan.

underside of the cup include: H. griseoalba, similar in size but with a gray cup; and H.
costifera, smaller and grayish-brown.

Helvetia leucomelaena (White-Footed Elfin Cup)

FRUITING BODY cup- or bowl-shaped, with a short stalk or base, (1) 2-5 (7) cm broad
and high. Fertile surface (interior of cup or“cap”) smooth, dark gray to dark brown, dark
grayish-brown, or blackish, the margin often finely scalloped, lobed, or split, especially
in age. Exterior colored like the interior above, shading into white or creamy below,
minutely roughened, not ribbed or only ribbed basally(i.e., appearing “gathered”). Flesh
thin, rather brittle. STALK shortand stout to practically absent, 0.5-2 (4) long,0.5-1.5 (3)
cm thick at apex, with distinct, broad, low, rounded, whitish ribs that terminate at the base
of the cup; usually white but at times dingy grayish; channelled in cross-section (especially
the upper portion). SPORES 20-23 (25) * 10-14 microns, elliptical to somewhat oblong,
smooth, with a single central oil droplet.
HABITAT: Scattered to densely gregarious or clustered(occasionally solitary) onground,
usually near or under pine and other conifers; widespread, but particularly common in
western North America. In our area it fruits in the winter and early spring, but is infre¬
quently encountered (or easily overlooked because of its dark color). Farther north and at
higher elevations it is quite common in the spring and early summer. In all regions it
favors bare, grassy, or hard-packed soil along roads and paths. The fruiting bodies begin
their development underground, and their stalks are often buried until maturity.

Helvetia leucomelaena. Left: Side view showing the short whitish ribbed stalk. Right: View of dark
inner surface and finely scalloped margin; note how short stalk is covered by dirt.
HELVELLA 809

EDIBILITY: I can find no information on it.

COMMENTS: Formerly known asPaxina leucomelas, this species is likely to be mistaken
for a cup fungus. However, the dark “bowl” and whitish, distinctly ribbed stalk plus the
fairly fragile flesh place it in Helvella. The stalk is sometimes so short as to be non-existent,
in which case the base of the bowl appears “gathered” (folded or ribbed).//, crassitunicata
and H. solitaria are two very similar western species that differ microscopically. The
first, however, usually fruits in the late summer and fall and has a northern distribution
(the Pacific Northwest and Alaska), while the latter has a more distinct stalk and usually
develops above the ground. None of these species have ribs which extend up the exterior
of the cup as in H. acetabulum and relatives, nor is their stalk as long as that of H. queletii,
nor as slender as that of H. macropus and relatives.

Helvetia queletii (Ribbed Elfin Saucer)

CAP 2-8 (12) cm broad, usually concave with opposite margins rolled up and in (in¬
curved) at first, becoming shallowly cup- or saucer-shaped to plane or even with a down-
curved margin in age. Fertile (upper) surface blackish to grayish-brown to brown (often
darker when young) or in one form pale brown to buffy-brown, smooth; margin often
splitting in age. Exterior (underside) same color or more often paler or grayer, minutely
roughened (granulose) to nearly smooth, without ribs. Flesh thin, rather brittle. STALK
(1) 2.5-7 (12) cm long, 0.5-2 (5) cm thick, equal or thicker below, deeply ribbed longi¬
tudinally, the ribs extending only to the base of the cup (cap); white (including the ribs)
or sometimes tinged buff, tan, or ochre. SPORES (17) 19-22 * 11-14 microns, elliptical
to oblong, smooth, with a large central oil droplet.
HABITAT: Solitary, scattered, or in small groups in forest humus or occasionally on
rotting wood, usually under hardwoods; widely distributed but not particularly common.
In most of North America it is a late spring and summer species, but in coastal Cali¬
fornia it fruits, like most Helvellas, in the winter and early spring.
EDIBILITY: I can find no information on it.
COMMENTS: This is a most distinctive species, as the photograph shows. The concave
to plane, grayish-brown to nearly black fertile surface plus the prominently ribbed or
fluted stalk are the principal fieldmarks. The stalk is much longer than that of H. leuco-
melaena and H. solitaria, and the ribs do not extend onto the underside of the cup (“cap”)

Helvella queletii. This beautiful species has a well-developed stalk whose ribs do not extend to margin
of cup. Note how opposite sides of “cup” are inrolled when young (specimen at right).

/
810 HELVELLACEAE

as in H. acetabulum, H. griseoalba, and H. costifera. The peculiar way in which opposite

sides of the “cup” are often rolled up when young (see photo on previous page) is sug¬
gestive of a young H. compressa, but that species does not have a ribbed stalk.

Helvetia macropus (Scurfy Elfin Cup)

CAP 0.8-3 (6) cm broad, sometimes closed up when young but soon opening to become
shallowly cup-shaped, rarely becoming plane in old age; margin at first incurved. Fertile
(upper or inside) surface usually gray to grayish-brown, varying to grayish-tan or grayish-
olive or grayish-buff, smooth. Exterior(sterile surface) colored like the interior or slightly
paler or grayer, hairy or minutely fibrillose-scaly or dandruffy. Flesh thin. STALK 1-5
(7) cm long, 2-5 mm thick, equal or often thickened at the base, not ribbed but sometimes
with wrinkles or pits at the base; colored like the exterior of cup above and hairy or scaly-
dandruffy, usually paler or whitish at the base; round or flattened but not chambered in
cross-section. SPORES (18) 20-25 * 10-12.5 microns, more or less spindle-shaped, finely
roughened or smooth, with one large oil droplet and a smaller one at each end.
HABIT AT: S olitary, scattered, or gregarious on ground or rotten wood under both hard¬
woods and conifers; widely distributed. It is one of the commoner summertime Helvellas
of eastern North America, but in our area it fruits in the winter and spring and is not
very numerous (or at least not often collected because it is so inconspicuous).
EDIBILITY: Unknown, but too small to be of value.
COMMENTS: This cute little elfin cup is best told by its overall grayish color and hairy
or scurfy-scaly sterile surface and stalk. The stalk is not ribbed as in H. leucomelaena or
H. acetabulum, and the elongated spores are unusual for a Helvella. Both short- and
relatively long-stemmed varieties are known. H. villosa is a very similar and widespread,
slightly darker species with a slightly less hairy stalk and a shallower cup that usually
becomes plane or even slightly convex in age (see photo below). I have found it several
times in our area, also in the winter and spring. H. cupuliformis is quite similar in shape,
but has a paler (often whitish) exterior or underside. H. corium is similar in shape but is
entirely black, and the lower portion of its stalk is often somewhat ribbed. It seems to have
a circumpolar (northern) distribution and often occurs in sandy soil or debris. H.pallidula
of eastern North America is a pale tan to pale grayish-brown, cup-shaped species. None
of these have the markedly ribbed stalks of H. queletii and H. acetabulum, and all have the
elliptical spores typical of most Helvellas, rather than the elongated spores of H. macropus.

Left: Helvella macropus, a young specimen with hairy-scurfy underside. Center: Helvella macropus,
mature specimen with a broader cup. Right: Helvella villosa (see comments above) resembles H.
macropus, but its cap or “cup” is often plane in age, as shown here.
Helvetia compressa. Left: Young specimens in which opposite sides of the cap are rolled up and in,
making the cap somewhat cuplike and hiding the fertile surface; sterile surface is whitish and minutely
hairy. Right: A mature saddle-shaped specimen which has faded somewhat (to brown).

Helvetia compressa (Compressed Elfin Saddle) Color Plate 204

CAP 1.5-5 cm high and broad, saddle-shaped to somewhat irregularly lobed (with 2-3
lobes) when mature, typically with a well-developed sinus (cleft), but opposite margins
rolled up and over the fertile surface when young; margin unrolling and often flaring in
age, free from the stalk. Fertile surface brown to dark brown or grayish-brown, smooth.
Underside (sterile surface) white or creamy to grayish-white, minutely hairy. Flesh thin,
rather brittle. STALK 3-10 cm long,0.3-1.2(1.7)cmthick, equalorthickerbelow, smooth
or finely downy, not ribbed; white to pale cream-colored; round or somewhat flattened
in cross-section, but not chambered; base sometimes pitted. SPORES 19-22 (25) * 12-15
microns, broadly elliptical, smooth, with a large central oil droplet.
HABITAT: Solitary to gregarious on ground in woods and at their edges, under trees,
etc.; known only from western North America. In our area it usually fruits in the late
winter and spring under redwood, oak, and various other trees. It seems to be the most
common of our non-fluted Helvellas.
EDIBILITY: Unknown.

Helvetia compressa, mature specimens. Note deep cleft (sinus) in the saddle-shaped cap. These small,
dark specimens could just as easily be referred to H. albella (see comments on next page).
812 HELVELLACEAE

COMMENTS: This attractive elfin saddle is best identified by its combination of brown
to grayish-brown cap with a finely hairy underside and slender, white, non-ribbed stem.
The transformation in the shape of the cap as it matures is apt to cause some confusion
unless intervening stages are found. When young the margin is curled over the fertile
surface so that the whitish underside is most visible and the cap is almost cuplike( see photo
at top of p. 811). In age, however, the margin unfurls and the saddle shape is more apparent
(see other photos on p. 811). H. albella is a very similar but smaller (cap 1 -2.5 cm broad and
high, stalk up to 5 cm long and 2-6 mm thick) species with a grayish-brown to blackish cap.
Some authors describe its underside as hairless while others maintain it is minutely hairy
when young. If the latter is true then its small size is the principal point of departure from
H. compressa. A small form meeting this description is not uncommon in California in
the winter and spring, usually under oak. Another closely related species, H. stevensii
(-H. connivens) is small to medium-sized but has a paler (tan to yellowish- or cinnamon-
buff) cap and slightly smaller spores. It has a wide distribution and also occurs in our area.
Both H. compressa and H. stevensii are distinguished from H. elastica by their upturned
cap margin when young, deeper and narrower cleft (sinus), and minutely hairy sterile
surface (see color plate), and from H. atra and relatives by their whitish stalk.

Helvetia leucopus (Elfin Miter)

cap 3 -7 cm broad and high, often with three or more lobes but sometimes with only two;
shape variable: miter-shaped or appearing saddle-shaped from three different angles, but
sometimes irregularly contorted or occasionally suggestive of an elephant head (one
longer lobe with two “ears”); margin nearly straight when young (not conspicuously
inrolled), the lower edge of each lobe typically joined to the stalk at one point. Fertile
surface typically more or less smooth and very dark (dark brown to dark grayish-brown to
blackish), occasionally mottled with lighter brown areas. Underside (sterile surface) white
or tinged faintly with the cap color, not minutely hairy. Flesh thin, rather brittle. ST ALK
4-12 cm long, 0.7-2 cm thick, equal or thicker below, often flattened or compressed but
not ribbed or deeply fluted; smooth (not hairy), often with hollows or holes at the base;
white or sometimes developing slight smoky-brown stains in age, often curved (giving it
the appearance of a bleached rib); hollow in cross-section. SPORES 20-23 * 14-15
microns, elliptical or slightly oblong, smooth, with a large central oil droplet.

Helvetia leucopus. Note dark cap which often has three or more distinct lobes. Stalk is whitish,
usually hollow, and fairly thick (over 5 mm).
HELVELLA 813

HABITAT: Scattered to gregarious on ground under trees, usually in the spring; widely
distributed. In our area I have found it under cottonwood in the late winter and spring,
but it grows in other habitats as well.
EDIBILITY: Not recommended. The European version is said to be edible, but I can
find no information on North American material.
COMMENTS: The above description is based on several dozen specimens that grew in
sandy soil under a cottonwood near Santa Cruz, California. It agrees perfectly with
European descriptions of H. monachella, a name now considered synonymous with H.
leucopus; H. albipes is yet another synonym. No matter what you call it, the dark cap
contrasts nicely with the white, unribbed stalk, making it a most attractive elfin saddle.
It is most apt to be confused with H. compressa and H. albella, but is usually larger and
stouter, with a more irregular, multi-lobed cap whose underside is hairless.

Helvetia elastica (Brown Elfin Saddle)

CAP 0.5^1 (6) cm broad and high, typically saddle-shaped or miter-shaped with a broad,
shallow sinus (cleft) between the lobes, but sometimes convex (with little or no sinus).
Fertile surface smooth, brown to tan or grayish-tan, the margin straight or somewhat
incurved toward the stalk; lobes free from the stalk or sometimes partially attached to it
or to each other. Underside (sterile surface) whitish to buff or pale tan, smooth, not hairy.
Flesh thin, rather brittle. STALK (2) 4-8 (14) cm long, (0.2) 0.4-1.2 cm thick, equal or
tapered upward, sometimes curved, smooth or indistinctly grooved at the base but not
fluted or ribbed; white to buff or pale yellowish-buff; round to slightly flattened in cross-
section, not chambered. SPORES 18-22(24) * 10-14 microns, elliptical or oblong, smooth
or warted, with one central oil droplet.
HABITAT: Solitary to widely scattered or gregarious in woods or at their edges, particu¬
larly near streams and paths; widely distributed. It occurs in our area in the fall, winter, and
early spring, but is not as common as H. compressa.
EDIBILITY: Unknown.
COMMENTS: Many authors (myself included) have applied the name H. elastica indis¬
criminately to a slew of smallish Helvellas with slender white, non-ribbed stems and
brownish, saddle-shaped caps. However, in the “true” H. elastica the cap is smooth(hair-
less) on its underside and usually has a broad, shallow cleft and a straight to slightly down-
curved margin, whereas in species such as H. compressa and H. stevensii the cleft is nar¬
rower and the lobes more upright, the underside is minutely hairy, and the margin is often
curled up and in (thus hiding the fertile surface) when young.

Helvetia atra (Dark Elfin Saddle)

CAP 0.7 -2 cm high and broad, distinctly saddle-shaped or occasionally 3-lobed when
mature, with a narrow sinus (cleft); margin free from the stalk or occasionally attached.
Fertile surface deep sooty-gray to black, smooth. Underside (sterile surface) pale gray to
blackish, hairless. Flesh thin, fairly brittle. ST ALK 1 -3.5 cm long, 2-5 mm thick, equal or
slightly thicker below, not fluted but occasionally with shallow pits at the base; dark
sooty-gray to black (often palest at base), round or flattened in cross-section but not
chambered. SPORES 17.5-20 * 10.5-12.5 microns, oblong-elliptical, smooth, with a
large central oil droplet.
HABITAT: Solitary, scattered, or in small groups on ground or rotten wood under both
hardwoods and conifers; widely distributed but rare (at least in North America), fruiting
mainly in the summer and fall. I have found it only once—in New Mexico under pine. It is
also reported from Alaska, Montana, and Washington.
814 HELVELLACEAE

EDIBILITY: Unknown.
COMMENTS: This diminutive elfin saddle is easily recognized by its black, distinctly
saddle-shaped cap and non-ribbed, blackish stalk. H. lacunosa is similar in color but larger
and much more common and it has a fluted and pitted (lacunose) stalk; H. corium(see
comments under H. macropus) is black but has a cuplike cap. H. atra has several close
relatives with a dark saddle-shaped cap and dark, non-ribbed stalk, including: H. sub¬
glabra of eastern N orth America, which has a brownish to gray (never black) cap, and two
widespread species with a minutely hairy or downy underside (sterile surface): H. ephip-
pium, which is the same size as H. atra, but grayish; and H. pezizoides, which is slightly
larger and dark brown to grayish-brown or blackish. None of these are worth eating.

Helvetia maculata (Fluted Brown Elfin Saddle)

CAP 1.2-6 cm broad and high, saddle-shaped or irregularly lobed at maturity, the margin
free from the stalk and at first rolled up (thus obscuring the fertile surface), but unfurling
or flaring out with age and sometimes splitting. Fertile surface brown to grayish-brown
or buffy-brown, often mottled with darker and lighter shades, smooth or slightly wrinkled.
Underside (fertile surface) creamy to yellowish, buff, or grayish, minutely hairy, some¬
times with a few ribs extending a little way up from the stalk. Flesh thin, rather brittle.
ST ALK 2-10 cm long, 0.5 -3 cm thick, equal or tapered above, deeply ribbed and sometimes
lacunose (pitted), white to pale buff, sometimes with grayish or brownish stains in age;
convoluted or chambered in cross-section. SPORES (18)20-23 * 12-13.5 microns, bluntly
elliptical or oblong, smooth, with one central oil droplet.
HABITAT: Solitary to gregarious in mixed woods and under conifers; widespread in
western North America. I’ve seen large fruitings locally in the winter and early spring, but
it is not nearly as common as H. lacunosa. In many regions it fruits in the summer and fall.
EDIBILITY: Unknown.
COMMENTS: This showy elfin saddle can be identified by its grayish-brown to brown
mottled cap and deeply ribbed stalk. It is reminiscent of H. crispa and H. lacunosa, but is
darker than the former and paler than the latter. It might also be confused with Gyromitra
californica, but is less fragile and lacks the strongly spreading or umbrella-like cap of that
species. H. fusca is said to be quite similar, but has distinct ribs on the underside of the
cap and lacks a flaring margin. I have not seen it.

Helvella maculata. Cap is brown to grayish-brown but never black. Note deeply fluted stalk.
Helvetia lacunosa is our most common elfin saddle. The cap is black or gray and usually lobed or
wrinkled, while the stalk is white to dark gray, always deeply fluted, and usually lacunose (i.e.,
with visible pits). Note how cross-section of stalk (bottom right) is chambered. In our area this
species occurs in many habitats but is especially abundant under pine.

Helvetia lacunosa (Fluted Black Elfin Saddle)

CAP 1-10 cm or more broad and high (but averaging 2-5 cm), saddle-shaped or more often
irregularly lobed or convoluted. Fertile surface smooth or wrinkled, black to grayish-
black or gray (or rarely whitish); margin typically attached to the stalk at several points
or intervals, often incurved (toward the stalk) when young. Underside (sterile surface)
grayish to grayish-brown or black (rarely white), smooth {not hairy), ribbed or unribbed.
Flesh thin, rather brittle. STALK 3-15 cm long, (0.5) 1-3 cm thick, equal or tapering up¬
ward, deeply ribbed and typically lacunose(i.e., the ribs branching to form elongated holes
or pockets); ribs often sharp and double-edged; white to gray or sometimes black (often
white when young and darker in age), the base often paler; convoluted and chambered in
cross-section. SPORES (12) 15-21 * (9) 11-14 microns, broadly elliptical to nearly round
to oblong, smooth or slightly roughened at maturity, with one central oil droplet.
HABITAT: Solitary, scattered, or in groups in woods and under trees; very widely dis¬
tributed, but especially common in western North America. In our area it is usually abun¬
dant in the winter and early spring in pine forests or on lawns or roadsides where pines
have been planted, but it also occurs with oak and Douglas-fir. In other regions it usually
fruits in the late summer and fall.
EDIBILITY: Edible when cooked and rated highly by some people. It can be sauteed, but
in my experience is rather chewy and bland. The tough stalks should be cooked separately
or discarded. It can also be dried and powdered for use as a seasoning.
COMMENTS: This species is by far the most common of our Helvellas, often appearing
in large groups or troops after winter rains. W hen fresh it is virtually unmistakable because
of its gray to black cap and deeply fluted stem, but it is frequently disfigured by a white,
moldy-looking parasite (Hypomyces cervinigenus). Such specimens should not be col¬
lected for eating. The cap is not brown as in H. maculata, and pale individuals can be
told from H. crispa by the partially attached rather than free cap margin. H. mitra is
an obsolete synonym. There are several similar species, including: H. palustris (-H. phi-
lonotis) of eastern North America, a very similar but smaller species that favors damp or
swampy places and has a deeply ribbed but not lacunose stalk; H. sulcata, which has a

815
Left: This fluted black elfin saddle (Helvetia lacunosa) shows its lacunose (pitted) stalk to good
advantage. (Rick Kerrigan) Center: Another example of Helvetia lacunosa {others can be seen in
photo on p. 815). Right: Helvetia crispa, a common species in eastern North America, but infrequent
in California. The pale (white to buff) cap and fluted stalk are its trademarks.

small, pale gray to black cap that is usually saddle-shaped with a deep, well-defined cleft
plus a ribbed but not lacunose stalk (it often grows on rotten wood); and H. phlebophora,
a small, slender, rare species with a more or less convex or umbrella-like cap. All three of
the latter species have been reported from the Midwest and / or eastern North America by
Nancy Weber, and may occur in the West. I can find no information on their edibility.

Helvetia crispa (Fluted White Elfin Saddle)

CAP l -5 cm broad and high, saddle-shaped to irregularly lobed at maturity; margin
typically at first rolled up, then unfurling and flaring, typically free from the stalk. Fertile
surface smooth or slightly wrinkled, white to creamy, buff, pale pinkish-buff, or tinged
yellowish. Underside (sterile surface) colored like the cap or slightly darker or grayer,
scurfy or minutely hairy. Flesh thin, rather brittle. STALK 3-10 cm long, 0.8-3 cm thick,
equal or tapering upward, deeply ribbed and often lacunose, white to pinkish-buff or
colored like the cap, sometimes darkening slightly in age; convoluted or chambered in
cross-section. SPORES (14) 17-21 (24) * 10-14 microns, elliptical or oblong, smooth,
with a large central oil droplet.
HABITAT: Solitary to gregarious on ground or very rotten wood under both hardwoods
and conifers; widely distributed and fairly common (especially in eastern North America),
but almost totally supplanted in our area by H. lacunosa. I have found it only twice—in
October and March under redwood and tanoak. In other areas it fruits in summer and fall.
EDIBILITY: To be avoided. Some sources report it as edible, but it should be tried
cautiously if at all—and only after cooking it thoroughly.
COMMENTS: The white to buff cap and deeply ribbed or fluted stalk are characteristic
of this beautiful elfin saddle, which is the type species of the genus Helvella. The cap is not
brown as in H. maculata, nor gray or black as in H. lacunosa. Pallid specimens of the
latter occasionally occur, but can be distinguished by their partially attached rather than
free cap margin. Other species: H. lactea has a white to creamy or buff cap and a fluted
stalk, but the underside of its cap is hairless.

816
 817

Cup Fungi
spores
PEZIZACEAE& Allies
IN THESE ubiquitous but inconspicuous fungi the fruiting body is usually concave (cup¬
shaped) to disclike (flat) or cushion-shaped (very slightly convex) and may or may not
have a stalk. In a few species it is ear-shaped, top-shaped, cabbage-like, or irregular. The
spore-bearing asci line the upper surface of the fruiting body or inner surface of the cup.
The spores are shot out of the asci with such terrific force that cup fungi, like other members
of the Pezizales, will often “smoke” visibly (spew out clouds of spores) when handled.
The cup fungi are a vast group embracing five families and nearly one hundred genera
—or more than twice that number if you count the minute cuplike members of the earth
tongue order (Helotiales). Obviously, they can’t be covered thoroughly in a book of this
kind, and no attempt is made to do so. Most cup fungi are difficult to identify anyway
and are too small to be worth eating. A few, such as the Pezizas, can be quite large, but are
still unlikely to show up on anyone’s list of best edibles.
Cup fungi occur in a wide range of habitats—on dung and manure, in grass, humus,
soil, and moss, on wood and foliage, and in recently burned areas. As the characters used
to delineate the five families (Pezizaceae, Sarcosomataceae, Sarcoscyphaceae, Pyro-
nemataceae, Ascobolaceae) and numerous genera are almost entirely microscopic, I have
rather arbitrarily divided the cup fungi into five groups based on color, size, and texture.
These are keyed below.
Key to the Pezizaceae & Allies
1. Fruiting body merely the bowl- or cup-shaped remains of a puffball, usually with traces of spore
powder inside (or if not, then usually very dry and light as a straw) (see Gasteromycetes, p. 676)
1. Not as above . 2
2. Fruiting body beginning as a hollow underground ball, then splitting at the top into several
starlike rays; fertile (inner) surface sometimes grayish or whitish but more often pinkish or
purplish or lilac; exterior without brown hairs . Sarcosphaera, p. 825
2. Not with above features (but may have some of them) . 3
3. Fruiting body an erect hollow club or closed urn that splits lengthwise from the top to form
several rays; found on or near dead hardwoods .Sarcosoma & Allies, p. 826
3. Not as above . 4
4. Fruiting body an irregular mass of cabbage-like, cauliflower-like, brainlike, or contorted tissue,
with or without a stalk . 5
4. Not as above; fruiting body cuplike, earlike, disclike, etc. (but sometimes contorted, especially
if growing in clusters) . 6
5. Stalk present, long (15-30 cm), brown, solid; fertile “head” brainlike, pitted, or cabbage-like,
usually beige or yellowish-brown; found under hardwoods in eastern North America; rare
. Wynnea sparassoides
5. Stalk absent or rudimentary; widely distributed but not common ... Peziza& Allies, p. 818
6. Fruiting bodies consistently slit down one side, usually erect or semi-erect (often standing on
one end like a rabbit’s ear); usually medium-sized, not huge or minute, often clustered or in
contorted masses, not growing on manure or dung; tips of asci not amyloid . Otidea, p. 831
6. Not as above; fruiting bodies sometimes slit down one side but not consistently so, and not
usually growing erect; sometimes growing on dung . 7
7. Fertile (upper or inner) surface of fruiting body brightly colored (red, orange, yellow, blue, or
green but not violet) . Aleuria&t Allies, p. 833
7. Fertile surface some shade of brown, black, tan, dingy yellowish, or violet, or sometimes with
a pinkish or lilac tinge . 8
8. Flesh gelatinous to rubbery-gelatinous, or if not then flesh rather tough (not breaking easily)
and fertile surface dark brown to black .Sarcosoma & Allies, p. 826
8. Not as above; flesh fragile or brittle to slightly rubbery, but usually breaking easily . 9
818 PEZIZACEAE & ALLIES

9. Stalk usually present and flesh rather tough; either fertile surface more or less pinkish-brown
with a scalloped or stellate margin or fertile surface yellow-brown to olive-brown and underside
(sterile surface) blackish .Sarcosoma& Allies, p. 826
9. Not as above . 10
10. Exterior or underside (sterile surface) of fruiting body clothed with brown or black hairs (the
hairs sometimes sparse); tips of asci not amyloid. Aleuria & Allies, p. 833
10. Not as above . 11
11. Fruiting body large to fairly small but not minute, usually (0.5) 1-10 cm broad, often flattened
(disclike) or spreading at maturity; fertile surface yellow-buff to brown to dark brown, or
violet, or occasionally whitish or tinged pinkish; asci with amyloid tips; usually growing soli¬
tary, scattered, or in small groups (a few species in clusters) .Peziza & Allies, below
11. Fruiting body fairly small to minute, 0.5-2 (3) cm broad, often remaining cup-shaped at maturity;
fertile surface variously colored but not often dark brown or violet unless very small; sometimes
growing in large swarms; tips of asci not amyloid . Aleuria & Allies, p. 833

PEZIZA & Allies

Small to fairly large fungi found in a wide variety of habitats. FRUITING BODY typically cup¬
shaped to flattened (disclike) at maturity, but in a few cases truffle-like or irregularly contorted.
Fertile (upper or inner) surface typically some shade of yellow-brown, brown, black, or purple {but
sometimes paler or tinged pinkish. Exterior (underside) smooth, scurfy, or roughened by warts but
not usually hairy. Flesh often rather brittle or at least breaking easily. STALK absent or if present
usually short and continuous with the cup and not normally ribbed. SPORES elliptical to round,
smooth or roughened at maturity, with or without oil droplets. Asci lining upper surface of cup or
disc, typically 8-spored, operculate, with amyloid tips.

THIS is the largest and most common genus of cup fungi. It is also the most mundane. The
majority of its species are dull or dark colored. A short stem is sometimes present but more
often absent. Although most species are cup- or disclike, a few are truffle-like (e.g., P.
ellipsospora) and one, P. proteana, sometimes produces compound fruiting bodies that
look like heads of cabbage.
Identification of Pezizas is difficult even with a microscope. Most species grow in soil,
humus, or rotten wood, but some have more distinctive milieu. For instance, P. vesiculosa
grows on dung, P. violacea on burnt ground, and P. domiciliana in bathrooms, cellars,
rugs, and other domestic settings.
Little is known about the edibility of Pezizas, and considering how difficult they are to
identify, the entire group is best avoided. Only a few of the many species are described
here. Two closely related genera, Plicaria and Pachyella, are also treated.

Key to Peziza & Allies

1. Fruiting body flattened or spreading, attached to substrate by numerous “roots” (rhizomes);
stalk absent; usually under conifers .(see Helvellaceae, p. 796)
1. Not as above; fruiting body if flattened not attached by numerous “roots” . 2
2. Growing on dung or manure . P. vesiculosa & others, p. 823
2. Not as above . 3
3. Fruiting body compound, somewhat reminiscent of a head of cabbage (i.e., composed of many
spoon-shaped or distorted lobes or “cups”); fertile surface whitish to pinkish, creamy, tan, or
lilac-tinged; often but not always growing in burned areas.P. proteana, p. 824
3. Not as above (but fruiting bodies can be clustered and somewhat distorted by mutual pressure)
. 4
4. Growing in charcoal or recently burned areas . 5
4. Not as above . . . ... 8
5. Fertile (upper) surface violet- or lilac-tinged when fresh.P. violacea & others, p. 824
5. Fertile surface creamy to pinkish, tan, yellowish, reddish, brown, or black, occasionally with a
lilac tinge, but if so then overall color quite pale . 6
PEZIZA& ALLIES 819

6. Fertile surface dark brown to black .Plicaria endocarpoides & others, p. 820
6. Fertile surface paler than above. 7
7. Fertile surface creamy to pinkish or faintly lilac-tinged; tips of asci amyloid P. proteana, p. 824
7. Fertile surface reddish to yellowish to tan to brown, grayish-brown, or sometimes distinctly
pink; tips of asci not amyloid .(see A leuria&c Allies, p. 833)
8. Growing in bathrooms, on rugs, in cars, and other domestic situations P. domiciliana, p. 822
8. Not as above; usually growing outdoors. 9
9. Growing underground or under the humus (but sometimes partially exposed); fruiting body
round, lobed, or saucer-shaped with a permanently inrolled margin; interior hollow or with
canals or solid and marbled with veins . P. sp. (unidentified) & others, p. 824
9. Not as above . 10
10. Fruiting body developing underground with only the mouth at ground level (like a hole in the
ground); interior brown to dark brown; margin often lobed or scalloped at maturity; growing
in sand, sandy soil, or sand dunes .P. ammophila (see Sarcosphaera crassa, p. 825)
10. Not as above . 11
11. Fruiting body shallowly cup-shaped to umbilicate (with a central depression) to flat or even with
a downcurved margin; fertile surface usually yellow-brown to tan, but sometimes darker
brown or reddish-brown; short stalk often present; nearly always found in the spring when or
soon after the snow melts, especially under mountain conifers; mature spores often apiculate,
with oil droplets; tips of asci not amyloid . (seeDiscina, p. 797)
11. Not with above features (but may have some of them) . 12
12. Fruiting body somewhat rubbery or rubbery-gelatinous, almost the entire underside attached to
substrate (not just the center); usually found on wet wood . 13
12. Not as above; fruiting body rather brittle and centrally attached, on wood or ground .... 14
13. Fruiting body small (less than 5 mm broad) Pachyella babingtonii(see Peziza repanda, p. 821)
13. Fruiting body larger (typically 2-8 cm broad) Pachyella clypeata(see Peziza repanda, p. 821)
14. Fruiting body small (usually less than 2.5 cm broad) and stalkless, dark brown to nearly black,
usually becoming shallowly saucer-shaped or disclike (flat) at maturity P. brunneoatra, p. 820
14. Not as above; fruiting body usually larger, or if small then paler or more deeply cup-shaped
or with a stalk . 15
15. Fruiting bodies usually slit down one side and often clustered, causing them to be somewhat
contorted; tips of asci not amyloid .(see Otidea, p. 831)
15. Not as above; fruiting bodies often round (spherical) when very young and flattened or
spreading in age, or at least not as above . 16
16. Fruiting body 1-3 cm broad, brown to grayish-brown, often with a short stalk or narrowed
base (especially when young), not normally flattening out in age; tips of asci not amyloid
.(seeAleuria& Allies, p. 833)
16. Not as above . 17
17. Flesh staining yellow or exuding a yellowish juice when squeezed or cut .
. P. succosa (see P. sylvestris, p. 821)
17. Not as above . 18
18. Fertile (upper or inner) surface grayish to dark grayish-brown to nearly black; base of fruiting
body often with obscure ribs .(see Helvetia, p. 805)
18. Not as above; fertile surface usually tan to brown, reddish-brown, etc., but not grayish or
blackish . 19
19. Exterior (underside) of fruiting body brown to reddish-brown even in dry weather .
. P. badia& others (see P. sylvestris, p. 821)
19. Exterior usually white to pale tan, at least toward base (but often brown if wet) . 20
20. Fruiting body usually flattening out in age, the fertile surface usually strongly veined, wrinkled,
or pebbled (especially toward center); usually found on well-drained or sandy soil(e.g., along
streams) in spring; tips of asci not amyloid; spores smooth, without oil droplets .
. (set Disciotis, p. 796)
20. Not with above features (but may have some of them); tips of asci amyloid. 21
21. Usually found on rotten wood, wood chip mulch, or humus rich in lignin; fruiting body often
quite large and spreading (flattened) in age; spores smooth .P. repanda, p. 821
21. Usually found on ground, sometimes on very rotten wood; fruiting body usually medium-sized,
the spores often roughened at maturity .P. sylvestris & many others, p. 821
Left: Plicaria endocarpoides, a blackish charcoal-loving disclike fungus. Right: Peziza sylvestris.
Young specimen at bottom is cuplike; older one at top resembles a torn piece of discarded rubber.

Plicaria endocarpoides
FRUITING BODY 1 -7 cm broad, at first cup-shaped but soon becoming flattened or
undulating. Fertile (upper) surface dark brown to black, smooth or roughened, sometimes
also wrinkled. Exterior(underside) same color or paler, smooth or roughened. Flesh rather
brittle, usually paler or browner than fertile surface. STALK absent or rudimentary.
SPORES 8-10 microns, round, smooth, typically with several small oil droplets.
HABITAT: Solitary to gregarious on burnt ground, old campfire sites, etc.; usually but
not always fruiting in the spring; widely distributed.
EDIBILITY: Who knows?
COMMENTS: Also known as P. leiocarpa, this charcoal-lover is easily told by its dark
brown to black fruiting body that is usually stalkless and flattened at maturity (see photo).
P. trachycarpa is a very similar but smaller (up to 2.5 cm broad) warted species with larger,
roughened spores. P. carbonaria should also be mentioned. All three of these grow in
ashes, have round spores, and are placed in Peziza by some taxonomists.

Peziza brunneoatra
FRUITING BODY 0.5 -2.5 cm broad, cup-shaped becoming flattened (disclike) in age.
Fertile (upper or inner) surface brown to brownish-black, sometimes with an olive tinge,
smooth. Exterior (underside) same color or slightly paler or redder (red-brown). Flesh
thin, brownish, brittle. STALK absent or present only as a very short, narrowed base.
SPORES 16-22 x 8-12 microns, elliptical, smooth becoming roughened (warty) or par¬
tially reticulate at maturity, with one or two oil droplets.
HABITAT: Scattered to gregarious or clustered on damp soil along roads and paths
through the woods, near streams, under trees, etc.; widely distributed but not particularly
common. I have found it several times in our area in the winter and spring.
EDIBILITY: Unquestionably inconsequential.
COMMENTS: The small size, dark color, and tendency to become disclike in old age
help to identify this species. It is a “cup fungus” in name only, for mature specimens are
usually disclike or even slightly convex. Plectania species are also quite dark, but are
tougher and more deeply cup-shaped, while Pachyella species usually grow on wood and
Plicaria species favor ashes.

820
Peziza sylvestris. This common woodland cup fungus resembles many others in its genus. Fertile
surface is brown; exterior is whitish or buff, at least when dry. See photo on p. 820 for more specimens.

Peziza sylvestris (Boring Brown Cup Fungus; Fairy Tub)

FRUITING BODY 2.5-8 cm broad, deeply cup-shaped when young, becoming shallowly
cup-shaped or sometimes flattened in age. Fertile (upper or inner) surface brown, smooth
or often somewhat wrinkled; margin sometimes wavy or scalloped. Exterior finely warted
or roughened to nearly smooth, whitish to pale tan when fresh. Flesh rather fragile.
STALK absent or rudimentary. SPORES 15-20 * 9-10 microns, elliptical, smooth or
becoming finely roughened at maturity, without oil droplets.
HABITAT: Solitary, scattered, orin groups or small clusters on ground, usually in woods;
widely distributed. It is common in our area, especially in the winter and spring, under
both hardwoods and conifers.
EDIBILITY: Unknown
COMMENTS: This is one of many terrestrial brown Pezizas that are best differentiated
microscopically. As a group they can be recognized by their brown color, moreorlesscup-
shaped fruiting body, fragile texture, and growth (usually) on the ground in the woods.
P. sylvestris seems to be the most common of the lot in coastal California, but may not be
so abundant in other regions. Other boring brown cup fungi include; P. echinospora
(-P. pustulata?), with a pale brown fertile surface and coarsely roughened or pimpled
whitish exterior; P. succosa, light brown with a whitish exterior and turning yellow (or
exuding a yellow juice) when broken; P. cerea, a yellowish-buff species; and three species
that often have a reddish tinge when fresh (i.e., they range from brown to reddish-brown to
brownish-orange): P. badia andP. badioconfusa, both medium-sized to fairly large witha
brown to reddish-brown exterior (the former with partially reticulate spores, the latter
with warty spores); and P. petersii, a smaller species (up to 4 cm broad) with a smooth
exterior and smaller spores (10-12 microns long). All of these Pezizas are cuplike at first
and are sometimes called “fairy tubs” because they look like miniature bathtubs when
filled with rainwater. In age, however, they can be quite flat or irregular in shape, or in the
words of Gary Lincoff, “may resemble discarded torn pieces of rubber” (see photo on p.
820). None are worth eating.

Peziza repanda (Spreading Brown Cup Fungus)

FRUITING BODY 4-13 cm broad or occasionally larger, cup-shaped when young but
expanding to nearly flat or often wavy (undulating), the margin often splitting. Fertile
(upper or inner) surface pale brown to medium brown, tan, or in age somewhat darker,
smooth to somewhat wrinkled or convoluted at the center. Exterior (underside) pallid.
Flesh fairly brittle. STALK absent or present only as a short, narrowed base. SPORES
14-18 x 8-10 microns, elliptical, smooth, without oil droplets.

821
Left: Peziza repanda, fairly young specimens which have not yet flattened out; note growth on wood.
Right: This unidentified Peziza (see description on pp. 824-825) is one of several truffle-like species
that grow underground or on the surface of the soil beneath the humus. Note irregular shape.

HABITAT: Solitary, gregarious, or in clusters on logs and branches (especially of hard¬

woods), lignin-rich humus, etc. (often in nurseries where wood chip mulch is used); widely
distributed. It is common in our area in the winter but fruits practically year-round.
EDIBILITY: Said to be edible, but easily confused with look-alikes of unknown edibility.
COMMENTS: The salient features of this brown cup fungus are its medium to large size,
growth on wood or wood chips, light brown color, and smooth spores. There are several
similar species, including: P. varia, with minutely roughened spores and a slightly grayer
fertile surface, widespread on rotting wood or occasionally in basements;P. emileia, with
minutely roughened spores and an ochre-brown fertile surface; andP. badioconfusa(see
comments under P. sylvestris), often found on or near rotting conifers. See also P. domi-
ciliana, which prefers to grow in domestic situations, and P. sylvestris, which usually grows
on the ground. Other species: Pachyella (formerly Peziza) grows on soggy logs or stumps,
but has a somewhat gelatinous-rubbery fruiting body that is broadly attached to the
substrate (rather than just at the center) and is usually flattened (disclike) or shallowly
cuplike. The most common species, Pachyella clypeata, grows 2-8 cm broad and has a
brown to chestnut-brown fertile surface. Pachyella babingtonii is minute (1-5 mm) and
cushion-shaped, reddish-brown to purplish-brown, and often somewhat translucent.

Peziza domiciliana (Domicile Cup Fungus) Color Plate 207

FRUITING BODY 2-10 cm broad or occasionally larger; cup-shaped when young and
often with a short stalk, becoming flattened or wavy in age but often retaining a central
depression (i.e., umbilicate); outline frequently irregular or somewhat angular in old age.
Fertile (upper) surface at first white or buff, but often darkening in age to tan or brown;
smooth to slightly wrinkled. Exterior(underside) same color or paler. Flesh rather fragile,
at times slightly waxy, sometimes turning yellow when broken. STALK often prominent
when young and inconspicuous or absent in age, short( when present) and stout, up to 1 cm
long, usually whitish. SPORES 11-15 * 6-10 microns, elliptical, smooth to very slightly
roughened, at times with two small oil droplets.
HABITAT: Solitary to gregarious or clustered on a wide range of domestic materials:
plaster, cement, sand, gravel, coal dust, carpets, fireplace ashes, etc. (It is said to favor
strongly alkaline substrates.) Hugo Sloane, Santa Cruz County’s incorrigible enfante
PEZIZA 823

tem'6/e-in-residence, has had specimens grow out of the wall above his bathtub—in two
different houses! It also grows in cellars, greenhouses, shower stalls, damp closets, under
porches, on wet rugs, behind refrigerators, around leaky water beds—and in my car!
EDIBILITY: If it were poisonous we would probably know by now, but I can find no
specific information on it.
COMMENTS: This white to tan or brown fungus is best identified by its propensity for
appearing in unexpected places. Several other cup fungi will occasionally grow indoors
(e.g., P. varia, P. petersii), but this one makes a habit of it. The color plate shows specimens
photographed in situ—in the carpeted “romper room” of a nursery school! It has been
shown that the fruiting bodies develop quite slowly, taking 3-5 weeks to mature. During
this time their shape, size, and color can change considerably.

Peziza vesiculosa (Common Dung Cup)

FRUITING BODY 2-8 cm broad, at first more or less round (spherical), then opening to
become cup-shaped with an inrolled (but often crimped or convoluted) margin. Fertile
(inner or upper) surface yellowish to yellow-brown to pale brown or buff, smooth or
wrinkled toward the center, sometimes darker brown in age. Exterior(underside) whitish
to buff or pale tan, minutely roughened or scurfy. Flesh rather fragile and soft. STALK
absent or present only as a narrowed basal point of attachment. SPORES 18-24 * 10-14
microns, elliptical, smooth, without oil droplets.
HABITAT: Solitary or more often gregarious (sometimes in large clusters) on manure,
dung, rotting straw, in corrals, around stables, gardens, and other fertilized areas, etc.;
widely distributed and common. In our area it fruits practically year-round, whenever it is
damp enough. I have seen massive clusters growing with Bolbitius vitellinus in a corral.
EDIBILITY: Not recommended. One source says it is poisonous unless well-cooked.
COMMENTS: This is one of our characteristic dung-inhabiting fungi. The habitat is its
most distinctive feature, and when it grows in fertilized soil it may be necessary to examine
it microscopically in order to distinguish it from P. repanda and other species. P.fimeti
also grows in dung, but is smaller (up to 2 cm) and dull brown, seldom grows in clusters,
and has smaller spores. For even more miniscule“dungcups,” see Cheilymenia coprinaria.

Peziza vesiculosa, a common dung lover. These are young, almst spherical specimens. As they mature
they will open out or even become flat; those growing in clusters will often be distorted.
824 PEZIZACEAE & ALLIES

Peziza violacea (Violet Cup Fungus)

FRUITING BODY 1 -3 (4) cm broad, at first nearly round but soon becomingcup-shaped,
then expanding to shallowly cup-shaped or disclike; often somewhat irregular in old age
with the margin splitting. Fertile (upper) surface smooth or slightly wrinkled and often
depressed at the center, violet to reddish-violet, often darker in age. Exterior (underside)
pallid to grayish or tinged violet, delicately powdered at least near margin. Flesh brittle,
thin, tinged violet. STALK absent or present as a short, narrowed base (especially when
young). SPORES 16-17 * 8-10 microns, elliptical, smooth, without oil droplets.
HABITAT: Solitary to gregarious on burnt ground (forest fire and campfire sites, etc.);
widely distributed, but not very common. In our area this species and P. praetervisa {ste-
comments) fruit in the winter and spring; elsewhere I’ve seen them in spring and summer.
EDIBILITY: U nknown, or at least I can find no information on it.
COMMENTS: This is one of several cup fungi that grow almost exclusively on burnt
ground. The violet color is distinctive, although another charcoal-lover, P. praetervisa,
is also violet (but usually darker: deep purple to purple-brown), with smaller, roughened
spores. P. proteana also likes ashes, but is paler (white to pinkish). For smaller charcoal-
lovers, see comments under Cheilymenia coprinaria and Geopyxis vulcanalis.

Peziza proteana (False Sparassis)

FRUITING BODY 1-6 cm broad and cup-shaped to disclike in the typical form, but
forming large cabbage-like clumps 10-30 or more cm in diameter in form sparassoides, the
“cups” in these fruiting bodies much distorted by mutual pressure and usually lopsided,
spoon-shaped, or completely misshapen (especially at center of clump). Fertile surface(s)
smooth or wrinkled, whitish to tan, often with a pinkish or lilac tinge; margin( s) often wavy.
Exterior same color or slightly paler or darker, or often lilac-tinged at or toward the base;
often slightly scurfy. Flesh thin, brittle. STALK absent or rudimentary. SPORES 10-13
* 4.5-7 microns, elliptical, minutely roughened, usually with two small oil droplets.
HABITAT: Solitary to gregarious or clustered on ground in woods, usually in burned
areas; widely distributed but not particularly common (form sparassoides is rare). I have
found it only twice in our area, in the winter and spring.
EDIBILITY: Edible if cooked thoroughly. One book lists form sparassoides as “choice.”
COMMENTS: The typical form of this cup fungus can be recognized by its pale color and
preference for burnt ground (it is not normally as dark or as purple as P. violacea and P.
praetervisa). The cabbage-like form was originally placed in a different genus, but is now
thought to be a growth form of P. proteana in which the “cups” are “bent out of shape” by
overcrowding( wouldn’t you be?). This form superficially resembles the cauliflower mush¬
room (Sparassis), but is differently colored, more brittle in texture, and bears its spores
in asci rather than on basidia. It has been suggested that “sparassioid” Pezizas such as
P. proteana link the hordes of regular cup-shaped species to the rare, convoluted, truffle¬
like forms (e.g., P. ellipsospora).

Peziza sp. (unidentified) (Truffle-Like Peziza)

FRUITING BODY 0.5 -5 or more cm broad, usually buried in ground, but sometimes
partly exposed; ranging from simple and cuplike with a strongly inrolled margin to com¬
plex and lobed (a loosely joined mass of folded chambers). Exterior smooth or finely hairy,
white to creamy when fresh, yellowing when handled and developing ochre to orangish to
rusty-yellow stains in age (and old or dried areas often becoming brown or reddish-
brown). Interior (fertile surface) sometimes intricately folded, with one to several large
PEZIZA 825

chambers which open to the exterior; colored like exterior. Flesh fragile; odor usually
sweet in age. STALK absent. SPORES 10-17 * 9-14 microns, broadly elliptical, minutely
roughened at maturity. Asci 8-spored, forming a palisade that lines the interior.
HABITAT: Solitary, scattered, or in groups under oak and other trees, usually growing
on the surface of the soil beneath the humus layer, but sometimes partially exposed; fairly
common in our area in the winter and spring.
EDIBILITY: Unknown, but too fragile to bother with.
COMMENTS: This is one of several truffle-like Pezizas. The internalizing of the fertile
surface is an obvious adaptation to growing underground (see chapter on truffles). The
above description is based on material collected near Santa Cruz, California, where it is
a common species (see photo on p. 822). It is apparently unnamed, although it approaches
P. (formerly Hydnotrya) ellipsospora, a sometimes large Californian species described as
being “purplish to brown” with slightly larger spores. Other truffle-like Pezizas include:
P. stuntzii, discovered under conifers in Washington, an aromatic species whose solid
interior is marbled with brown veins; andP. gautierioides, also found in the Pacific North¬
west, but growing quite deep in the soil (up to 6 inches down!), usually in the spring. All
of these can be confused with other underground Ascomycetes(e.g., Hydnotrya), but their
amyloid or amyloid-tipped asci place them in Peziza.

SARCOSPHAERA
THIS distinctive genus includes a single widespread species, described below. It usually
develops underground as a hollow ball, but when it surfaces the wall usually splits into
several lobes at the top to form a crown. In the underground stage it is apt to be mistaken
for a truffle, and was once given its own truffle genus, Caulocarpa. It is now placed along¬
side Peziza in the Pezizaceae because its asci have amyloid tips.

Sarcosphaera crassa (Crown Fungus)

FRUITING BODY beginning as a hollow, round to flattened orlobed ball 3-10 cm broad,
usually with a “soft spot” or slight depression at the top; wall usually splitting at maturity
from this spot downward to form several (usually 6-10) pointed segments or rays which
open up part way to form a deep crownlike cup; 5-20 cm broad when expanded. Fertile
(inner) surface smooth or breaking into fine scales, usually whitish to grayish at first, but
becoming grayish-pink to pinkish, lilac, purple, or purple-brown in age (especially after
splitting open). Exterior whitish to creamy, scurfy or roughened, usually dirt-incrusted.
Flesh rather thick but brittle, whitish. STALK usually (but not always) present as a short,
thick, narrowed base up to 3 cm long. SPORES 14-22 ><7-9 microns, elliptical with blunt
or truncate ends, smooth or very slightly roughened, with 1-3 (usually 2) oil droplets.

Sarcosphaera crassa, rather small specimens. This distinctive fungus begins as an underground ball
(right) which is hollow inside (center). Eventually, however, it splits at the top into starlike rays (left).
Sarcosphaera crassa. Left: Large, young specimens that are just beginning to split into rays. Right:
Older specimens. Large one at top has flattened out more than is usual. Flesh is thick but fragile.

HABITAT: Solitary to gregarious or in clusters of 2-5 individuals, developing at or below

ground level but usually exposed or partly exposed at maturity; widespread, but especially
common under western conifers. It is often found in the spring (while morel hunting), but
also fruits in the summer and fall. It is common under pine and other conifers in the Sierra
Nevada, Cascades, and Rocky Mountains, but I have yet to find it in our area.
EDIBILITY: Not recommended. It is rated highly by some but is difficult to clean and a
few people are adversely affected by it. Cook it thoroughly if you decide to try it. O.K.
Miller describes the texture as “a little like a rubber eraser that’s been softened by time.”
COMMENTS: Also known as S. eximia and S. coronaria, this is a curious and highly
distinctive fungus. Specimens which haven’t split open are likely to be mistakenfortruffles
(especially if they’re growing underground), but are easily distinguished by their com¬
pletely hollow interior. Older individuals, on the other hand, might be confused with
Scleroderma geaster, but can be told by their pinkish to purplish color. Other species:
Peziza ammophila is somewhat similar in shape when mature, but has a brown to dark
brown interior and is cup-shaped when young. It grows in sand dunes orsandy soilaround
the world, but is not common. Geopora species can also be similar, but have fuzzy brown
hairs on the exterior and are not pinkish- or purple-tinged. Neournula pouchetii has a
pinkish fertile surface, but is smaller and always cuplike (see key to Sarcosoma & Allies).

SARCOSOMA & Allies

Small to medium-large fungi found on ground or rotten wood. FRUITING BODY variously
shaped but usually cuplike to urnlike or top-shaped. Fertile (upper or inner) surface usually dark
brown to black, sometimes paler but not brightly colored. Exterior (sterile surface) often but not
always minutely hairy. Flesh gelatinous or very rubbery, or if not then rather tough (i.e., not
breaking easily). STALK present or absent. SPORES elliptical to round, smooth or roughened,
with or without oil droplets. Asci lining upper surface of fruiting body or interior of cup, mostly
8-spored, operculate, with non-amyloid tips.

THESE dark, tough to rubbery or gelatinous cup fungi are not necessarily cup-shaped.
The paler, non-gelatinous types can usually be separated from Peziza and Helvetia by
their tougher texture. The thickly gelatinous ones are more apt to be mistaken for jelly
fungi, but produce their spores in asci and are usually darker or differently shaped.
Sarcosoma, with gelatinous flesh, and Plectania and Urnula, with tough flesh, are the
three principal genera treated here. Along with a few other genera they comprise the
family Sarcosomataceae. They are saprophytic on humus and dead wood, and like most
Ascomycetes, are especially prevalent in the spring. Although distinctive, they are unpala¬
table except in an emergency because of their texture. Four species are described here and
others are keyed out, including gelatinous relatives of the earth tongues (Helotiales).
826
SARCOSOMA & ALLIES 827

Key to Sarcosoma & Allies

l. Fruiting body an erect, hollow club or closed urn that splits lengthwise from the top to form
several rays; fertile (inner) surface pallid to yellowish, exterior brown and hairy; found on or
near dead hardwoods; southern (fairly common in Texas) .... Choriactis (-Vrnula) geaster
1. Not as above . 2
2. Fruiting body shallowly cup-shaped or ear-shaped or like a piece of seaweed, dark brown to
reddish-brown or purplish but not normally black unless it dries out, 2-10 cm broad; flesh thin
and rubbery or rubbery-gelatinous; sterile surface (exterior) minutely hairy; spores borne
on basidia which often (but not always) line the lower surface of the fruiting body.
. (see Auricularia auricula, p. 675)
2. Not as above; spores borne in asci which line the upper surface of fruiting body. 3
3. Fruiting body rounded and nearly closed becoming shallowly cup-shaped in age; stalk short or
absent; fertile (upper or inner) surface brown to reddish-brown; exterior with a dense covering
of dark hairs; flesh thick and rubbery or gelatinous; found in groups or clusters on hardwood
sticks and branches in eastern North America and tropics (one report from California) ....
. Galiella (-Bulgaria) rufa
3. Not as above . 4
4. Fruiting body 1-5 mm broad or if larger then shallowly cup-shaped to disclike (flat) at maturity
(not top-shaped), stalkless, and broadly attached to the substrate (i.e., only the margin free);
flesh rather waxy; found on wet logs; asci with amyloid tips . . . (seePeziza&. Allies, p. 818)
4. Not as above . 5
5. Fruiting body top-shaped to cup-shaped or irregular and very rubbery or gelatinous; flesh
usually gelatinous and sometimes translucent, at least when young and fresh . 6
5. Not as above; fruiting body usually cup-shaped or urnlike or occasionally disclike, sometimes
with a long stalk; flesh tough to slightly rubbery, but not usually gelatinous . 9
6. Growing on hardwoods (usually dead); fruiting body cylindrical to top-shaped to shallowly
cup-shaped and usually less than 4 cm broad or if larger then fruiting body lobed or irregular
(like a jelly fungus) and not black; asci not operculate .(see Helotiales, p. 865)
6. Not as above; growing on ground or dead wood, usually under conifers; fruiting body round to
cylindrical to top-shaped or shallowly cup-shaped, dark brown to purple-brown to black,
2-12 cm broad when mature; asci operculate(i.e., with “lids”) .7
7. Found in eastern North America (rarely in West?); base of fruiting body usually as broad as the
top.Sarcosoma globosum (see S. mexicana, 828)
7. Found in western North America; base usually tapered (narrower than top). 8
8. Fruiting body 4-12 cm or more broad when mature; flesh thickly gelatinous; mature spores with
1-3 oil droplets .Sarcosoma mexicana, p. 828
8. Fruiting body 2-5 (7) cm broad; flesh gelatinous when young but often less so in age; mature
spores lacking obvious oil droplets . Sarcosoma latahensis (see S. mexicana, p. 828)
9. Stalk present . 10
9. Stalk absent or rudimentary . 16
10. Fertile (inner or upper) surface more or less pinkish-brown; margin of cup scalloped or lobed;
stalk whitish; found in northern North America, usually under conifers; rare .
.Neournula (-Vrnula) pouchetii
10. Not as above; fertile surface usually brown to black, at least in age . 11
11. Fertile surface tan to brown, yellow-brown, or olive-brown (but may blacken in age) .... 12
11. Fertile surface dark brown to black, even when young . 13
12. Exterior (underside of cup) wtih sparse brown hairs; stalk usually ribbed or appearing
“gathered” .(see Aleuria & Allies, p. 833)
12. Underside black or nearly so; not as above .. . Plectania melaena (see P. nannfe/dtii, p. 830)
13. Fruiting body urn-shaped to deeply cup-shaped, 4-12 cm high and 3-10 cm broad; found mainly
on hardwood sticks and logs in eastern North America (also reported from Alaska) .
. Vrnula craterium & others, p. 829
13. Not as above; fruiting body shallowly cup-shaped or smaller than above; widespread .... 14
14. Stalk 2-6 cm long, 2-4 mm thick, not ribbed; fruiting under mountain conifers in the spring,
when or shortly after the snow melts .Plectania nannfeldtii & others, p. 830
14. Not with above features . 15
Sarcosoma mexicana. Specimen on right has been sliced open to show the thickly gelatinous interior.
Specimen on left shows the wrinkled blackish exterior.

15. Stalk well-developed, 2-5 (7) mm thick; cup fleshy or fragile; exterior of cup lacking orangish
granules; spores with one large central oil droplet .(see Helvetia, p. 805)
15. Not as above; fruiting body tough, not breaking easily; stalk usually present as a narrowed base
beneath cup; exterior usually black, sometimes with orangish granules .
. Plectania melastoma, p. 829
16. Fruiting body small and shallowly cup-shaped to disclike, usually less than 1 cm broad; usually
found on ground in association with a minute orange cup fungus called Byssonectria
aggregata .Nannfeldtiella aggregata
16. Not as above . 17
17. Found on burnt ground or in dung, or if not, then flesh fragile . 18
17. Not as above; flesh usually tough .Plectania melastoma & others, p. 829
18. Fruiting body small to minute; found on dung or burnt ground, sometimes in swarms .
.(see Aleuria & Allies, p. 833)
18. Fruiting body usually larger than 1 cm broad; found on ground or in scorched areas but not in
dung .(see Peziza & Allies, p. 818)

Sarcosoma mexicana (Starving Man’s Licorice; Giant Gel Cup)

FRUITING BODY 5-10 (20) cm broad and 3-10 (15) cm high, rubbery, sometimes cup¬
shaped but more often more or less top-shaped (i.e., with a narrowed base and broader,
flattened to slightly concave “cap”); margin often lobed. Fertile (upper) surface black or
sometimes dark brown. Exterior dark gray to black, finely velvety (especially above),
usually narrowed below to form a thick “stalk” which is often deeply wrinkled or ribbed or
has large pockets. Flesh (interior of fruiting body) a thick, watery-gelatinous mass, clear
gray to black or brownish. STALK usually present as a narrowed base (see above).
SPORES 23-34 * 10-14 microns, elliptical to somewhat sausage-shaped, smooth, with
one to three oil droplets.
HABITAT: Solitary to gregarious or clustered on rotting wood or duff under conifers;
fruiting in the late winter, spring, summer, and early fall. It is known only from western
North America and Mexico, and has been characterized as “rare.” However, it is some¬
times very common (along with S. latahensis—see comments) in the mountains of Oregon
and northern California during the morel season (May and June) or shortly after the snow
melts. I have not seen it in our area, but S. latahensis occurs occasionally.
EDIBILITY: Unknown. As its common name implies, you would really have to be hungry
to be tempted by it.
COMMENTS: Originally known as Bulgaria mexicana, this is one of my fifty “five
favorite fleshy fungal fructifications.” Its black color, thickly gelatinous flesh (best seen
by slicing it open lengthwise as shown in above photo), and growth with conifers make it
virtually unmistakable. It is muchlargerthan“PoorMan’sLicorice”(Bulgariainquinans),
and is associated with conifers rather than hardwoods (at least in my experience). S. lata¬
hensis is a similar but slightly smaller (2-7.5 cm broad), purple-brown to black species

828
 ft

Sarcosoma latahensis (see comments under S. mexicana, which resembles it very closely). Specimens
on right and left are being viewed from the top; those in center, from the side (the one at the bottom
has been sliced open to show the gelatinous flesh).

that is common under western conifers, particularly at higher elevations. Its flesh is
gelatinous when young but often becomes tougher and less gelatinous in age, plus its
spores lack oil droplets at maturity. Another species, S. globosum, may occur in the West
but is much more common in eastern North America. It is massive, black, and gelatinous
like S. mexicana, but its water-and-gel-filled base is very broad (not tapered) and may
“leak” when collected!

Urnula craterium (Crater Cup; Devil’s Urn)

FRUITING BODY 4-12 cm high and 3-6 (10) cm broad, usually urn-shaped (i.e., with a
narrowed base or stalk); upper portion at first closed, then opening to form a deep cup or
“urn”; margin remaining incurved for some time, usually scalloped, sometimes torn at
maturity. Fertile (inner) surface smooth or scurfy, dark brown to black. Exterior variable
in color: dull pinkish-gray to dark brown and scurfy at first, often smoother and blacker
in age. Flesh tough and fibrous or leathery, black or very dark. STALK 2-4 cm long,
5-10 mm thick, continuous with and colored like exterior of cup, or darker; usually with
a patch of dark brown to black mycelial hairs emanating from the base. SPORES
24-36 x 10-15 microns, elliptical to spindle-shaped, smooth.

HABIT AT: Solitary or more often in groups or clusters on or near rotting hardwood sticks
and logs (the wood often buried); common in eastern North America in the spring.
EDIBILITY: Too tough to be worth eating.
COMMENTS: This dark brown to black urn-shaped fungus is one of the first fungi to
appear each spring in the hardwood forests of eastern North America. It differs from
Plectania melastoma by its deeper cup and better developed stalk, but the two species
might just as well be merged into one genus. I can find no record of it from the west coast,
but a similar species, U. hiemalis, has been found in Alberta and Alaska. It might con¬
ceivably be mistaken for a Craterellus, but the growth in the spring plus the fertile upper
or inner (rather than lower or outer) surface and slightly different shape distinguish it.

Plectania melastoma (Black Cup Fungus)

FRUITING BODY 1-2.5 cm broad and 1-3 (4) cm high; at first nearly round (or with a
stemlike base), then opening slowly at the top and eventually becoming more or less cup¬
shaped; margin usually remaining incurved for a long time but sometimes splitting in
places. Fertile (upper or inner) surface smooth or sometimes with veinlike markings when

829
Plectania melastoma. This small black cup fungus can be recognized by its color, tough texture,
and frequently wrinkled underside. Note how young specimens (right) are nearly closed at the top.

dry, often glistening when wet, black or deep brownish-black. Exterior tough (but with
a semi-gelatinous inner layer when wet), minutely hairy, strongly wrinkled or veined,
black, but often with a rusty-orange tinge near the margin from the presence of minute
orange granules. Flesh tough or cartilaginous, not brittle. STALK absent or present as
a short, stout, narrowed base; continuous with and colored like exterior of cup, the base
often with wiry black mycelial threads that extend into the substrate. SPORES 20-28
* 8-12 microns, elliptical or spindle-shaped, smooth, with oil droplets when immature.
HABITAT: Solitary or more often in small groups or clusters on decaying sticks and
other debris of both hardwoods and conifers; widely distributed and not uncommon, but
easily overlooked, fruiting mainly in the spring. In our area this species or something very
similar (see comments) occurs under oak in the late winter and early spring. I usually find
it when I’m foraging for Craterellus cornucopioides, perhaps because both are black.
EDIBILITY: Unknown
COMMENTS: The black color, wrinkled exterior (or underside), small size, and tough
texture are the principal fieldmarks of this attractive cup fungus. The typical form with
rusty-orange granules usually grows under conifers. Our local oak-loving version usually
lacks visible granules, but orange granules can be seen under the microscope. Bulgaria
melastoma is an older name for it. Two very similar black cup fungi with little or no stalk
and no orange granules are also worth mentioning: P. milled has a stellate margin (i.e.,
with starlike points) and elliptical spores, while P. (-Pseudoplectania) nigrella has round
spores. Both usually occur under conifers. See also Urnula craterium, which has a larger,
deeper (urnlike) fruiting body.

Plectania nannfeldtii (Black Snowbank Cup Fungus)

FRUITING BODY consisting of a shallow cup mounted on a well-developed, slender
stalk. Cup 0.5-2 (3) cm broad, the margin at first incurved. Fertile (inner or upper) surface
smooth, black. Exterior also blackish but delicately hairy. Flesh black, rather tough or
cartilaginous. STALK always present and typically rather long and slender, 2-6 cm long,
2-4 mm thick, more or less equal, not ribbed, black, often with black mycelium at base.
SPORES 21-30 (35) x 10-14 microns, elliptical, smooth or slightly roughened, without
oil droplets.
HABITAT: Solitary to gregarious in duff and debris or on buried or rotten wood under
conifers (particularly fir and spruce); fruiting in the spring shortly after the snow melts
or even developing under the snow. It is known only from western North America and is
fairly common at higher elevations (but easily overlooked). I have not seen it in our area.
EDIBILITY: Unknown.
COMMENTS: This attractive black cup fungus can be distinguished in the field by its
color, small size, well-developed stalk, and growth in the spring under conifers. Helvetia

830
Plectania nannfeldtii. This small black cup fungus has a long thin stalk and often grows near snow.

corium (see comments under H. macropus) is somewhat similar but larger, not as tough,
and often shows ribs on the stalk. P. (-Pseudoplectania) melaena also grows in the spring
under conifers, but is slightly larger and has a yellow-brown to olive-brown cap or cup
when young (but blackens in age) and a short or long stalk. For species with little or no
stalk, see Plectania melastoma.

OTIDEA
Medium-sized, mostly terrestrial fungi. FRUITING BODY usually erect or semi-erect (often
standing on one end), usually lopsided and open or slit on one side, sometimes shaped like a rabbit's
ear, at other times more cuplike. Fertile (inner) surface variously colored, usually smooth. Exterior
smooth or scurfy but not usually hairy. Flesh usually rather brittle, not gelatinous. STALK absent
or present as a short, thick base. SPORES elliptical, smooth, usually with two oil droplets. Asci
lining inside or upper surface of fruiting body, typically 8-spored, operculate, tips not amyloid.

THIS is a small but common group of lopsided or earlike cup cungi. The fruiting bodies
are usually erect (i.e., they stand on end) and slit down one side, and often occur in groups
or contorted clusters. Many other cup fungi can be open on one side or lopsided, but Otidea
is the only common genus that is consistently slit or lopsided and erect or semi-erect.
The dozen or so North American Otideas are differentiated primarily on microscopic
characters, but can be divided into two groups based on their shape. One group has an
elongated, erect, rabbit-ear-like fruiting body; the other is more open and cuplike and
often truncate (chopped off) at the top. To avoid otidealogical debate, only one species
from each group is described here. Otidea belongs to the Pyronemataceae.

Key to Otidea
1. Fruiting bodies arising in groups or clusters from an underground “tuber” (sclerotium), shaped
more or less like rabbit ears, 5-15 cm tall; fertile (inner) surface orange to pinkish to reddish-
brown, often blackening in age; exterior dark brown to blackish; found under hardwoods in
eastern North America, usually in the summer; rare, but often occurring in spectacular
numbers when it fruits .Wynnea americana
1. Not as above; fruiting bodies not arising from an underground “tuber” . 2
2. Fruiting body bright yellow to orange but usually developing dark bluish or greenish stains;
found mainly in spring under northern and mountain conifers (seeCaloscyphafulgens, p.837)
2. Not as above . 3
3. Fertile (inner) surface of fruiting body pale yellow to orange, ochraceous, or pinkish .4
3. Fertile surface predominantly tan to brown, sometimes with brighter areas . 7

831
832 PEZIZACEAE & ALLIES

4. Fruiting body pale to bright yellow, usually with a truncate (broad, flattened) apex .
. O. concinna & O. cantharella (see O. onotica, below)
4. Not as above . 5
5. Fertile surface bright orange; underside usually whitish; fruiting bodies only sometimes lop¬
sided or slit down one side (particularly when clustered) .(seeAleuria & Allies, p. 833)
5. Not as above . 6
6. Fertile surface dull orange to yellowish or tinged pinkish.O. onotica, below
6. Fertile surface yellowish, the exterior usually brownish . . O. leporina (see O. onotica, below)
7. Fruiting bodies typically elongated and erect (standing on end like rabbit ears) . 8
7. Fruiting bodies typically broadened and flattened somewhat on top (truncate) or cuplike or
distorted (especially when clustered); usually growing semi-erect . 9
8. Fertile surface yellowish-brown.O. leporina (see O. onotica, below)
8. Fertile surface usually deep reddish-brown to vinaceous-brown .
.O. smithii <fe Wynnella silvicola (see O. alutacea, below)
9. Fruiting bodies often nearly round (spherical) when young and flattened in age, usually solitary
or in small groups but not often clustered; tips of asci amyloid . (see Peziza & Allies, p. 818)
9. Not as above; fruiting bodies often clustered and distorted by mutual pressure, not normally
flattening out in age nor spherical when young; asci not amyloid O. alutacea & others, below

Otidea alutacea (Brown Clustered Ear Cup)

FRUITING BODY 2-6 cm high and 2-4 cm broad, shape variable: usually cup-shaped but
lop-sided (the shorter side split lengthwise or open) and semi-erect, but often irregularly
wavy or contorted when growing in clusters; apex often truncate when growing erect.
Fertile surface (interior) smooth, tan or light brown to grayish-brown or brown. Exterior
often slightly scurfy, pale to dull brown (or yellowish in one variety). Flesh brittle. STALK
absent or present as a narrowed, whitish, downy base. SPORES 14-16 x 7-9 microns (or
smaller in one variety), elliptical, smooth, typically with two oil droplets.
HABITAT: Scattered to densely clustered in forest humus, usually under conifers; fairly
common in western North America. In our area this species and its look-alikes (see
comments) is frequent in the fall and winter, particularly under Douglas-fir and oak.
EDIBILITY: Unknown—better chucked than plucked.
COMMENTS: The brownish color, lopsided fruiting bodies that are often contorted,
broadened at the apex, and split down one side, plus the tendency to grow in dense clusters
are the distinguishing features of this undistinguished fungus. There are a number of
similar brownish Otideas, including: O. bufonia, dark brown, usually growing in clusters;
O. rainierensis, found in the Pacific Northwest and normally not clustered; O. abietina,
medium to dark brown and often cup-shaped, but with much larger spores (18-22 microns
long); and O. grandis, brown but frequently with reddish-orange patches on the fertile
surface and a thicker, more prominent stalk. There are also several brown Otideas with
an elongated, erect, spoon-shaped to earlike (not truncate or cup-shaped) fruiting body
like that of O. onotica. These include: O. smithii, deep reddish-brown to vinaceous-brown
and fairly large (up to 8 cm high), with a thick stalklike base, fairly common under conifers
in the Pacific Northwest and northern California; and Wynnella silvicola (-O. auricula),
a northern species that is colored like O. smithii but has larger spores (22-25 microns long).

Otidea onotica (Donkey Ears)

FRUITING BODY (3) 5-10 cm high and (2) 4-6 cm broad (more if completely expanded);
shape variable but usually spoon-shaped or like an elongated ear (standing erect on one
end with one side open or slit); margin at first inrolled, but expanding somewhat in age.
Fertile surface (interior) smooth, ochraceous to dull orange to orange-buff or yellowish,
often with a pinkish or rosy tint when fresh. Exterior similarly colored but without a
Left: Otidea alutacea, rather pale (tan) specimens. Note rather erect growth habit. Right: Otidea
onotica. Note erect growth habit and also how the fruiting body is slit down to the base on one side.

pinkish tinge, often slightly scurfy. Flesh thin, pallid, brittle. STALK present as a whitish,
narrowed, hairy or downy base that arises from a litter-binding mycelium. SPORES
12-14 x 6-8 microns, elliptical, smooth, with two oil droplets. Paraphyses (sterile cells)
strongly hooked.
HABITAT: Scattered or more often in groups or clusters under both hardwoods and
conifers; widely distributed. In our area it fruits in the winter and spring, but is not as
numerous as O. alutacea.
EDIBILITY: Edible according to some, but one study revealed the presence of the toxin
MMH. In other words, it is better chucked than plucked.
COMMENTS: The erect growth habit and earlike fruiting body plus the ochraceous or
orangish to pinkish-tinged fertile surface separate this species from most other cup fungi.
The apex of the fruiting body is not broadly flattened or truncate as is typical of O. alutacea.
Other species: O. leporina (“Rabbit Ears”) has a yellowish-brown interior and brownish
exterior, but is otherwise quite similar. O. concinna and O. cantharella are both pale
to bright yellow, but often have a broadened or truncate apex. All of these species are
widely distributed. For duller or browner Otideas, see comments under O. alutacea.

ALEURIA & Allies

Mostly small to minute fungi found on dung, soil, moss, wood, foliage, and ashes. FRUITING
BODY usually cup-shaped to disclike (flattened) or cushion-shaped, often gregarious. Fertile
(upper) surface often brightly colored, usually smooth. Sterile surface (underside) often hairy, but
sometimes bald. Flesh usually fragile, but sometimes rather tough. STALK absent or present.
SPORES round to elliptical or elongated, smooth or roughened, with or without oil droplets,
thick-walled when immature in Ascobolaceae, otherwise thin-walled. Asci lining upper surface
of fruiting body, typically 8-spored, operculate, not amyloid; thin-walled in Pyronemataceae
and Ascobolaceae, thick-walled in Sarcoscyphaceae.

THIS is a motley multitude of miniscule to medium-sized cup- and disclike fungi with
non-amyloid, operculate asci and fragile to fairly tough but not gelatinous flesh. In
contrast to Peziza, many of the species are brightly colored, and those that aren’t are
usually small. A few, such as the common orange peel fungus, Aleuria aurantia, are
conspicuous, but most are so minute or mundane that only the most avid devotees of
diminutiveness (see p. 224) will notice them.

833
834 PEZIZACEAE & ALLIES

These cup fungi, like others, grow in a wide range of habitats. Many digest dung; others
occur in swarms on scorched earth and were among the first organisms to colonize the ash-
covered wasteland around Mount St. Helens after that volcano erupted. Space permits
descriptions of only a few species, nearly all of which belong to the Pyronemataceae (not
to be confused with the Pyrenomycetes or flask fungi). Exceptions are the scarlet cup
fungus, Sarcoscypha coccinea, and its relatives, which are placed in a separate family, the
Sarcoscyphaceae, because of their tougher texture and a number of microscopic dif¬
ferences. A third family, the Ascobolaceae, is only briefly mentioned here. Most of its
constituents are minute, dark-spored dung addicts of interest only to scatologists, asco-
mycologists, students of Fungi 112B, and other assorted eccentrics who deal with dung
on a daily basis.
Key to Aleuria & Allies
l. Growing on dung or manure . 2
1. Not as above . 3
2. Fruiting body typically at least 1 cm broad when mature, nearly round (spherical) when young
becoming cup-shaped to nearly flat in age, yellowish to brown, without prominent hairs;
asci with amyloid tips .(see Peziza & Allies, p. 818)
2. Not as above; fruiting body small or minute, often with hairs .
.Cheilymenia coprinaria & many others, p. 838
3. Exterior (underside) of cup or disc clothed with brown to black hairs; margin often fringed
with dark hairs also . 4
3. Not as above; exterior either hairless or with white or pale hairs . 12
4. Most or all of fruiting body immersed in the ground with only the top (mouth) showing,
making it look like a hole in the ground . (see Geopora, p. 846)
4. Not as above . 5
5. Fertile (upper or inner) surface yellow, orange, or red . 6
5. Fertile surface white to creamy, tan, gray, brown, etc. 9
6. Fruiting body small or minute, growing on burnt soil or charred wood .
.Anthracobia melaloma & others (see Cheilymenia coprinaria, p. 838)
6. Not as above; growing in soil, humus, or on wood but not usually in burned areas . 7
7. Fruiting body tough or corky and thick-fleshed; fertile surface orange to red or sometimes yel¬
lowish; exterior dark brown to black; found in eastern North America Wolfina aurantiopsis
7. Not as above . 8
8. Underside of fruiting body with minute brown hairs; fertile surface bright orange .
.Melastiza chateri(sec Scutellinia scutellata, p. 839)
8. Underside of fruiting body with fairly obvious hairs which fringe the margin like eyelashes;
fertile surface bright red to orange-red or orange .... Scutellinia scutellata & others, p. 839
9. Fertile (upper or inner) surface white to grayish; stalk absent or rudimentary . 10
9. Fertile surface white to grayish, brownish, or darker; stalk usually present (but often short) 11
10. Fruiting body typically 1-3 cm broad .Humaria hemispherica, p. 839
10. Fruiting body typically less than 1 cm broad .
. Trichophaea boudieri & others (see Humaria hemispherica, p. 839)
11. Stalk ribbed, often short; fertile surface white to tan or brownish but not gray .
.Jafnea semitosta (see Humaria hemispherica, p. 839)
11. Stalk not ribbed, or if so then fertile surface grayish or blackish .(see Helvetia, p. 805)
12. Growing on recently fallen branches or foliage of conifers; fruiting body small or miniscule,
stalkless, often brightly colored; asci operculate .
.Pithya vulgaris & others (see Sarcoscypha coccinea, p. 836)
12. Not as above . 13
13. Fertile (upper or inner) surface bright red to scarlet; exterior usually whitish or with white hairs;
usually growing on wood or buried sticks . 14
13. Not as above . 16
14. Fruiting body very small (less than 1 cm broad); exterior of cup with long hairs . 15
14. Fruiting body larger and/ or hairs on exterior short . . Sarcoscypha coccinea & others, p. 836
 ALEURIA& ALLIES 835

15. Fruiting body arising from an elongated rootlike structure (several often arising together from
the same “root”) . Microstomaprotacta (see Sarcoscypha coccinea, p. 836)
15. Not as above .Microstoma floccosa (see Sarcoscypha coccinea, p. 836)
16. Fruiting body yellow to orange but soon developing dark bluish to greenish or olive stains; found
under northern and mountain conifers in spring and early summer Caloscyphafulgens, p. 837
16. Not as above (but fruiting body may be yellow or orange) . 17
17. Fruiting body bright yellow, minute (less than 5 mm broad), occurring in swarms on wood; asci
inoperculate (without “lids”) .(see Helotiales, p. 865)
17. Not as above . 18
18. Fruiting body minute (1-6 mm broad) and disclike to cushion-shaped; growing mainly in burned
areas, often in swarms or masses . 19
18. Not as above; either larger or distinctly cuplike or consistently growing in other habitats . 20
19. Fertile (upper or inner) surface orange to yellow-orange to reddish .
.Pyronema omphalodes & others (see Cheilymenia coprinaria, p. 838)
19. Not as above; fertile surface usually darker .
.Ascobolus carbonarius & others (see Cheilymenia coprinaria, p. 838)
20. Fertile (upper or inner) surface bright orange to yellow-orange . 21
20. Not as above (but fertile surface may have a dull yellowish or pale orange tinge) . 23
. 21. Stalk absent or rudimentary; very common .Aleuria aurantia, p. 837
21. Stalk present, at least in many specimens . 22
22. Fertile surface bright orange to bright yellow-orange . Aleuria rhenana & others, p. 836
22. Fertile surface pale orange to pale or dingy yellowish .Geopyxis vulcanalis, p. 840
23. Growing in recently burned areas . 24
23. Not as above . 26
24. Fertile surface pink to reddish; stalk absent or rudimentary .
.Tarzetta rosea (see Geopyxis vulcanalis, p. 840)
24. Not as above; fertile surface differently colored (including brick-red) . 25
25. Fertile surface more or less brick-red . Geopyxis carbonaria (see G. vulcanalis, p. 840)
25. Not as above . 26
26. Fruiting body turquoise to blue-green or at least tinged those colors; found on wood
.(see Helotiales, p. 865)
26. Not as above; differently colored . 27
27. Fruiting body with a stalk that arises from a swollen tuberlike structure (sclerotium) immersed
in the substrate; asci inoperculate .(see Helotiales, p. 865)
27. Not as above . 28
28. Fruiting body very small (typically less than 7 mm broad); stalk when present very thin; often
found on living plant parts (leaves, stems, etc.); asci inoperculate . .. (see Helotiales, p. 865)
28. Not as above; usually found on ground or dead wood; asci operculate (with “lids”) . 29
29. Fruiting body tough or leathery, the flesh very thin; exterior usually with white or grayish
hairs; common on dead hardwood sticks, branches, etc; spores borne on basidia .
. . . .. (see Stereaceae & Allies, p. 604)
29. Not as above . 30
30. Fertile surface pale orange to pale or dingy yellowish; fruiting body usually less than 1.5 cm
broad . Geopyxis vulcanalis & others, p. 840
30. Not as above; fertile surface grayish-tan to tan or brown and/or fruiting body larger .... 31
31. Stalk present (at least in most specimens) . 32
31. Stalk absent or rudimentary . 34
32. Exterior of cup and upper stalk usually hairy or densely scurfy when fresh; fertile surface
often dark; stalk well-developed though sometimes short .(see Helvetia, p. 805)
32. Not as above . 33
33. Fruiting body 1-3 cm broad . Tarzetta catinus (see Geopyxis vulcanalis, p. 840)
33. Fruiting body typically 1.5 cm broad or less Tarzetta cupularis (see Geopyxis vulcanalis, p. 840)
34. Fertile surface dark brown to blackish; fruiting body usually saucer-shaped or disclike (flat)
in age; tips of asci amyloid .(see Peziza & Allies, p. 818)
34. Fertile surface brown to yellowish; tips of asci not amyloid .
.Tarzetta bronca (see Geopyxis vulcanalis, p. 840)
836 PEZIZACEAE & ALLIES

Sarcoscypha coccinea (Scarlet Cup Fungus) Color Plate 210

FRUITING BODY 2-5 (6) cm broad when mature, more or less cup-shaped, the margin
usually incurved, often tattered in old age. Fertile (inner or upper) surface bright red to
scarlet, sometimes fading to reddish-orange in age, smooth. Exterior whitish, covered
with minute hairs. Flesh thin but not particularly brittle. STALK absent or more often
present, up to 4 cm long, 3-7 mm thick; minutely hairy and white, tapered downward.
SPORES 24-40 x 10-14 microns, elliptical, smooth.
HABITAT: Solitary or in groups on buried or fallen hardwood sticks or branches; widely
distributed and fairly common in the winter and early spring (or late fall in some regions).
In our area I have found it in abundance in a riparian woodland composed of willow,
alder, buckeye, and cottonwood.
EDIBILITY: Said to be edible; I haven’t tried it.
COMMENTS: Formerly known as Plectania coccinea, this beautiful cold weather cup
fungus is easily told by its bright red fertile surface. The marginof the cup is not fringed with
dark hairs as in Scutellinia scutellata, and the exterior or underside is whitish. The length
of the stem seems to depend partly on how deep its food source (stick) is buried. Other
species: S. occidentalis is a similar but smaller (up to 1.5 cm broad) eastern species with
a well-developed (1-4 cm long) stalk and smaller spores. Microstoma (-Anthopeziza,
Sarcoscypha) floccosa is a minute (up to 1.5 cm high and 1 cm broad) bright red species
whose exterior is clothed with long white hairs. It is fairly common on downed sticks in
eastern North America, but I have not seen it in the V^cst.Microstomaprotacta(-Plectania
hiemalis) is somewhat similar to M. floccosa, but its stalk arises from a hard, elongated
rootlike structure and is often branched above, giving rise to up to a dozen bright red
cups. It is widely distributed but rather rare. Several closely related genera of bright red
to orange or pink cup fungi occur in the tropics, including Cookeina and Phillipsia,
with striate or banded spores. Finally, there are several small stalkless, disclike species
that grow on the recently fallen branches or foliage of conifers. These species include:
Pithya vulgaris, yellow to orange or reddish-orange, semi-gelatinous, and up to 1 cm
broad, usually found in the spring on branches of fir and other conifers; P. cupressina,
similar but not gelatinous, found on cedar or cypress; and Pseudopithyella miniscula,
a miniscule (1-2 mm) scarlet species.

Aleuria rhenana (Stalked Orange Peel Fungus) Color Plate 209

FRUITING BODY 1 -2 cm broad, cup-shaped with a stalk. Fertile (inner or upper) surface
bright orange to yellow-orange, smooth. Exterior white or whitish and minutely hairy or
downy. Flesh thin, brittle. STALK present, 1-3 cm long, 2-5 mm thick, slender, equal or
tapered downward, colored like the exterior; base often arising from a dense mass of white
mycelium that may bind several stalks together. SPORES 20-23 x 11-13 microns, ellip-

Aleuria rhenana is a small orange or yellow-orange cup fungus with a well-developed stalk. It often
grows in clusters, as shown here (at left) and in the color plate.
ALEURIA 837

tical, coarsely reticulate at maturity.

HABITAT: Gregarious, often in small clusters, on ground or moss in woods (usually
under conifers); widely distributed but infrequent. I have found it only twice—in Mt.
Rainier National Park in Washington, in September, and near San Francisco in January.
EDIBILITY: Presumably consumable, but much too small and rare to be of value.
COMMENTS: This petite cup fungus is the same color as its cosmopolitan cousin, A.
aurantia, but is much rarer and usually smaller, possesses a stalk, and likes to grow in
dainty clusters (see color plate). Leucoscypha (-Neottiella, Aleuria) rutilans is a similar
moss-inhabiting species with slightly larger spores and longer hairs on its underside.

Aleuria aurantia (Orange Peel Fungus) Color Plate 208

FRUITING BODY 1-10 cm broad, sometimes nearly round at first but soon becoming
cup-shaped to saucer-shaped to flattened or wavy, or sometimes irregularly contorted
(especially if clustered). Fertile (upper or inner) surface bright orange to golden-orange,
fading somewhat in age, more or less smooth; margin often wavy or lobed. Exterior
(underside) pallid or at least paler, smooth orminutelydowny. Flesh thin, brittle or fragile.
STALK absent or rudimentary. SPORES 18-24 * 9-11 microns, elliptical, coarsely
reticulate or ridged at maturity, typically with two oil droplets.
HABITAT: Scattered to gregarious or in fused clusters on ground, fruiting mainly in the
fall and winter in our area; widely distributed and very common. It seems to prefer bare
soil or sand along roads, paths, landslides, etc., but also grows in grass or moss.
EDIBILITY: Edible and highly rated by one authority, but bland according to others.
One of my colleagues uses it raw in salads, but it is so thin-fleshed and fragile that it hardly
seems worth the trouble to collect it.
COMMENTS: The orange peel fungus is most likely to be mistaken for one of the old
orange peels that frequently litter our woods and roadsides. It is much more fragile, how¬
ever, and less common. Its size and shape vary considerably depending on environmental
conditions, but the “aleuring” bright orange color and absence of a stalk are constant.
Some species of Otidea are orangish, but have a more erect rather than prostrate growth
habit, while A. rhenana is smaller and has a stqlk. A variety of A. aurantia with smaller
spores (13-15 microns long) occurs in our area. See also Melastiza chateri (under Scutel-
linia scutellata), a somewhat similar but smaller species with brown hairs on its exterior.

Caloscyphafulgens Color Plate 211

(Snowbank Orange Peel Fungus)
FRUITING BODY 1-4 (6) cm broad, sometimes nearly spherical when young but be¬
coming cup-shaped or flatter in age, sometimes slit down one side and appearing lop¬
sided; margin inrolled when young. Fertile (upper or inner) surface smooth or slightly
wrinkled, bright yellow-orange to orange, sometimes with dark bluish to olive-green
stains. Exterior hairless, colored like interior but usually with more pronounced blue or
greenish stains, especially toward margin. Flesh thin, brittle. STALK absent or present
only as a short, narrowed whitish base. SPORES 6-8 microns, round, smooth.
HABITAT: Scattered to gregarious or clustered in damp soil or duff under conifers,
fruiting in the spring and early summer shortly after the snow melts; widely distributed,
but especially common in the mountains of western North America. It is one of the charac¬
teristic spring mushrooms of the Sierra Nevada, Cascades, and Rocky Mountains.
EDIBILITY: I can find no information on it.
838 PEZIZACEAE & ALLIES

COMMENTS: The yellow-orange color and dark blue or greenish stains that make it
look like a moldy orange peel are the hallmarks of this springtime cup fungus. The my¬
celium apparently parasitizes the seeds of conifers (mainly spruce and fir) and clusters
of fruiting bodies often arise where squirrels stash their seed-containing cones. An albino
form of this species with bluish stains has been found in Idaho.

Cheilymenia coprinaria (Eyelash Dung Cup)

FRUITING BODY (1) 3-7 (10) mm broad, at first closed but soon opening to become
shallowly cup-shaped to disclike or somewhat cushion-shaped. Fertile (upper) surface
orange to pale orange becoming yellow or brownish in age, smooth; margin fringed with
minute dark brown hairs, often wavy. Exterior (underside) paler, also clothed with dark
hairs. Flesh thin. STALK absent. SPORES (14) 17-22 (25) x 8-12 microns, elliptical,
smooth, without oil droplets.
HABITAT: Solitary to densely gregarious on dung and manure or compost, etc.; fruiting
in wet weather, cosmopolitan (along with its numerous look-alikes) but seldom noticed
because of its small size.
EDIBILITY: Who knows? Who cares?
COMMENTS: This species is one of several small, difficult-to-distinguish, dung-loving
cup fungi. Although the hairs on the exterior are quite conspicuous, in the words of one
specialist, “they may be overlooked in the field, where the nature of the substrate dis¬
courages close scrutiny.” Similar yellow to orange dung-lovers include: C. theleboides,
with paler hairs (sometimes also growing on soil, humus, or “spent hops”); C. stercorea,
with branched dark brown hairs; Coprobia granulata, minute (1-2 mm), orange, and hair¬
less. Several similar disclike to cushionlike species occur in burned areas, often in vast
numbers, or in the heated (sterilized) soil in greenhouses. These include: Anthracobia
macrocystis, with a reddish fertile surface and brown hairs on its exterior; A. melaloma,
with a yellowish-brown to ochre-orange fertile surface and brown hairs on its exterior;
Trichophaea abundans, minute and whitish with pale brown hairs, growing on plaster as
well as burnt ground; Pyronema omphalodes, very common in confluent masses, with a
pale orange to reddish-orange, minute (1-3 mm), hairless fruiting body and elliptical
sores; Pulvinula carbonaria, also minute and pale to bright orange and hairless, but with
round spores; and Pulvinula archeri, similar to the previous species but with smaller
(7-9 microns), round spores. Also worth mentioning is Ascobolus, which usually has
a minute dark (greenish to dark brown or black), disclike fruiting body. Most of its
species grow on dung, but one, A. carbonarius, grows in swarms on burnt ground. For
small cup fungi that do not grow in dung or burned areas, see Scutellinia scutellata, and
for larger and more deeply cup-shaped, ash-loving or terrestrial species, see Geopyxis
vulcanalis. Better yet, go get some exercise!

Cheilymenia coprinaria in its favorite milieu—a “road apple” (piece of horse dung). Note small size,
disclike fruiting body, and the long hairs or bristles protruding from its margin.
Scutellinia scutellata typically grows in groups (left) and is easily recognized by its orange to red color
and the dark hairs that fringe its margin (right). Unfortunately, the hairs do not show up as well in these
black-and-whites as they do in the original color photographs. (Ray Gipson, Dan Harper)

Scutellinia scutellata (Eyelash Pixie Cup)

FRUITING BODY 0 2-1.5 cm broad, at first nearly round (spherical), but soon opening
to form a shallow cup and eventually disclike (flattened). Fertile (upper) surface smooth,
bright red to scarlet to orange (or rarely paler with a pinkish cast); margin conspicuously
ciliate (fringed with dark brown or blackish hairs up to 1 mm long). Exterior (underside)
also clothed with dark hairs. Flesh very thin. STALK absent. SPORES (15) 17-19 (23)
* (9) 11-14 (17) microns, elliptical, minutely warted, with one or more oil droplets.
HABITAT: Gregarious on rotten wood or damp soil (or occasionally on ashes* wet leaves,
or conks); widely distributed and common, but easily overlooked because of its small size.
In our area it fruits in the winter and spring.
EDIBILITY: Unknown, but much too puny to be of importance.
COMMENTS: This is another easily-recognized cup fungus. The bright red to orange
fertile surface and ciliate (eyelash-like) margin are good field characters. S. umbrarum
is a very similar, widespread, terrestrial species with a slightly larger fruiting body (up to 2
cm broad), larger spores, and shorter, less conspicuous hairs. S. erinaceus is also similar,
but is orange to yellow and smaller (2-5 mm broad), has smooth spores, and grows on
wood. Cheilymenia crucipila is a minute (1-4 mm) orange to orange-red, terrestrial species
with paler, shorter hairsand smooth spores that lack oil droplets. Lamprospora species are
minute and hairless. Finally there is Melastiza chaleri, a bright orange terrestrial species
that is 0.5-2 cm broad and has minute brown hairs on its exterior, especially near the mar¬
gin. For similar dung- and ash-lovers, see comments under Cheilymenia coprinaria.

Humaria hemispherica (Hairy Fairy Cup)

FRUITING BODY 1-3 cm broad, at first nearly round (spherical), gradually opening up
to become cup-shaped. Fertile surface (interior) white or whitish to grayish, smooth;
margin fringed with brown hairs. Exterior densely clothed with stiff brown hairs. Flesh
thin. STALK absent or present only as a rather abruptly narrowed base. SPORES 20-24
x 10-12 microns, elliptical, smooth or minutely warted, with 2 or sometimes 3 oil droplets.
HABITAT: Solitary, scattered, or in groups on ground or occasionally rotten wood;
widely distributed and fairly common under both hardwoods and conifers, usually
fruiting in the summer and fall. I have yet to find it in our area, but it may well occur.
EDIBILITY: Who knows? Who cares? Do you?
COMMENTS: The combination of pallid fertile surface and brown hairy exterior make

839
840 PEZIZACEAE & ALLIES

this an easy cup fungus to recognize. Jafneasemitosta is a larger (2-5 cm broad) species with
a creamy-white to tan or brown interior, a brown exterior clothed with scattered soft brown
hairs, plus a short ribbed stalk; it is fairly common in eastern North America. Other hairy
species: Trichophaea boudieri and T. bullata have a pale gray to whitish interior and
brown hairy exterior, but are much smaller (1-6 mm broad) and grow on wet soil under
conifers; T. abundans is a minute whitish species that grows in burned areas. For more
colorful hairy or ciliate species, see Scutellinia scutellata and Cheilymenia coprinaria.

Geopyxis vulcanalis (Vulcan Pixie Cup)

FRUITING BODY 0.3-1 (2) cm broad, nearly round (spherical) when young, becoming
deeply cup-shaped and then often flattening out in age. Fertile (upper or inner) surface
smooth, pale orange to pale or dingy yellowish, the margin usually finely scalloped. Ex¬
terior paler or whitish, usually powdery or downy when young but often entirely smooth
in age. Flesh thin, fragile. STALK usually present, up to 5 mm long and 1-3 mm thick,
sometimes so short as to be practically absent, equal or tapered downward, colored like
exterior of cup. SPORES 14-21 * 8-11 microns, elliptical, smooth, without oil droplets.
HABITAT: Scattered to densely gregarious in duff or moss under conifers, or in burned
areas; widely distributed. I have seen large fruitings locally in the fall, winter, and spring.
EDIBILITY: Unknown, but much too puny to be of value.
COMMENTS: This pixieish cup fungus and its close relatives are easily told by their small,
deeply cup-shaped (at least when young) fruiting body that often has a finely scalloped
margin. In G. vulcanalis a short stalk is usually present, but in some of the other species
(see below) it is lacking. G. carbonaria is a similar species that grows only in burned areas.
It has a brick-red fertile surface and normally remains cup-shaped rather than expanding.
Tarzetta species closely resemble Geopyxis in size and appearance, but as currently de¬
fined, have spores with two prominent oil droplets. Their ranks include: Tarzetta cupu-
laris, widespread under conifers, in moss, on burnt ground, etc., which resembles a
miniature goblet with its dainty stalk and grayish-tan to tan or brownish cup; T. rosea,
found in burned areas, with a pink to reddish cup and little or no stalk; and two somewhat
larger (1-3 cm broad), yellowish to brownish, woodland species: T. brortca of eastern
North America, with little or no stalk, and T. catinus, widespread (including California),
with a stalk. Most of these species have a scalloped margin or “lip” as in G. vulcanalis, and
were originally placed in Geopyxis, Pustularia, and/ or Peziza.

Geopyxis vulcanalis. Note small size, gregarious nature, and the small stalk below the cup which is
often covered by dirt (visible in specimen at top).
 841

T ruffles
TUBERALES
TRUFFLES are seldom seen because they grow underground.* Most of them look like
tiny potatoes or rocks, but reveal a system of canals, veins, and/ or cavities when sliced
open. They are likely to be confused only with the aptly named false truffles (Hymeno-
gastrales), which also grow underground but bear their spores on basidia rather than in
asci and typically have a minutely chambered rather than marbled, channelled, or hollow
interior. (For a more detailed comparison, see footnote at bottom of p. 844.)
The truffles are thought to be cup fungi which have gone underground. The evolutionary
pathway leading from a cup fungus to a truffle is exquisitely illustrated by the genus Geo-
pora. SomeGeoporus are truffle-like and subterranean, while others are cuplike and partly
exposed (i.e., hollow with a large mouth at ground level that opens to the air as shown on
p. 935). Some of the underground Geoporas are also hollow, but the mouth is oriented
randomly (at the side, bottom, etc.) and others have become greatly infolded so that the
interior is channel led or chambered (see photo on p.847) rather than hollow. The infolding
of the tissue is advantageous because it greatly increases the surface area for producing
spores (the spore-bearing asci line the canals or chambers inside the fruiting body). In
the “true” truffles (e.g., Tuber), this trend is carried to its logical extreme—the interior
of the fruiting body is marbled but solid (it presumably evolved through the fusing or
merging of folded tissue) and the asci are imbedded randomly in the tissue rather than
lining the canals or chambers.
As might be expected, truffles exhibit several other special adaptations to their under¬
ground lifestyle. In Geopora the spores are forcibly discharged as in the cup fungi, but
other truffles have lost the ability—and necessity—to discharge them because they do not
depend on the wind for spore dispersal. Instead, their spores are spread by various truffle¬
eating animals (rodents, deer, pigs, insects, slugs, etc.). The spores pass through the ani¬
mals’ digestive systems unscathed, and a microscopic analysis of the spore content of
animal droppings can give you a pretty good idea of which truffles and false truffles grow
in your area! Some animals, such as the California red-backed vole (a sort of burrowing
mouse) tunnel through the soil eating nothing but truffles and are thus restricted to the
coastal fog belt, where truffles and false truffles occur year-round.
To attract attention and make themselves desirable, most truffles have developed
distinctive odors and flavors. However, the odor and flavor do not normally become
strong until well after the truffle is mature, thereby insuring that the eater of the truffle will
ingest a large number of viable spores. Furthermore, truffles do not need to develop as
rapidly as epigeous (above-ground) fungi because they are insulated from sudden changes
in the weather. Instead, the maturation process takes place gradually, over a period of
several weeks or months, although a few spores often develop earlier coupled with a slight
scent, perhaps as a safeguard against a prolonged cold or hot spell that would inhibit
further development of the fruiting body.
Truffles, particularly species of Tuber, have been eaten for centuries. Unfortunately,
the fabled truffles of France and Italy have become a fetish of the rich. Due to their rarity
and the difficulty in finding them, they have acquired considerable snob appeal and retail
for more than $500 a pound! Their flavor and aroma are so powerful that a little goes a long
way, but to a person of modest means such as myself, nothing edible is worth that much!
Since truffles grow underground, we humans, with our underdeveloped noses, have

♦When speaking of truffles and false truffles, the terms “underground,” “subterranean,” “hypogeous,” and
“buried” are used rather loosely to mean beneath the surface of the ground, either in the humus layer or in the soil
itself or in the interface between the two.
Left: The famous “Black Diamond” of France, Tuber melanosporum, sliced open to show the
marbled interior (see pp. 854-855 for more details). Right: Microscopic view of the surface of a Tuber
spore, showing the alveolate (pitted-reticulate) pattern typical of many species. (Herb Saylor)

trouble finding them without “hired hounds.” Goats have been used to track down
truffles in Sardinia and bear cubs have been employed in Russia, but pigs and dogs are
the most accomplished truffle hunters. Some truffles contain pig sex hormones, meaning
that pigs have a natural nose (and lust!) for truffles. They require little or no training, but
must be physically restrained from devouring their quarry, and are hard to control even
when there are no truffles about. Acorns are sometimes given as nutritional recompense
for finding a truffle (a pitiful substitute, if you ask me) or the pig is muzzled and pulled
away just as it begins to dig up the truffle with its exceptional snout. Another problem with
pigs is that they tire easily, and must be carted to and from the truffle grounds if they are
distant.
Dogs, on the other hand, are tireless and devoted, and care more for humans than
truffles. In fact, most dogs loathe truffles and must be painstakingly trained to seek them
out. There are schools in Italy devoted exclusively to this purpose, and a seasoned truffle
hound commands a steep price. S hort-legged breeds are traditionally popular, presumably
because they’re closer to the ground. Both pigs and dogs, incidentally, can detect truffles
from as far away as 50 yards, and there is one case on record of a dog that jumped a hedge,
crossed a field, and “secured his prize” under a beech tree at least 100 yards away!
Perhaps you are now convinced that you need canine or porcine companions to find
truffles. Well, let me state, unequivocally, that you don’t. True, the odds against casually
bumping into a truffle are great, but you can find truffles by making a concerted effort
to find them. This means getting down on your hands and knees and systematically sifting
through the forest humus and soil, paying special attention to “truffle tracks”: squirrel
diggings, small cracks in the soil caused by those that develop close to the surface, strange
and compelling odors (some seasoned trufflers claim they can smell them out!), and an
occasional cloud of “truffle flies” hovering over the buried object of their affections.
Truffles, in fact, are far easier to find than most people realize. Looking for them is both
challenging and fun, like hunting for buried treasure or panning for gold, but without the
monetary incentive. (Entrepeneurs, are you listening? The expensive truffles of Europe
are not known to occur in the U nited States and most of our native species are notas richly
flavored.)
Why, then, do so few mushroom hunters look for truffles? Perhaps because they don’t
know how, when, or where. (As evidence of this assertion, I offer the chapter on truffles
in the first edition of Mushrooms Demystified!) The how, the when, and the where are
delightfully described by Harold E. Parks in the following excerpt from a 1921 article in
the scientific journal, Mycologia. A resident of San Jose, California, Parks was one of
California’s earliest and most avid trufflers, and he has had numerous species of truffles
and false truffles named after him.
842
TUBERALES 843

Even when one knows the ground thoroughly it is surprising how little of it may be
covered on a day of good collecting. Frequently two or three hours will be spent in
working over the ground under a single large oak, and on several occasions an entire
afternoon has been spent in one place . . .
The equipment of the truffle hunter is important. I use a wheel on many trips, as the
roads are excellent and the stops are very frequent in some places. It is easily hidden
in the brush when I leave the roadways and take to the high hills, and it makes accessible
places otherwise out of one’s reach. To the wheel is strapped a small combination rake
and hoe with a four-foot handle. This implement is very useful in climbing, raking and
digging and furnishes good protection in a snake country, as I well know. A short-
handled hoe useful for work in thick brush, a trowel, knife, tweezers, lens, kodak,
plenty of newspapers and a large number of small pasteboard cartridge boxes obtained
from a shooting gallery. These small boxes are very useful in handling the many small
specimens or single individual specimens, while large collections are wrapped in the
paper. Lunch and thermos bottle complete the outfit, and all are packed compactly in
the large canvas bags used by newsboys. These bags ride comfortably with a large load
evenly distributed over the shoulders.
In the earlier parts of the season the edges of the forests and the small groups of trees
are usually the best places for operations, although frequently the dense forest will yield
good specimens. Late in the season the best places are to be found deep in the forest,
where the ground retains more moisture. When the collector finds a favorable place
for operations the rake comes into use and a small area is raked free of leaves and humus.
W atch must be kept in the leaves for certain species... Other species will appear entirely
exposed on the surface of the earth [under the leaves] and some will be just beneath
the surface and out of sight. Excavation may be continued to a depth of a foot, at which
depth most species will cease to be found. Care should be taken at all stages, especially
near the surface, to avoid injury to specimens, but they will often be injured in spite of
it, and many of the dark-colored species will require very careful search and sifting of
the soil. The rewards are more often blistered hands and an aching back than truffles,
but there are also some intensely exciting moments . ..

To these remarks I would add only this: Digging up the forest can be unsightly as well
as destructive, so do it on a small scale, in scattered places over a large area, don’t go
truffling in locales traditionally frequented by mushroom hunters (they have a right to
undisturbed duff!), and always cover up the soil you expose, leaving the environment as
close to its original state as possible.
Truffles are mycorrhizal. This means they can be found wherever there are trees and
shrubs, but like the false truffles, they are especially abundant and diverse along the west
coast. They are normally terrestrial, but can also occur inside very rotten wood that has
been permeated by tree rootlets. This is particularly true in dry areas like the Sierra Nevada,
where the rotten logs are a major source of moisture for both the trees and the truffles.
In our area they seem to favor evergreen oaks (live oak and tanoak) and conifers such as
Douglas-fir. Because truffles develop slowly, they are usually found at the end of the
mushroom season (February-July in our area). Some, such as Tuber gibbosum, are
excellent esculents; others are mediocre and still others have yet to be tried.
Although a microscope is often required, truffles are not as difficult to identify as false
truffles (for one thing, there are far fewer species). A fairly extensive—but by no means
comprehensive—selection is offered here in the hope that it will stimulate mushroom
hunters to start looking for these clandestine denizens of our forests. Since I am by no
means an expert on truffles, I have gleaned much of the information in this chapter from
articles by Helen Gilkey and James Trappe, the past and presentauthorities on the subject.
Anyone who takes truffles seriously should consult these articles(see Suggested Readings
and References) or join the North American Truffling Society.
Modern taxonomists try to show the truffles’ relationships to the cup fungi by scattering
them among several families in the Pezizales, just as they place many false truffles with
the Agaricales. The Tuberales, in other words, is a defunct and artificial—but convenient
844 TUBERALES

—category that is used here because it facilitates identification. In the following key,
the truffles have been divided into several natural groups. An attempt has been made to
use field characters, but microscopic features have unavoidably come into play—
particularly the shape and ornamentation of the mature spores (you must have at least
one mature or partially mature specimen!). Truffle spores, incidentally, are exceptionally
ornate. Some are spiny like porcupines or pitted like golf balls, others are covered with a
geometrical network of ridges (see photo on p. 842), still others are warted, pegged,
or smooth.

Key to the Tuberales

1. Spores borne on basidia* .(see Hymenogastrales & Allies, p. 739)
1. Spores borne inside asci* . 2
2. Found in the deserts of the Southwest; fruiting body more or less round and somewhat puffball¬
like, i.e., developing underground but sometimes emerging at maturity, then drying out and
blowing about in the wind; asci brown under the microscope .Carbomyces
2. Not as above . 3
3. Fruiting body earthball-like, i.e., consisting of a thick (2-5 mm) tough outer wall and a single
large inner cavity which is soon filled with tissue; internal tissue at first white and cottony,
becoming divided into several chambers by white sterile bands, then becoming dark brown to
blackish and powdery when mature (the asci disintegrating quickly and the sterile bands not
evident in age) .Elaphomyces, p. 862
3. Not as above; mature spore mass not powdery . 4
4. Spores borne inside the fruiting body (or on inside surfaces); common . 5
4. Spores borne externally (on outside surface of fruiting body); rare. 23
5. Interior of fruiting body either hollow or with empty chambers or with open veins or empty
canals formed by infolding of the fruiting body wall, or occasionally with chambers that are
loosely stuffed with cottony hyphae . 6
5. Interior solid, with pockets or zones of fertile tissue and meandering sterile veins . 16
6. Fruiting body hollow inside (with one or sometimes two large empty chambers which may be
round or convoluted or canal-like due to infolding of the outer wall) . 7
6. Interior of fruiting body with separate canals or several separate chambers . 9
7. Exterior of fruiting body with small rounded to angular warts, typically with one or more open¬
ings to the interior .Genea& Genabea, p. 849
7. Exterior not finely warted; opening(s) present or absent . 8

*As already pointed out, this fundamental difference can only be seen with a microscope (and then only after the
basidia or asci have formed and before they disintegrate). However, the false truffles (underground Basidiomycetes)
and truffles (underground Ascomycetes) can often be differentiated in the field by the following characters:

If the interior is gelatinous, it is a false truffle.

If the interior has a columella (e.g., a branched or unbranched internal stalk or well-developed sterile base),
it is probably a false truffle (exception: Fischerula subcaulis, an Ascomycete).
If the interior is solid and marbled with veins, it is probably a truffle.
If the interior is composed of numerous minute holes or empty chambers (giving it a sponge-like ap¬
pearance) it is probably a false truffle.
If the interior shows the embryonic beginning of cap, gills, and stalk, it might be a young Amanita or other
gilled mushroom!
If the fruiting body is very hard with a solid interior that flakes or chips off like wax, it is probably a truffle.
If the interior is completely hollow or has several large hollows or is composed of one mazelike hollow,
it is probably a truffle.
If the wall (peridium) of the fruiting body is very thick (several mm) and the interior is not hollow and the
fruiting body has a distinct base, it is probably a false truffle or earthball.
If the outer wall of the fruiting body is very thick and the interior is cottony or powdery and the fruiting body
lacks an obvious base, it is probably a truffle.
If the wall of the fruiting body is infolded to form numerous empty canals or veins orcavitiesthat often open
to the exterior, it is probably a truffle.
If the exterior is covered with rootlike mycelial threads (rhizomorphs) it is probably a false truffle.
If the exterior is covered with warts (often small or large), it is probably a truffle.

There are also several mycorrhizal Zygomycetes (e.g., Endogone and Glomus) with truffle-like fruiting bodies.
These fungi are not treated in this book because they have neither asci nor basidia. Instead sexual spores are
formed by the conjugation of “mother cells” (gametangia), and asexual spores are often formed on hyphae.
Most species have gigantic spores (100-200 microns!) and some, such as the common Endogone lactiflua, exude
a latex when cut.
TUBERALES 845

8. Fruiting body a large hollow ball (but often lobed or flattened); inside surface white to grayish,
pinkish, or purplish, not warted; outer wall often splitting into lobes (at top) in age; common,
especially under northern and mountain conifers .(see Sarcosphaera, p. 825)
8. Not as above . 11
9. Exterior of fruiting body with rounded to angular warts . 10
9. Exterior of fruiting body smooth or hairy, but not distinctly warted . 11
10. Fruiting body black to brownish to reddish or orange, often (but not always) with a tuft of my¬
celium at the base; spores smooth even at maturity.Balsamia & Allies, p. 852
10. Fruiting body white to yellowish or yellow-gray, or if not then interior warted like exterior; basal
tuft present or absent; spores ornamented at maturity .Genea& Genabea, p. 849
11. Exterior of fruiting body with fuzzy brown hairs (i.e., tomentose); interior a hollow chamber
or with open canals formed by complex infolding of the fruiting body wall; spores forcibly
discharged, smooth at maturity; fairly common, especially under conifers . Geopora, p. 846
11. Not as above; exterior not brown and tomentose . 12
12. Spores smooth and round at maturity; chambers of the fruiting body typically stuffed with
cottony hyphae; tips of asci not amyloid; rare (at least on west coast) .Stephensia
12. Not with above combination of features. 13
13. Asci with amyloid tips (i.e., tips staining bluish in iodine solution) (see Peziza & Allies, p. 818)
13. Asci not amyloid . 14
14. Interior of fruiting body hollow or with open veins, canals, or chambers formed by complex
infolding of the fruiting body wall; spores ornamented at maturity; especially common under
northern or mountain conifers (often inside rotten wood), but also found in other habitats
.Hydnotrya, p. 848
14. Not as above . 15
15. Spores elliptical and smooth at maturity; channels or canals inside fruiting body usually empty
.Balsamia & Allies, p. 852
15. Not as above; spores ornamented (at least at maturity) .Tuber & Allies, p. 854
16. Columella (sterile column or base or rudimentary internal stalk) present, or if not then a very
distinct basal pad of mycelium present; exterior of fruiting body pallid to pinkish-gray to
brownish; interior pinkish-gray to grayish-purple to nearly black with narrow white veins;
spores very large (60-100 microns), elliptical, brown at maturity and ornamented with obscure
spines; found under conifers in the Pacific Northwest .Fischerula subcaulis
16. Not as above; columella absent .•. 17
17. Asci with amyloid tips or weakly amyloid throughout; asci arranged in a distinct palisade
(hymenium); spores elliptical . 18
17. Asci not amyloid; asci arranged in a palisade or randomly imbedded in tissue; common . . 21
18. Spores round, ornamented with spines or pegs at maturity; rare.Tuber & Allies, p. 854
18. Not as above; occasional . 19
19. Asci weakly amyloid throughout; fruiting body white to yellowish or yellow-brown; rare
(known from coastal California) . 20
19. Asci amyloid mainly or only at their tips; fruiting body often darker than above; widespread
.(see Peziza & Allies, p. 818)
20. Spores less than 20 microns long .Hydnotryopsis setchellii
20. Spores averaging 20 microns or more long.Hydnotryopsis compactus
21. Fruiting body with a fatty or gristle-like consistency (especially the interior), usually whitish
or buff when fresh; spores round and ornamented at maturity .Tuber & Allies, p. 854
21. Not as above; texture not gristle-like . 22
22. Spores smooth even when mature; exterior of fruiting body often warted; interior pallid when
mature or grayish to olive with pallid veins (rarely brown) .Balsamia & Allies, p. 852
22. Spores ornamented at maturity; exterior of fruiting body warted in some species, but more often
smooth; interior pallid when immature but usually brown or reddish-brown with pallid veins
when mature (but sometimes grayish or olive) .Tuber & Allies, p. 854
23. Fruiting body pale brown to brown or purplish; asci faintly amyloid; spores mostly 20 microns
or more broad, hyaline (colorless) under the microscope . Sphaerosoma
23. Fruiting body yellowish to olive or brown; asci not amyloid; spores 8-25 microns broad, hyaline
to yellowish or brown under the microscope .Sphaerozone
846 TUBERALES

GEOPORA (Fuzzy Truffles)

Small to medium-sized fungi growing underground or at ground level. FRUITING BODY usually
more or less round to cuplike, with one (or sometimes more) openings to the interior. Exterior
typically brown and hairy or fuzzy (tomentose). INTERIOR complexly folded or channelled in
one species, hollow in the others. STALK, columella, and basal mycelial tuft typically absent.
SPORES elliptical to nearly round, smooth, hyaline (colorless) under the microscope, forcibly
discharged. Asci arranged in a distinct palisade(hymenium) lining the inside surface of the fruiting
body or the internal folds (if present), mostly 8-spored, not amyloid.

THIS genus is best recognized by its fuzzy brown exterior. Some species, such as G. areni¬
cola, are traditionally grouped with the cup fungi (in genus Sepultaria) because of their
hollow fruiting body that grows just below the soil surface with only the large opening or
“mouth” at the top exposed to the air. G. cooperi, on the other hand, has traditionally
been treated with the truffles because it grows underground and has a complexly folded
interior and one or more irregularly oriented openings. These apparently disparate fungi
are linked by species such as G. clausa, which is hollow like G. arenicola, but grows under¬
ground and has an apical, basal, or lateral opening.
Geopora is currently placed in the Pyronemataceae, alongside Aleuria, Otidea, Geo¬
pyxis, and many other cup fungi. About a dozen species are known, most of them in the
mode of G. arenicola. However, G. cooperi seems to be the most common species in Cali¬
fornia. It is said to be a good edible, but I can find no information on other members of
the genus.
Key to Geopora
1. Wall of fruiting body complexly infolded to create numerous canals or chambers inside the
fruiting body . 2
1. Not as above; interior of fruiting body with a simple hollow . 3
2. Exterior of fruiting body only slightly hairy if at all; odor often sweet or garlicky when fully
mature; spores ornamented at maturity; often (but not always) growing inside rotten wood
. (see Hydnotrya, p. 848)
2. Not as above; exterior distinctly hairy or fuzzy; spores smooth .G. cooperi, below
3. Opening or“mouth” of fruiting body irregularly oriented(at top, base, or side); usually growing
underground .G. clausa (see G. arenicola, p. 847)
3. Opening or “mouth” always at top; fruiting body immersed or partly immersed in the ground
with the mouth exposed to the air .4
4. Fertile surface (interior) orangish to reddish or sometimes yellowish .
.G. aurantia& G. pellita(see G. arenicola, p. 847)
4. Fertile surface white to pale brownish or sometimes drying yellowish . 5
5. Fruiting body up to 1 cm broad .G. arenosa(see G. arenicola, p. 847)
5. Fruiting body l A cm broad .G. arenicola & others, p. 847

Geopora cooperi (Fuzzy Truffle)

FRUITING BODY usually buried or partially buried, round or nearly round (but often
squirrel-eaten), 2-7 (10) cm broad. Exterior fuzzy or velvety fromacoating oflight brown
to dark brown hairs, usually furrowed. INTERIOR white to creamy or yellowish-tan,
usually streaked with tan or brown, deeply convoluted, the folds often touching each
other but leaving at least some open spaces or“canals” between them. Odor usually mild,
but in one form resembling fermented cider. SPORES 18-27 (30) * (10) 13-21 microns,
broadly elliptical in one form, round or nearly round in another; hyaline (colorless) under
the microscope, smooth, with one oil droplet. Asci usually 8-spored, forming a distinct
palisade (hymenium) that lines the open surfaces of the folds.
HABITAT: Solitary, scattered, or gregarious on or in the ground under both hardwoods
Geopora cooperi. Note the convoluted interior (specimen on left) and fuzzy exterior (specimen on
right). It is usually found under conifers, often in sandy soil.

and conifers (but especially the latter); widely distributed in western North America and
locally common, especially under mountain conifers during the spring, summer, and fall.
It favors pine in coastal California and pine, fir, or spruce in the Sierra Nevada and else¬
where; in Alaska it has been found under willow and aspen. It develops underground
but may surface (or be dug up by squirrels) in age, and so is often seen by casual collectors.
EDIBILITY: Edible. Rodents are very fond of it and so are some humans.
COMMENTS: This is one of our largest truffles and also one of the more distinctive.
Its telltale traits are the fuzzy brown exterior and convoluted interior. The latter is simply
a mass of folded tissue (see photograph), with the spore-bearing asci lining the empty
spaces or“canals” between the folds. The spores are shot out of the asci as in the cup fungi.
It has numerous synonyms, including G. harknessii and G. magnata.

Geopora arenicola (Hole In The Ground)

FRUITING BODY at first closed and buried in ground, then opening at the top and
becoming more or less cup-shaped at maturity, the margin remaining incurved or often
splitting stellately (in starlike lobes) in age; 1-4 cm broad. Exterior brown and densely
clothed with flexible brown hairs that bind surrounding dirt or sand. INTERIOR (fertile
surface) pallid to creamy or grayish, often becoming yellowish, tan, or brownish in age;
smooth. Flesh brittle to rather tough. STALK absent or rudimentary. SPORES 23-30 x
12-17 microns, elliptical to spindle-shaped, smooth, usually with one( rarely two) large oil
droplets. Asci lining inner surface of cup, typically 8-spored, not amyloid.
HABIT AT: Scattered to densely gregarious or clustered in sand or silt, disturbed ground,
etc., usually immersed in the soil with only the mouth showing; widespread but not
common, or at least not often noticed. I have found it once in our area, in the spring.
EDIBILITY: Academic—it is practically impossible to get rid of the sand or dirt!
COMMENTS: Better known asSepultaria arenicola, this species looks like a hole in the
ground or an insect burrow or worm tunnel( see photo on p. 935). In old age it is clearly cup¬
like, however, and often splits into lobes (as shown in photo). The hairy brown exterior
distinguishes it from Sarcosphaera crassa and other cup fungi that develop in the ground,
and relates it to G. cooperi, which has a complexly folded rather than hollow interior. The
fuzzy brown hairs can be seen with a hand lens by gently brushing away some of the dirt.
Other species: G. longii of the South west is similar but has nearly round spores; G. arenosa
is also similar but much smaller; G. aurantia is similar but has a reddish to orange or
egg-yellow interior and rigid hairs on its exterior; G. pellita has a yellowish to pale orange
interior, but its hairs are not rigid; G. clausa(-Hydnocystis californica) has a fuzzy brown
exterior and its mouth is oriented randomly with respect to the fruiting body (i.e., at the top,
bottom, or side). It usually grows underground, but is rare in our area.

847
848 TUBERALES

HYDNOTRYA (Wood Truffles)

Small to medium-sized woodland fungi found in soil or very rotten wood. FRUITING BODY
roundish to lobed or brainlike. Exterior variously colored, smooth to scurfy or occasionally slightly
fuzzy, but not conspicuously hairy or warted. INTERIOR variable in structure, sometimes com¬
posed of a single chamber, but more often with several to many chambers or canals formed by
complex infolding and fusing of the outer wall. STALK and columella absent; mycelial tuft also
absent. SPORES round to elliptical or cubical, smooth at first but becoming roughened, warted,
ridged, pitted, or spiny at maturity; yellowish to brown under the microscope. Asci typically borne
in a palisade (hymenium) lining the chambers) or canals, not amyloid.

THE fruiting bodies of this genus are extremely variable in size and shape. Some species
are hollow inside, others are complexly folded. All have open canals or chambers, non¬
amyloid asci, and spores which suggest a kinship to the elfin saddles and false morels
(Helvellaceae). Geopora exhibits a similar range of variation, but has a hairier or fuzzier
exterior and smooth spores. Genea and Genabea differ in having a distinctly warted
exterior and interior.
Hydnotryas are sometimes called “wood truffles” because several species can fre¬
quently be found inside rotten wood (they also grow in soil). As pointed out earlier, the
fact that they grow inside wood does not necessarily mean they are wood-rotters. Rather,
they could be associated with tree rootlets that penetrate the wood in search of moisture.
A dozen species of Hydnotrya are known; half occur in North America. They are fairly
common in the Sierra Nevada and other mountain ranges, but rare or absent in coastal
California. A single species is described here and several others are discussed.

Key to Hydnotrya
1. Exterior of fruiting body brown to dark brown to dark reddish-brown; interior complexly
folded or convoluted; odor often strong (sweet or garlicky) when fully mature', spores round
(but often knobby) . H. cerebriformis& H. tulasnei(see H. variiformis, below)
1. Exterior whitish to buff, cinnamon-buff, pinkish-cinnamon, or sometimes brown; interior
complexly folded or a simple hollow; odor not usually as above; spores elliptical. 2
2. Interior usually a simple hollow; spores often cubical; usually found in ground .
.H. cubispora (see H. variiformis, below)
2. Interior ranging from a simple hollow to complexly folded; spores not cubical; found in ground
or inside rotten wood . 3
3. Asci with amyloid tips; common in coastal California .(see Peziza & Allies, p. 818)
3. Not as above; asci not amyloid .H. variiformis & others, below

Hydnotrya variiformis
FRUITING BODY 0.7-4 cm broad, round to somewhat flattened, depressed, or lobed.
Exterior minutely velvety, whitish to creamy to buff or yellowish to cinnamon-buff or
brownish, not warted. INTERIOR variable in configuration, but small specimens often
containing a simple cavity with a prominent opening, and larger ones usually with several
chambers or narrow, branching canals formed by crowding and infolding of the outer
wall; canals usually empty but their sides often fused; white or pallid, but the hymenium
(fertile tissue) often brownish to pinkish-orange at maturity. SPORES 32-36 * 24-28
microns, elliptical, smooth becoming minutely pitted and wrinkled at maturity, yellowish-
brown under the microscope. Asci borne in a palisade (hymenium) that lines the canals or
cavity, typically 8-spored.

HABITAT: Solitary or in groups in soil or inside very rotten wood under conifers; oc¬
casional (along with H. cerebriformis—see comments) in the Sierra Nevada, Cascades,
and other western mountains. It fruits in the spring, summer, and early fall.
EDIBILITY: Edible? I can find no specific information on it.
HYDNOTRYA 849

COMMENTS: This species and H. cerebriformis (see below) can usually be told in the
field by their fondness for growing inside rotten wood plus their complexly folded interior
(at least in large specimens) and non-amyloid asci. The exterior lacks the warts of Genabea
cerebriformis and the brown hairs of Geopora cooperi. H. cerebriformis is similar to
H. variiformis and grows in similar habitats. It has a more consistently complex or con¬
voluted interior (not unlike that of Geopora cooperi, shown on p. 847), is usually slightly
darker than H. variiformis (dull reddish-brown to dark purple-brown), typically has a
strong garlicky odor when mature, and has round, minutely spiny spores. H. tulasnei is a
widespread odoriferous species that is very similar to H. cerebriformis; it is also brown to
reddish-brown, but has coarsely warted spores. H. cubispora is a widely distributed,
brownish to pinkish-cinnamon, usually terrestrial species with a more or less hollow (but
lobed) interior and spores which are often cubical. H. michaelis (-H. yukonensis) is a
rather rare northern species with elliptical warted spores and a convoluted interior.

GENEA& GENABEA (Geode Truffles)

Small underground woodland fungi. FRUITING BODY round to strikingly lobed or brainlike,
usually with one or more openings to the interior. Exterior minutely warted, variously colored,
bald or with hairs. INTERIOR usually warted; hollow or with empty, mazelike canals formed by
infolding or inward projections of the outer wall. STALK and columella absent, but basal tuft of
mycelium often present in Genea. SPORES round to elliptical, warted or spiny at maturity (but
ornamentation may dissolve in KOH or Melzer’s reagent!), pale or colorless (Genea) to brown
(Genabea) under the microscope. Asci arranged in a palisade (hymenium) lining the surfaces of
the inner cavity or canals; typically 8-spored, not amyloid.

THESE small truffles are easily recognized by their finely warted, often lobed fruiting
bodies with a hollow or partly hollow, warted, geode-like interior. Geneas typically have
a single, often irregularly shaped cavity and warted spores, while Genabeas usually
have a more complex or mazelike interior and spiny spores. In addition, the western
species differ in color: reddish to brown or black in Genea, white or creamy in Genabea.
The origins and affinities of Genea and Genabea are unknown; together they form the
family Geneaceae. Both are fairly common, at least in California. They seem to fruit closer
to the surface of the ground than many truffles and also have an earlier season, appearing
in late November in our area and continuing on into the spring. I can find no information
on their edibility. Their small size is hardly an asset, but they seem to be very popular with
our local wild pigs. (You can sometimes find them where pigs have been foraging.) Genea
has over 20 known species, whereas Genabea includes only a handful. Three Geneas and
one Genabea are described here and several others are keyed out.

Key to Genea & Genabea

1. Exterior of fruiting body or at least the “mouth” with brown hairs . 2
1. Not as above . 3
2. “Mouth” of fruiting body fringed with stiff hairs Genea kraspedestoma{see G. arenaria, p. 851)
2. Not as above . Genea arenaria & others, p. 851
3. Exterior distinctly reddish; interior often reddish- or pinkish-tinged Genea intermedia, p. 851
3. Not as above (but exterior may be very dark reddish-brown) . 4
4. Exterior of fruiting body dark reddish-brown to dark brown to almost black when fresh .. 5
4. Exterior white to buff, yellowish, light brown, or medium brown when fresh . 6
5. Found in West; spores warted .Genea harknessii & others, p. 850
5. Found in eastern North America; spores spiny Genabeafragilis(see G. cerebriformis, p. 851)
6. Exterior of fruiting body white to creamy or pale yellow; interior usually complex (mazelike or
with many separate chambers); spores spiny .Genabea cerebriformis & others, p. 851
6. Exterior light yellow-brown to light brown or brown; interior usually a single cavity (but the
cavity often convoluted); spores warted . Genea compacta (see G.harknessii, p. 850)
Many geode truffles look like bits of knobby coral. This one is Genea harknessii. (Herb Saylor)

Genea harknessii (Dark Geode Truffle)

FRUITING BODY underground, 0.5-2.5 cm broad, round to flattened to very knobby
and irregularly lobed, usually with an apical opening to the interior and a tuft of mycelium
at the base. Exterior dark reddish-brown to dark brown to dark gray or black (but often
with a thin whitish covering of hyphae when very young), divided into small, often pyra¬
midal warts. INTERIOR basically hollow, but often interrupted by irregular projections
of sterile tissue from the outer wall; inner surface of wall warted and dark brown to blackish
or bluish-gray; sterile tissue white to grayish. SPORES averaging 24-28 x 22-27 microns,
elliptical to nearly round, hyaline (colorless) under the microscope, smooth at first but
finely warted at maturity. Asci 8-spored, arranged in a palisade (hymenium).
HABITAT: Solitary to gregarious in humus or soil under oak, manzanita, coyote bush
(Baccharis) and other trees and shrubs; known only from the west coast. It is fairly com¬
mon (for a truffle) in California in the winter and spring, especially in February-March.
I’ve found this species, G. gardneri, and G. compacta (see comments) under live oak. It
also occurs in the Sierra Nevada. Like other Geneas, it often grows just below the ground
or on the soil surface beneath the humus, and is difficult to see because of its color.
EDIBILITY: Prized by pigs, but I can find no mention of humans eating it.
COMMENTS: The hollow interior and dark warted exterior separate this common
species from most other truffles. The shape ranges from nearly round to elaborately lobed
or cerebriform(brainlike). Round specimens are reminiscent of miniature geodes because
of their warted internal cavity, while the more knobby specimens look like bits of coral or
piles of intertwined worms (see above photo). Other species: G. gardneri is very similar
but has larger, coarsely warted spores; it grows under oak in coastal California. G. com¬
pacta is a similar but much paler (brown to yellowish-brown), knobby species with a
hollow, convoluted interior (see photo below). I have found it several times under oak
in the spring. It is edible but tasteless.

Left: Genea gardneri (see comments above). Note black hollow interior (below) and warted exterior.
(Herb Saylor) Right: Genea compacta (see comments above), a common yellowish to brown, oak-
loving species. Note knobby fruiting body, large “mouth” (specimen at center), and hollow interior.
GENEA 851

Genea intermedia (Red Geode Truffle)

FRUITING BODY underground, 0.5-2.5 cm broad, sometimes roundish(especially when
young), but more often lobed or knobby, with or without an apical opening to the interior
and a basal mycelial tuft. Exterior reddish to reddish-brown, vinaceous, or vinaceous-
purple, the lobes or protuberances superimposed with minute warts. INTERIOR
basically hollow, but often interrupted by irregular projections of sterile tissue from the
outer wall; inner surface warted, pink to whitish; sterile tissue also white to pinkish.
SPORES averaging 36-40 microns, round, finely warted at maturity. Asci arranged in a
palisade (hymenium), typically 8-spored.
HABITAT: Solitary to gregarious in soil in woods; known only from Oregon and Cali¬
fornia. It is not uncommon under conifers in the Sierra Nevada in the spring; I have yet
to find it on the coast.
EDIBILITY: Unknown.
COMMENTS: The beautiful reddish, lobed and warted exterior plus the hollow interior
make this one of the few truffles that can be recognized instantaneously in the field. No
other Genea is as red, at least in California. (G. harknessii can be dark reddish-brown
when fresh but soon blackens after picking.)

Genea arenaria (Hairy Geode Truffle)

FRUITING BODY underground, 1 -3 cm broad, usually irregularly lobed or coarsely
knobby (at least at maturity), with or without an opening to the interior and a basal tuft of
mycelium. Exterior brown to pale brown, divided into small, often pyramidal warts and
covered with scattered long brown to dark brown hairs. INTERIOR basically hollow, but
the cavity usually irregular in shape due to infolding of the outer wall; inner surface finely
warted and colored like the exterior or paler; sterile tissue whitish. SPORES averaging
22-32 x (16) 20-24 microns, mostly elliptical, with minute scattered warts. Asci arranged
in a palisade (hymenium), typically 8-spored.
HABITAT: Solitary, scattered, or in small groups in soil under trees (mainly live oak);
known only from California and Oregon. It typically fruits from the late fall through the
early spring (November-April) and is not uncommon in our area. However, it seldom
occurs in quantity and is easily overlooked because of its brown color.
EDIBILITY: Unknown.
COMMENTS: This species is best recognized by the highly irregular shape at maturity,
the hollow (but usually folded) interior, and the presence of hairs on the warted brownish
exterior. Other species: G. hispidula is a similar species known from eastern North
America; G. kraspedestoma is a reddish-brown to brownish Californian with a circular
apical opening that is fringed by stiff incurved hairs; it has small spores( only 12-20 microns
long) and was originally collected near Almaden, California, under oak. G. compacta (see
comments under G. harknessii) is also similar, but lacks obvious hairs on its exterior.

Genabea cerebriformis (White Geode Truffle)

FRUITING BODY underground, 0.4-1.5 (2.5) cm broad (but usually under 1 cm); smaller
specimens often roundish, larger ones usually quite irregular (knobby, lobed, or brain¬
like) in shape and typically with several openings to the interior; lacking a basal tuft of
mycelium. Exterior covered by small, more or less conical warts superimposed on each
knob; white to yellowish-white or yellow-gray. INTERIOR often a single cavity in small
specimens, but in larger ones usually consisting of a mazelike system of canals formed by
852 TUBERALES

infolding and inward projections of the wall; colored more or less like exterior. Odor
mild or strong. SPORES 28-44 microns, round, smooth at first but covered with long,
slender spines at maturity; hyaline (colorless) to grayish-yellow (in age) under the micro¬
scope. Asci arranged in a palisade (hymenium), typically 8-spored.
HABITAT: Solitary to gregarious in soil or humus in woods and under trees; fairly
common (for a truffle) in western North America under various trees, but especially fond
of Douglas-fir. In California it fruits, like other truffles, in the winter, spring, and early
summer. Although small, its light color makes it fairly conspicuous.
EDIBILITY: I can find no information on it; too small to be of much value.
COMMENTS: Formerly known as Genea (or Myrmecocystis) cerebriformis, this is one
of our most distinctive truffles. The yellowish-gray to white color plus the irregularly con¬
voluted and warted exterior and complex interior with open canals (in larger specimens)
distinguish it. The interior is vaguely reminiscent of Geopora cooperi, but that species is
much larger and has a fuzzy brown exterior. Other species: Genabea fragilis (the type
species of the genus) is a blackish species reported from Europe and Quebec; G. spino-
spora is a whitish species that has been found in Virginia.

BALSAMIA & Allies (Smooth-Spored Truffles)

Small to medium-sized underground woodland fungi. FRUITING BODY round to somewhat
flattened or irregular, often with a depression or cavity leading to the interior. Exterior variously
colored, warted in Balsamia, warted or smooth in Barssia and Picoa. INTERIOR solid with sterile
veins and pockets of fertile tissue (Picoa), or with open or hyphae-stuffed veins or canals (Barssia
& Picoa); variously colored. ST ALK and columella absent, but a basal tuft of mycelium sometimes
present in Balsamia. SPORES smooth at maturity, usually elliptical or elongated but sometimes
round; colorless under the microscope in Barssia and Balsamia, sometimes brown in Picoa. Asci
typically 8-spored, not amyloid, arranged in a palisade or randomly imbedded in the tissue.

THREE genera are treated here: Balsamia, Barssia, and Picoa. They are intermediate in
aspect between the geode truffles (Genea and Genabea) and the true truffles (Tuber and
allies). As a unit they are difficult to distinguish in the field because their unifying feature
is microscopic: the spores are smooth even at maturity. On an individual basis, however,
the three genera are easier to recognize. For instance, Balsamia can be told by the frequent
presence of a basal mycelial tuft plus its warted exterior and pale (white to yellowish)
marbled interior. Barssia typically has a broad depression or “mouth” at the top of the
fruiting body and several open canals which empty into it. Picoa, on the other hand, has
a solid interior plus a brown to blackish exterior. It is easily confused with the true truffles,
but has a greener or grayer interior and smooth spores.
Balsamia, Barssia, and Picoa constitute the family Balsamiaceae, but their relationships
to other families are unclear. They occur in a variety of habitats and seem to fruit relatively
early for truffles, at least in our area. All three genera are small; one species from each is
described here.
Key to Balsamia & Allies
1. Interior of fruiting body solid and usually gray to greenish-gray to greenish-blue at maturity;
exterior slate-violet to black, minutely warted; basal mycelial tuft absent .
.Picoa carthusiana, p. 854
1. Not as above; interior of fruiting body not solid, or if solid then remaining pallid .2
2. Interior of fruiting body with open canals which empty into a broad central depression at or near
the top of fruiting body; exterior warted or not warted; basal mycelial tuft absent .
. Barssia oregonensis, p. 853
2. Not as above; internal veins or canals empty or stuffed with hyphae; exterior finely warted;
tuft of mycelium often present at base . Balsamia magnata, p. 853
Balsamia magnata. Note marbled or veined interior and finely warted exterior. Unlike Tuber, the
interior does not darken appreciably at maturity. (Herb Saylor)

Balsamia magnata
FRUITING BODY underground, 0.5-2 cm broad, round to somewhat compressed or
flattened, the apex usually infolded and the base often with a tuft of mycelium. Exterior
divided into numerous rounded to pointed warts, occasionally with small depressions;
color variable: bright orange to reddish-brown to brownish-pink or occasionally black
(but may be whitish when very young). INTERIOR white to pale yellowish, even when
mature; composed of crowded folds which form mazelike canals, the canals united or
separated into several chambers and either open or filled loosely with cottony hyphae;
canals usually converging at the apex or sometimes at several points. SPORES 20-24 *
12-14 microns, variable in shape (cylindrical to elliptical to nearly round), smooth at
maturity, hyaline (colorless) under the microscope, usually with three oil droplets. Asci
mostly imbedded in the tissue between the veins or canals; typically 8-spored.
HABITAT: Solitary to gregarious (usually the latter) in soil under various trees and
shrubs (oak, pine, madrone, etc.); common (for a truffle) in California and Oregon in the
winter and spring; also reported from Arizona.
EDIBILITY: I can find no information on it.
COMMENTS: The prominently warted orange to reddish-brown exterior plus the pallid
interior composed of open or stuffed, often united or converging canals are characteristic
of this rather common truffle. It is most likely to be confused in the field with Pachyphloeus
citrinus (which can also be bright orange), but the smooth spores and pale interior dis¬
tinguish it. Pseudobalsamia magnata is an older alias, and the names P. alba and P.
nigrens have been used for the whitish and black forms (species?), respectively.

Barssia oregonensis (Depressed Truffle)

FRUITING BODY 1-2.5 cm broad, roundish but usually more or less flattened or slightly
lobed, typically with a prominent depression or cavity at the top; firm but not hard, without
a mycelial tuft at base. Exterior smooth to roughened or finely warted, pale ochre-buff to
orange-cinnamon to brick-red or reddish-brown. INTERIOR composed of empty, un¬
connected, broad or narrow canals, many of which empty into the depression (i.e., more
or less solid except for the canals); white to pale gray, even at maturity. SPORES 24-36
x 12-21 microns, oblong-elliptical, smooth, colorless under the microscope. Asci mostly
8-spored, forming a palisade (hymenium) that lines the canals.
HABITAT: Solitary or in small groups in soil in woods, associated with Douglas-fir and
possibly other trees; known only from the West (California, Oregon, Idaho). Like most
truffles, it is commonest in the spring and early summer. I have yet to find it in our area,
but see comments.
EDIBILITY: Presumably edible; I haven’t tried it.

853
854 TUBERALES

COMMENTS: This truffle is best recognized by its color, the frequent presence of a
prominent depression into which several canals empty, and the white or pallid interior. It
might be mistaken for a Genea or Genabea, but is differently colored, not as warted, and
has smooth spores at maturity. An unidentified yellowish to pale orange Barssia with a
prominent broad apical depression has been found recently in the Guadelupe Mines area
near Almaden, California (an area as richly endowed with truffles as it is with mercury).

Picoa carthusiana (Oregon Black Truffle)

FRUITING BODY 0.5—4.5 (8) cm broad, round or nearly round to slightly irregular,
without a mycelial tuft at the base. Exterior minutely warted, black to dusky slate-violet.
INTERIOR more or less solid, composed of large pockets of fertile tissue marbled with
paler (whitish to buff) sterile veins; fertile tissue whitish to buff when young but becoming
grayish-green to greenish-blue in age; sometimes exuding a clear latex when fresh which
slowly (overnight) stains white paper pale violet. SPORES (56) 74-84 * 20-35 microns,
lemon- or spindle-shaped, smooth, typically with one giant oil droplet at maturity; pallid
to greenish-yellow becoming brown at maturity (under the microscope). Asci typically
8-spored, imbedded in the tissue (not forming a palisade).
HABITAT: Solitary, scattered, or in small groups in soil and humus in woods; known
from Europe and the western United States. In Oregon and California it favors Douglas-
fir. In our area it fruits in the spring and summer, but is more common in Oregon, where it
occurs earlier. It has been found at Point Reyes by Herb Saylor and Dennis Desjardin,
in July.
EDIBILITY: Edible and delicious raw, according to truffle connoisseur Gary Menser.
COMMENTS: In the words of Herb Saylor (Mycena News, May 1983), “The general
aspect of this fungus is that of a piece of animal dung, with which we frequently confused
it while making the collection. It is possible that this may be one reason why it is infrequent¬
ly collected, as animal dung of similar size and color was common in the area.” The solid
interior might lead to confusion with Tuber and allies, but its greenish to grayish color
at maturity plus the large, smooth, spindle-shaped spores and minutely warted, blackish
exterior form a distinctive set of characters.

TUBER & Allies (True Truffles)

Small to fairly large, underground mycorrhizal fungi. FRUITING BODY round to copiously
lobed, often hard (especially in Tuber). Exterior sometimes warted in Tuber and Pachvphloeus,
otherwise not; variously colored. INTERIOR typically solid and firm, usually marbled, often
waxy, usually white or pallid when young but usually with dark fertile tissue at maturity or pockets
of fertile tissue separated by paler walls. STALK and columella absent; basal mycelial tuft
also absent (except in some species of Pachvphloeus). SPORES ornamented with spines, warts,
pegs, pits, or ridges at maturity) but smooth when young), round to elliptical in Tuber, round in
other genera; light to dark brown at maturity. Asci typically 1-6-spored in Tuber, 4-8-spored in
other genera, randomly imbedded in the tissue between the veins (or sometimes forming a palisade
or hymemum in Pachyphloeus and Choiromyces); not amyloid.

THE “true” truffles or “earth nuts,” as they are sometimes called, can be told from other
truffles by their solid, marbled interior and ornamented spores. There are two families,
the Tuberaceae (with one principal genus, Tuber) andtheTerfeziaceae. Tuber is the largest
and most famous genus of truffles. It includes the fabled black truffle (T. melanosporum
—see photo on p. 842) and white truffle (T. magnatum) of Europe as well as a number of
species endemic to North America. Tuber is an easy genus to recognize. The fruiting body
is hard and easily mistaken for a small rock or acorn. The interior is solid and marbled
TUBER & ALLIES 855

(typically whitish when young but become brown or black with white veins at maturity),
and has the consistency of wax, i.e., it flakes or chips like a candle. The exterior of the
fruiting body is smooth in some species and warted in others, and may or may not be lobed.
Microscopically, Tuber is distinct by virtue of its relatively large, round to elliptical, geo¬
metrically-patterned spores and one- to six-spored asci that are imbedded randomly in
the tissue between the veins. However, most species of Tuber are practically indistin¬
guishable from each other when young (i.e., without mature spores) and not much easier
to differentiate at maturity. A few are distinctive in color, odor, and habitat, but most can
only be identified by examining the spores under the microscope. Even then it isn’t easy,
because the ornamentation of the spores changes as they mature and the size is notoriously
variable (a one-spored ascus tends to produce significantly larger spores than a two- or
four-spored ascus in the same fruiting body). In other words, Tuber may be an easy genus
to recognize, but the identification of its species often requires the services of a specialist.
The second family of “true” truffles, theTerfeziaceae, encompasses five genera. Micro¬
scopically these genera differ from Tuber in several respects (see the key), but they can
often be told in the field on an individual basis. Pachyphloeus, for instance, has a more or
less round, warted fruiting body with a grayish-olive to blackish-olive interior marbled
with paler veins, and it often has a tuft of mycelium at its base; Delastria, on the other hand,
is often pink- or reddish-tinged; Terfezia is partial to sandy soil in arid or semi-arid regions;
Hydnobolites has a very distinctive gristly or fatty texture and pale color, while Choiro-
myces is even harder than Tuber and tends to be rougher and more copiously lobed.
Tuber is a fairly sizable genus, with about 60 known species and an equal number of
synonyms. Roughly half of these species occur in California and Oregon, making the west
coast the best truffle territory in North America. Tubers take an inordinately long time
to mature—several weeks or even months. In our area they typically begin developing in
the winter, which means they mature in the spring (March-June), after most other mush¬
rooms have long departed. They are mycorrhizal with both hardwoods and conifers, but
are particularly abundant under oak and Douglas-fir. Some of our species (e.g., T.
gibbosum) are good edibles, though not as distinctively flavored, perhaps, as their Euro¬
pean counterparts. Many other North American species have yet to be tried. As already
mentioned, differentiating the various species can be extremely difficult. Fortunately,
none are known to be poisonous. Alas, the famous European truffles (T. magnatum,
T. melanosporum, T. aestivum) do not seem to occur here, though special truffle hounds
have been flown in from Italy to look for them.
Terfezia is the largest genus in the Terfeziaceae. However, its dozen or so species occur
mostly in southern and/ or arid regions, and have yet to be found in California. The other
four genera in the Terfeziaceae are very small. Little is known of the edibility of the N orth
American representatives, but Terfezia arenaria, a large (5-12 cm) Mediterranean species
that grows in sandy soil (often with rock rose or Cistus) is prized in Islamic countries and
was a favorite with the Romans and Greeks. Four common Tubers and three members of
the Terfeziaceae are described here, and several others are keyed out.

Key to Tuber & Allies

1. Fruiting body white or pale-colored when fresh; interior with a very distinctive gristle-like
texture .Hydnobolites californicus, p. 857
1. Not as above; interior not gristle-like . 2
2. Exterior of fruiting body black to slate-violet or greenish-black, usually warted; interior usually
greenish, grayish, or blackish at maturity (with paler veins). 3
2. Not with above features .4
3. Spores smooth; widely distributed, but on west coast occurring mainly with Douglas-fir ....
.(seeBalsamia& Allies, p. 852)
3. Spores ornamented at maturity; found mainly in eastern North America and Europe .
.Pachyphloeus melanoxanthus (see P. citrinus, p. 856)
856 TUBERALES

4. Exterior of fruiting body usually warted and often brightly colored; interior either remaining
pallid at maturity or becoming olive to grayish to blackish with paler veins; mycelial tuft often
present at base of fruiting body; spores smooth or ornamented with spines or pegs . 5
4. Not as above; mature interior usually brown to reddish with white veins (but usually pallid when
young); exterior warted or not; mycelial tuft usually absent; mature spores ornamented . . 6
5. Spores smooth; interior whitish or pallid even in age .(seeBalsamia& Allies, p. 852)
5. Spores ornamented with pegs at maturity; interior pallid when young but becoming olive,
grayish, or darker at maturity . Pachyphloeus citrinus & others, below
6. Exterior of fruiting body covered with warts (warts often small).7
6. Exterior of fruiting body smooth, cracked, downy, pitted, etc., but not warted . 8
7. Found in eastern North America; exterior tawny becoming distinctly reddish or brown at
maturity; interior usually brick-red or reddish-brown with paler veins (at maturity) .
. T. canaliculatum (see T. gibbosum, p. 858)
7. Not as above; found in western North America T. murinum & others (see T. gibbosum, p. 858)
8. All of the spores round at maturity and alveolate (pitted-reticulate) .9
8. Spores spiny or alveolate at maturity, at least some of them elliptical or broadly elliptical 10
9. Fruiting body 1-10 cm broad, often lobed and very hard; spores with numerous small pits
like those on a golf ball .Choiromyces alveolatus, p. 858
9. Not as above; fruiting body 1 -3 (5) cm broad; exterior often with minute white hairs or patches of
hairs (i.e., pubescent) . T. calijornicum & others, p. 860
10. Associated with Douglas-fir; odor often garlicky when mature and the peridium (skin) often
cracking in age; spores alveolate (pitted-reticulate) . T. gibbosum, p. 858
10. Not as above . 11
11. Found in Texas and along the Gulf Coast . T. texensis (see T. gibbosum, p. 858)
11. Not as above . 12
12. Spores spiny; exterior of fruiting body brown to cinnamon-colored when mature (but usually
paler when young) . T. rufum, p. 861
12. Spores alveolate (pitted-reticulate); exterior of fruiting body usually some shade of brown or
yellowish-brown when mature, but sometimes reddish-brown (especially when old) .
. T. separans & many others, p. 859

Pachyphloeus citrinus (Berry Truffle) Color Plate 213

FRUITING BODY underground, 0.5 -3 cm broad, more or less round (spherical) to
slightly lobed or tapered below, firm but not hard; apex often with a depression, circular
furrow, or cluster of furrows; base often with a tuft of mycelium. Exterior usually divided
into polygonal warts; color variable, but usually bright to dull orange to brown. IN¬
TERIOR more or less solid, composed of sterile veins which often converge toward the
apex or depression and form elongated pockets of fertile tissue between them; entirely
whitish when young, becoming grayish to grayish-olive with paler (white to yellowish)
veins, and eventually becoming dark olive to blackish with yellow to pale olive veins. Odor
of mature specimens sometimes pungent (“like rotting weeds”—Herb Saylor) after col¬
lecting, at other times mild. SPORES (11) 13-21 microns, round, smooth and hyaline
(colorless under the microscope) at first, becoming spiny, and at maturity the spines
enlarging into broader, conical to truncate warts or “pegs” that look like miniature golf
tees; usually yellowish at maturity. Asci typically 8-spored, forming an irregular palisade
(hymenium) along the sterile veins and/ or randomly imbedded in the surrounding tissue.
HABITAT: Solitary to gregarious in soil under both hardwoods and conifers; very widely
distributed (throughout most of North America and Europe). In our area it fruits, like
other truffles, in the winter, spring, and early summer. I have found it several times under
tanoak and madrone in June and July, and it has turned up repeatedly under live oak in
the Guadelupe Mines area near Almaden, California.
EDIBILITY: Specimens I sampled had little taste, but were immature.
PACHYPHLOEUS 857

COMMENTS: This species appears to be one of the most widespread of all the truffles.
The combination of orange to brown, warted exterior and solid, grayish-olive to blackish
interior is distinctive. The “pegs” on the spores are also unusual—at maturity each is
usually tipped with a small depression that makes it look like a golf tee. The fruiting bodies
are not nearly as hard as Tubers, and are usually rounder. Local material is usually bright
orange when immature and has the aspect of a madrone berry (see color plate) or the fruit
from a strawberry tree (a European madrone). Other species: P. virescens is said to be
similar, but has a dull green exterior and yellower interior. It was originally collected in
Los Gatos, California, but is also reported from Nebraska! P. melanoxanthus of eastern
North America and Europe has a black to greenish-black, warted exterior and a grayish
to blackish interior marbled with hollow or greenish veins (at maturity) and sometimes
has a short “stalk” of mycelial fibers. P. conglomeratus has slightly amyloid asci.

Hydnobolites californicus (G ristly T ruffle)

FRUITING BODY underground, 0.5-4 cm broad, roundish to oval, usually lobed or
folded but quite compact, gristly or fatty and rubbery in texture (especially the interior),
not hard. Exterior often roughened but lacking warts, whitish to buff or dingy yellowish,
often becoming ochre-tan or dingy brown in age or as it dries. INTERIOR more or less
solid, with several narrow, meandering sterile veins or canals; white to slightly grayish,
but often discoloring brown when cut and dried or in old age. Odor mild or sometimes
musty at maturity. SPORES 14-18 (24) microns, round, very coarsely alveolate at
maturity, the ridges of the alveoli(pits) projecting like needles from the edges of the spore;
hyaline (colorless) to yellowish or pale brown under microscope. Asci mostly 8-spored,
imbedded in the tissue between the veins, not arranged in a palisade (hymenium).
HABITAT: Solitary, scattered, or in groups or pockets of several individuals in soil and
humus under trees and in woods. As its name implies, it was originally discovered in Cali¬
fornia, but appears to be widely distributed in North America. In our area it is fairly com¬
mon from January to July under many trees, but especially oak.

EDIBILITY: Prized by slugs, despised by humans. The texture is fatty but the flavor is not.
COMMENTS: This truffle is easily recognized by its pale color and gristly or fatty con¬
sistency. The latter feature is particularly striking and almost without parallel among
the truffles. The interior is whitish except in old age and the fruiting body is never as hard
as a Tuber. The very coarsely alveolate spores are also diagnostic. Other species://, cere-
briformis of Europe is said to have larger spores; it has also been found in Iowa.

Hydnobolites californicus. Note pale color and lobed fruiting body.

858 TUBERALES

Choiromyces alveolatus (Hard Truffle)

FRUITING BODY usually underground, (0.5) 1-10 cm broad, nearly round or more
often lobed or knobby (potato-like); very hard, without a mycelial tuft at the base. Exterior
usually roughened or minutely downy, whitish when very young becoming yellowish to
tawny, brown, or rusty-brown in age. INTERIOR more or less solid and very firm, consis¬
tency rather like hard wax; at first white or pallid, but becoming marbled with darker
(yellowish to orange, yellow-brown, or rusty-brown) veins or chambers which are usually
solid but occasionally empty. Odor mild or distinctive (see comments). SPORES 20-30
(36) microns, round, smooth at first but covered with numerous rounded pits (alveolate)
like a golf ball at maturity; yellowish to brown under the microscope. Asci mostly 8-spored
(but many appearing M-spored in younger specimens), arranged in a palisade (hy-
menium) lining the veins or chambers (but see comments).
HABITAT: Solitary to gregarious in soil under trees and in the woods; known only from
western North America, occasional (if you’re looking for truffles) in the late winter, spring,
and early summer. It is particularly numerous in the Sierra Nevada, but also occurs along
the coast. Like the Tubers, it takes several weeks or months to mature.
EDIBILITY: Tempting and probably edible, but I can find no mention of anyone eating it.
COMMENTS: This is a variable species as evidenced by its plethora of pseudonyms (e.g.,
C. cookei, Piersonia alveolata, P. bispora). It is likely to be mistaken for a Tuber because
of its solidity, but microscopic examination reveals that the spores are finely pitted like
golf balls and the spore-bearing cells (asci) are arranged in nests or a palisade rather than
being randomly imbedded in the tissue. (Apparently the spores tend to form first in“nests”
which represent the inner termination of the veins; it is only in older specimens that they
line the entire lengths of the veins.) It is one of our largest truffles, capable of attaining the
size of a fist! The odor is sometimes distinctive. Helen Gilkey says of one collection: “Odor
at first resembling desiccated coconut, changing as [it] dries to that of strong cream cheese.”

Tuber gibbosum (Oregon White Truffle)

FRUITING BODY usually underground, 1.5-5 (8) cm broad, nearly round to irregularly
knobby or potato-like; firm or hard, without a basal mycelial tuft. Exterior minutely
downy or irregularly roughened but not warted, whitish when young becoming pale buff
to tan or brown, then usually developing darker (reddish to purple-brown) areas when
fully mature; often cracking in age. INTERIOR solid, marbled, crisp; whitish when
young, the fertile tissue becoming brown to dark brown to brick-red when mature, the
meandering sterile veins remaining whitish. Odor usually strong and garlicky when fully
mature. SPORES 35-52 * 17-40 microns, elliptical or elongated, reticulate-alveolate
(ridged and pitted with shallow depressions) when mature but smooth when very young;
dark yellow-brown to brown under the microscope when mature. Asci mostly 1- to
6-spored, randomly imbedded in the tissue between the veins.
HABITAT: Solitary, scattered, or gregarious in woods and at their edges, associated
mainly if not exclusively with Douglas-fir (usually trees between the ages of 8 and 65
years); found from California to British Columbia, but especially common in Oregon.
Although it normally grows underground, I have found specimens on the surface. (They
were probably dug up by squirrels, then rejected for reasons known only to squirrels.)
EDIBILITY: Edible and choice, but widespread collecting can be destructive! It smells
like the white truffle of Europe (T. magnatum), and some people proclaim it just as good.
It can now be bought—for a slightly more reasonable price than T. magnatum.
COMMENTS: Species of Tuber are often difficult to identify in the field, but this one
can be told by its growth with Douglas-fir, relatively large size (when mature), tendency
Left: Choiromyces alveolatus (p. 858). This specimen is over 5 cm broad! (Herb Saylor) Right: Close-
up of the marbled interior of a mature Tuber separans. When young the interior is whitish. Other
Tubers have very similar interiors. See next page and color plate for views of exterior.

to develop cracks in age, and strong garlicky odor (when present). Like other Tubers,
it has a marbled white-and-brown(or reddish-brown) interior when mature. The narrowly
elliptical spores are also very distinctive, providing you have a microscope. Other species:
T. besseyi is similar, but has an “olive-buff’ exterior and slightly longer spores. T. canali-
culatum is a sizable choice edible with a distinctly warted, brown to reddish or tawny
exterior. It is found in the summer and fall in eastern N orth America (a region not known
for its truffles). Several western truffles also have a warted exterior (e.g., T. murinum,
T. linsdalei, T. gardneri, and T. harknessii), but they are difficult to distinguish without
a microscope. T. texensis (of Texas, naturally) should also be mentioned. For other species
with elliptical, alveolate spores, see T. separans.

Tuber separans (Acorn Truffle) Color Plate 212

FRUITING BODY underground, very firm or hard; (0.7) 1^4.5 cm broad, round to oval
or sometimes lobed (potato-like), but usually with no more than three major lobes; lacking
a basal mycelial tuft. Exterior smooth to very minutely roughened but not warted; whitish
when very young but soon becoming uniformly pale brown to dingy yellow-brown (about
the color of an oak gall), then eventually developing dark reddish-brown to dark brown
areas (or darkening overall) as it reaches full maturity. INTERIOR solid, very firm and
crisp, flaking or chipping like wax, marbled; white when young, the fertile tissue becoming
light brown and then dark brown with age and the meandering sterile veins remaining
whitish. Odor at maturity slight, difficult to describe. SPORES (30) 34-58 * 28-50 (56)
microns, broadly elliptical to round, brown and alveolate (pitted-reticulate) at maturity,
with few to many pits. Asci 1-4 (6)-spored, imbedded randomly in tissue between veins.

HABITAT: Widely scattered to gregarious in soil under oak and other hardwoods; known
only from the west coast, fruiting mainly in the spring and early summer (at least in our
area). I have found more than sixty specimens growing together in loose soil, associated
with tanoak or possibly madrone, in June and July.
EDIBILITY: Edible. It is mild or slightly nutty like Boletus edulis, but much crisper.
Slice it thinly and saute very briefly (about one minute) or the flavor will be lost.
COMMENTS: The above description is drawn from a single large collection made near
Santa Cruz, California, and thus may not represent the full range of variation within the
species. The yellowish to brown color of the hard, marble- to walnut-sized specimens was
quite constant, and they were frequently confused with acorns buried in the duff (see
photo on p. 860). The growth with hardwoods and lack of a strong odor at maturity distin¬
guish it from T. gibbosum, while the exterior is not pubescent as in T. californicum

859
Can you find the acorn among these prime examples of Tuber separans? If not, you aren’t alone—
I didn’t discover it until I’d brought home all of these truffles, which were found under a single tanoak.
Many other objects or “pseudotubers” (rocks, animal dung, etc.) can also be mistaken for truffles!

and the different color and alveolate spores separate it from T. rufum. However, there are
many very similar species with elliptical to nearly round, alveolate spores that can only be
differentiated with great difficulty. Part of the problem is that there are no up-to-date
keys available for the North American species, and the spore sizes (an important
feature) are useful only when correlated with the number of spores in each ascus. In other
words, the identification of most Tubers is best left to truffle experts such asJamesTrappe
(who identified the above-mentioned collection as T. separans). Among the many other
Tubers with elliptical, alveolate spores are: T. monticola, a rare species found under
conifers in the Sierra Nevada; T. citrinum, also rare, with a smooth, pale yellow exterior
in youth; T. dryophilum, a very widely distributed species with smaller, coarsely alveolate
spores and a yellowish-brown exterior at maturity; T. levissimum, a thicker-skinned
species that is also widespread; T. shearii, with large, coarsely alveolate, broadly elliptical
spores; and T. irradians, with many nearly round, coarsely alveolate spores. All of these
species have spores about the same size as T. separans or smaller. See also T. gibbosum.

Tuber californicum (California T ruffle)

FRUITING BODY underground, (1) 1.5-3 (5) cm broad, round to oval or more often
irregularly lobed and/ or pitted; very firm, without a basal mycelial tuft. Exterior minutely
but evenly pubescent (covered with tiny hairs) and whitish when young, becoming mot¬
tled with darker (olive to dingy ochre to brown) areas as it matures, but usually retaining
patches of the white pubescence; sometimes cracked in age. INTERIOR solid, firm,
marbled, chipping like wax, whitish when young, becoming dark brown with large
meandering white sterile veins at maturity. Odor often distinctive when old (rather cheesy
or like “gourmet” crackers). SPORES (30) 39-52 microns, round, alveolate (reticulate-
pitted), brown at maturity. Asci 1 -4 (6)-spored, imbedded randomly in flesh between veins.
HABITAT: Solitary to gregarious in or on soil (but under humus) in woods and at their
edges; common (for a truffle) in California and Oregon, also reported from Idaho and
Ohio. In our area it is sometimes abundant under oak in the late winter and spring, but I
have also collected it on numerous occasions under Douglas-fir as late asJuly. The fruiting
bodies are often attacked by slugs, nematodes, and fly larvae.

860
Tuber californicum. This common species always has round (spherical) spores, but can usually be
recognized in the field by the patches of white pubescence on its exterior.

EDIBILITY: Edible. Some people detect a bitter taste, but the specimens I sampled had
a very strong mushroomy flavor that would go well in sauces or gravies.
COMMENTS: The critical feature of this Tuber is its uniformly round( not round toellip-
tical), alveolate spores. However, it can usually be told in the field by its pubescent or
downy exterior (use a hand lens!) and tendency to be quite knobby. T. sphaerosporum
also has uniformly round spores, but it lacks the pubescent exterior of T. californicum and
has fewer and larger pits on its spores. It occurs in eastern North America and Gary
Menser has found it under willow in Colorado. For alveolate-spored species with at least
some elliptical spores, see T. separans and T. gibbosum.

Tuber rufum (Cinnamon Truffle)

FRUITING BODY underground, 0.5-2.5 (3.5) cm broad, round to somewhat irregular or
potato-like, with or without one or more furrows; very firm or hard, without a basal tuft
of mycelium. Exterior smooth or broken up into patches or “eyes” but not conspicuously
warted, color variable: brown to cinnamon when mature (brown to orange- or reddish-
brown), usually duller and lighter (whitish to light brown) when young; sometimes with
whitish, pinkish, or golden areas mixed with darker shades. INTERIOR solid, firm, chip¬
ping like wax, marbled; whitish when young becoming grayish-brown to brown in age;
sterile veins large, remaining whitish. Odor not very distinctive. SPORES 20-48 * 17-32
microns, elliptical to nearly round, brown and covered with spines at maturity. Asci
1-4 (7)-spored, randomly imbedded in the tissue between the veins.
HABITAT: Solitary to gregarious or clustered beneath the soil under oaks and other
trees; widely distributed and common. In California it occurs principally with live oak
and is our most common Tuber. I have collected hundreds of marble-sized specimens
under oaks in the spring. It also fruits in the late winter, but takes several weeks to mature.

Tuber rufum (also known as T. candidum) is our most common truffle. The paler specimens in this
photo are younger, the darker ones older. Note the relatively smooth exterior and marbled interior.
862 TUBERALES

EDIBILITY: Edible and fairly good; it has a faintly nutty flavor.

COMMENTS: Also known as T. candidum and more exactly called T. rufum var. niti-
dum, this common truffle is best told in the field by its modest size and brown to cinnamon-
colored peridium (at maturity), and in the laboratory by its spiny rather than pitted or reti¬
culate spores. The latter feature sets it apart from most other North American Tubers. As
in other Tubers, the interior is solid and marbled. Other species: T. harknessii is an oak-
loving western species with spiny spores and a distinctly warted exterior.

ELAPHOMYCES (Deer Truffles)

Small to medium-sized, underground, mycorrhizal fungi frequently incrusted with dirt. FRUIT¬
ING BODY round to oval, usually hard and lacking an obvious base. Exterior smooth or more
often covered with small warts. Peridium (skin) thick (usually 2-5 mm), tough and usually hard.
INTERIOR a single large cavity soon filled with cottony white tissue that becomes dark (brown
to black) and powdery at maturity; sterile white bands of tissue often visible in intermediate stages.
STALK and columella absent; sterile base and mycelial tuft also absent. SPORES usually dark
brown to blackish at maturity, often so dark under the microscope that the ornamentation is dif¬
ficult to make out; round, ornamented with warts or spines. Asci typically round at maturity (often
irregular when young), 8-spored, not forming a hymenium, soon disintegrating, not amyloid.

THESE thick-skinned truffles are unique among the Ascomycetes in having a dark
powdery spore mass at maturity. The powdery texture, which results from early disinte¬
gration of the asci,* plus the thick skin can lead to confusion with the earthballs (Sclero¬
derma). However, earthballs usually grow near the surface of the ground and have a
distinct base or point of attachment to their substrate, and are not usually incrusted with
rootlets (mycorrhizae), whereas Elaphomyces species grow up to 12 inches under the soil,
lack a distinct base, and are usually incrusted with dirt and mycorrhizae.
Elaphomyces differs from other truffle genera in several additional respects. The wall
of the fruiting body, in addition to being thick and hard, is marbled in some species (when
sectioned). Also, the interior, although apparently homogeneous in old age, is actually
divided into several large chambers by thick white bands of sterile tissue when younger.
Finally, Elaphomyces is the only genus of fungi to be parasitized by certain species of
Cordyceps (see p. 879). For these reasons, among others, Elaphomyces has traditionally
been separated from other truffles and placed in its own order and family. However, it
is now thought to be less dissimilar to other truffles than once believed.
Elaphomyces is mycorrhizal with both hardwoods and conifers. In our area it fruits
throughout the mushroom season and can even be found in the summer if you look hard
enough (or dig deep enough!). It is among the most abundant of all underground mush¬
rooms, but is seldom seen or collected, perhaps because the soil-incrusted fruiting bodies
look like balls of dirt. In Europe, species of Elaphomyces have been used as aphrodesiacs
and cheap truffle substitutes, but their rindlike skin is too tough and the mature spore mass
too powdery to be worth eating. Over 30 species have been described, based largely on
microscopic criteria. Two common and farflung representatives are depicted here.
Key to Elaphomyces
1. Fruiting body with a distinct base or point of attachment to its substrate; growing underground
or above it; spores borne on basidia .(seeScleroderma, p. 707)
1. Fruiting body usually lacking an obvious base, but exterior often incrusted with dirt and my¬
corrhizal rootlets; growing underground; spores borne inside asci .2
2. Peridium (skin) marbled when sliced open (as shown on next page) E. muricatusgroup, p. 863
2. Peridium not marbled when sectioned .E. granulatus group, p. 864

*The early disintegration of the asci is a unique and puzzling feature of Elaphomyces that has caused even myco¬
logists to confuse it with Scleroderma. The spores are often highly compressed and irregular in shape while inside
the asci, presumably as a result of pushing against the ascus wall. It may very well be that this pressure causes the
asci to pop like balloons!
Elaphomyces muricatus group, mature specimens. Note how spore mass is dark and powdery in the
two sectioned specimens (but the central core is still cottony in one of them), and how the exterior of
central specimen is incrusted with dirt and mycorrhizal rootlets.

Elaphomyces muricatus group (Marbled Deer Truffle)

FRUITING BODY underground, 2-5 cm broad, round to oval or somewhat lobed, very
firm. Exterior yellow-brown to ochre-brown and covered with minute, hard, pointed
warts that give it a pimpled appearance, but the warts usually obscured by a crust of soil
and tiny rootlets (mycorrhizae) that is easily stripped or brushed away. Peridium (skin)
thick (2-5 mm), hard and rindlike, marbled when sectioned (dark brown to purplish-
brown with whitish to vinaceous-tinged veins). INTERIOR at first hollow, soon stuffed
with cottony white hyphae, then darkening to grayish, lilac, or purplish and divided into
chambers by sterile bands, finally become brownish-black to black and uniformly
powdery. Odor not very distinctive. SPORES 18-30 * microns, round, warted or warted-
spiny, dark brown to black under the microscope. Asci round to pear-shaped or irregular,
mostly 8-spored, not forming a palisade (hymenium) and disintegrating soon after the
spores form (and before they are completely mature).

Elaphomyces muricatus group. Close-up of a sectioned specimen, showing the thick marbled peri¬
dium (skin). The spore mass (interior) is still cottony and divided into chambers by white sterile tissue.
Elaphomyces muricatus group. Note how the thick skin is marbled in sectioned specimen at left, and
how the exterior is finely warted in specimen on right. E. granulatus and its close relatives (not illus¬
trated) look quite similar, but do not have a marbled peridium.

HABITAT: Solitary to gregarious in soil or duff in woods; widely distributed and fairly
common (if you’re digging for truffles). It is said to prefer pine woods, but in our area I
have found it as early as October under knobcone pine and manzanita and as late as July
under tanoak and madrone.
EDIBILITY: Unknown (but see comments on edibility of E. granulatus).
COMMENTS: This lesser-known cousin of E. granulatus is best told by its marbled
peridium (see photographs). Like other Elaphomyces species, it is sometimes parasitized
by Cordyceps, and is easily told from other truffles by its thick rindlike skin and dark,
cottony to powdery mature spore mass (interior). The latter features can lead to confusion
with the earthballs (Scleroderma), which, however, do not have a marbled peridium.
Other species of Elaphomyces with a marbled peridium (e.g., E. verrucosum, E. varie-
gatus) differ from E. muricatus microscopically.

Elaphomyces granulatus group (Common Deer Truffle)

FRUITING BODY underground, 2-5 cm broad, round to somewhat oval, very firm.
Exterior usually covered with small hard warts (but these often hidden by a crust of soil,
yellowish mycelium, and/or mycorrhizal rootlets); sometimes pallid when young but
usually pale to dingy ochraceous or yellow-brown. Peridium (skin) rindlike, very firm and
thick (2-5 mm), showing a very thin yellowish outer layer (when sectioned) and a thick
white to grayish inner layer that may feature darker (brown) zones, but not marbled.
INTERIOR at first hollow, soon stuffed with cottony tissue, eventually becoming
powdery when spores mature; white at first, soon grayish to purplish (and often separated
into chambers by whitish bands), finally becoming blackish and powdery. Odor not very
distinctive. SPORES 24-45 (65) microns, round, thick-walled, blackish-brown to very
dark reddish-brown under the microscope, ornamented with short spines or warts. Asci
mostly 8-spored, round to pear-shaped, not forming a hymenium, disintegrating before
the spores are fully mature.
HABITAT: Solitary, scattered, or gregarious in soil or humus under conifers or less
commonly hardwoods; widely distributed and very common, but seldom seen by the
average mushroom hunter because it grows underground. It is usually found 2-3 inches
(5-8 cm) below the surface, often imbedded in or resting on clay soil at the point where it
meets the humus layer. In many regions it is parasitized by species of Cordyceps—which
serve as an above-ground indicator of its presence—but I have yet to observe this in our
area. It fruits throughout the mushroom season and can sometimes be gathered by the
bushel. I have found it under hemlock and pine in northern California, and Herb Saylor
reports prolific fruitings from Mendocino County. Alexander Smith calls it“perhaps the
most common hypogeous [underground] fungus in North America”—and he should
know, since he has collected so many of them!
ELAPHOMYCES 865

EDIBILITY: Edible according to some reports, but not choice. In Europe it has been used
for centuries as an aphrodesiac and truffle-substitute.
COMMENTS: The thick rindlike skin and purple-gray to black, cottony to powdery
interior distinguish Elaphomyces from all other underground Ascomycetes. E. granulatus
and its close relatives differ from the E. muricatus group in having a non-marbled peridium
(as seen in sectioned specimens). The earthball genus Scleroderma is similar, but produces
spores on basidia, usually has a distinct base or point of attachment to the substrate, lacks
the small hard warts and outer crust of soil and mycorrhizae so frequently found in E.
granulatus, and usually grows nearer to the ground surface (or often above it). Other spe¬
cies of Elaphomyces with a non-marbled peridium are best differentiated microscopically.
One, E. subviscidus, has a smooth skin, dark brown mature spore mass, and smaller spores.

Earth Tongues
HELOTIALES
THIS large order includes hundreds of small stalked or cuplike Discomycetes with in-
operculate asci (i.e., each ascus has a pore at its tip through which the spores are expelled,
but no operculum or “lid” as in the Pezizales). The most conspicuous members of this
order are called earth tongues because of their clublike to tongue-shaped fruiting bodies.
Many others are cup-shaped or disclike, with or without a stalk.
The larger members of this order are saprophytic on soil, humus, and wood, while most
of the smaller types are parasitic or saprophytic on plant stems, leaves, and other tissues.
None are prized edibles, being too small or too tough or too small and too tough to bother
eating. There are several families and over 150 genera in the Helotiales. These are defined
largely on the basis of microscopic features, and only a smattering of the larger or more
colorful species are described here. These have been divided into five groups, keyed below,
based on the shape and texture of the fruiting body.
Key to the Helotiales
1. Fruiting body with a cap and stalk, the cap rounded to convex or wrinkled but not cuplike
(concave) or disclike . 2
1. Fruiting body variously shaped, sometimes with an enlarged “head,” but without a clearly
differentiated, rounded to convex or wrinkled cap . 4
2. Stalk very thin (usually less than 2 mm); cap bladderlike (hollow), whitish to yellowish; found in
eastern North America, often clustered; spores borne on basidia (see Aphyllophorales, p. 548)
2. Not as above; spores borne inside asci . 3
3. Cap with a sterile underside and an abrupt edge or margin (but the margin usually inrolled or
tucked in toward the stalk) .Leotia& Cudonia, p. 872
3. Not as above; cap merely an enlarged, differentiated “head” that lacks an abrupt margin and
sterile underside . 4
4. Flesh gelatinous or rubbery-gelatinous; fruiting body variously shaped but not clublike, pinkish
to reddish, purplish, brown, or black; growing on wood . Bulgaria & Allies, p. 875
4. Not as above . 5
5. Fruiting body cuplike or disclike, with or without a stalk . Ciboria & Allies, p. 877
5. Fruiting bod erect, clublike or with an enlarged or flattened “head” . 6
6. Fruiting body with large internal chambers or compartments .(see Helvellaceae, p. 796)
6. Not as above . 7
7. Entire fruiting body black or sometimes dark brown; at least some of the spores brown under
the microscope . Geoglossum & Trichoglossum, p. 866
7. Not as above; fruiting body usually lighter or brighter than above; spores hyaline (colorless)
under the microscope or tinged yellow .Microglossum, Spathularia, & Allies, p. 868
866 HELOTIALES

GEOGLOSSUM & TRICHOGLOSSUM

(Black Earth Tongues)
Small fungi usually found on ground or moss, sometimes on rotten wood. FRUITING BODY
upright, usually clublike, with or without an enlarged “head”; dark brown to black. Flesh thin,
tough, not gelatinous. STALK present, usually slender; smooth or velvety. SPORES very long
and narrow, usually septate, smooth, brown under the microscope (at least some of them). Asci
typically 8-spored, lining the upper portion of the fruiting body or the “head” if one is present;
inoperculate and not amyloid, with a pore at the tip.

THESE attractive little Ascomycetes are easily recognized by their black (or occasionally
dark brown) clublike fruiting bodies. Their color separates them from the fairy clubs
(Clavariaceae) and other earth tongues, and they lack the pimpled surface or white spore
powder so often found inXylaria. Their very long(up to250 microns!) brown partitioned
(septate) spores are also distinctive.
Both Geoglossum and Trichoglossum are charter members of the family Geoglossaceae.
In the more common of the two, Trichoglossum, brown lance-shaped cells called setae
protrude from the surface of the fruiting body, giving it a velvety texture. In Geoglossum,
the setae are absent (at least in the fertile portion), and the texture varies from smooth
to viscid to only slightly velvety.
The black earth tongues are saprophytic on humus, soil, moss, or occasionally rotten
wood. Several of the more than two dozen North American species are common, but all
are difficult to distinguish from their surroundings because of their dark color and small
size. They are also difficult to distinguish from each other, even with a microscope. For
this reason, only one representative from each genus is described here. Neither is worth
eating.
Key to Geoglossum & Trichoglossum
1. Flesh usually white; exterior of fruiting body often roughened or minutely pimpled but not hairy
or velvety; growing on wood (but wood often buried); asci borne in flasklike “nests” (peri-
thecia) imbedded in the fruiting body .(see Pyrenomycetes, p. 878)
1. Not as above . 2
2. Surface of fruiting body distinctly viscid when moist, often glistening, not velvety.
. G. glutinosum & others, below
2. Fruiting body not viscid .:. 3
3. Surface of fruiting body (especially stalk) distinctly velvety; fruiting body often (but not always!)
with a spade-shaped or flattened “head”. T. hirsutum & others, p. 867
3. Not as above; fruiting body not velvety or only very slightly so, usually clublike or twisted, but
sometimes with a distinct “head” . . . G. nigritum & many others (see G. glutinosum, below)

Geoglossum glutinosum (Viscid Black Earth Tongue)

FRUITING BODY 1.5-6 cm tall, cylindrical to club-shaped. Upper (fertile) portion 3-6
mm wide, often flattened or slightly twisted but otherwise not sharply differentiated from
lower portion (stalk); surface black, smooth, viscid (at least when moist). Flesh tough,
usually brownish, not gelatinous. STALK occupying lower 1 / 3-2/3 of fruiting body, 2-3
mm thick, dark brown to black, viscid when moist, usually smooth. SPORES 60-90 * 4-5
microns, greatly elongated, smooth, brown under the microscope, with 0-7 (usually 3 or 7)
septa (partitions).
HABITAT: Solitary, scattered, or in small groups in humus, moss, or sometimes rotten
wood, usually in the woods; widely distributed but infrequently encountered. I’ve found
it several times in our area in the late fall, winter, and early spring, but never in quantity.
EDIBILITY: I know of no one who bothers collecting Geoglossums, but Captain Charles
Mcllvaine says of this species; “Over a quart found in one patch. Stewed it is delicious.”
Left: Geoglossum glutinosum. Note all the debris clinging to the viscid fruiting bodies. Right: Geo-
glossum nigritum (see comments under G. glutinosum) is a fairly common species with a clublike or
twisted fruiting body that is not flagrantly velvety.

COMMENTS: This dark earth tongue has many look-alikes (see below), but is one of the
few species with a distinctly viscid, glistening surface when moist. Similar viscid species
include: G. affine, rare, with shorter spores; and G. difforme, larger (3-12 cm tall), whose
spores have 8-15 septa. The genus Geoglossum also includes many similar dark brown to
black, non-viscid earth tongues that can only be differentiated microscopically. These
species are not as velvety as Trichoglossum and do not often have a well-defined “head.”
Some of the more common and widespread ones are: G. glabrum, a smooth-stalked species
up to 10 cm tall; G. simile, the most common species in eastern North America, stalk often
scurfy or minutely scaly; and G. nigritum (see photo), the most common species in our area
(but more widely distributed), with strongly curved paraphyses (sterile cells) whose tips
are scarcely enlarged. All of these have brown spores, but some Geoglossums have both
brown and hyaline (colorless) spores, including: G.fallax, whose hyaline spores are non-
septate; and G. alveolatum and G. intermedium, whose hyaline spores are septate. There
are also several similar, dark Microglossum species whose spores are ^//hyaline, including:
Microglossum atropurpureum, fruiting body dark brown to purplish or black; M.
fumosum, yellow-brown to brown; and M. olivaceum (see comments under M. viride),
greenish-brown to dark brown with very short spores (10-18 microns long).

Trichoglossum hirsutum (Velvety Black Earth Tongue)

FRUITING BODY 2-8 cm tall, cylindrical to club-shaped or more often with a distinct
“head” (i.e., fruiting body shaped more or less like the tongue of a bell). Fertile “head” when
distinct 3-8 mm broad, usually flattened laterally or compressed, oval or elongated to
spade- or arrowhead-shaped; surface dry, minutely hairy or velvety, sometimes wrinkled,
black. Flesh thin, tough, usually brownish. STALK thin (1-4 mm thick), more or less
equal, densely velvety, often twisted or curved, tough, black. SPORES 80-195 (210)* 5-7
microns, greatly elongated, smooth, brown under the microscope, typically with 15 septa
(partitions) when mature, but some varieties with consistently fewer or more septa. Both
stalk and head lined with long brown sterile cells (setae). Asci 8-spored.
HABITAT: Solitary, scattered, gregarious, or tufted in humus, moss, or soil (or occa¬
sionally on rotten wood), usually in woods; very widely distributed and common. It is the
most abundant earth tongue in our area, sometimes carpeting large tracts of humus with

867
Trichoglossum hirsutum. Note the often spade-shaped “head” of this common black earth tongue.
Left: Typical fruiting bodies. Right: These specimens were photographed with a strobe in an effort
to highlight some of the minute hairs that give them a velvety texture.

its little black clubs. In seems to favor habitats shunned by other mushrooms (e.g., red¬
wood), and usually fruits in the winter or spring. Like other black fungi, it is difficult to see.
EDIBILITY: Supposedly edible, but much too tough to be worthwhile.
COMMENTS: This dainty earth tongue is easily recognized by its black velvety fruiting
body. In California specimens the fertile portion is usually (but not always) set off from
the stem as a thickened or flattened, often spade-shaped “head.” The wonderful velvety
texture is caused by hundreds of minute projecting hairs or spines (setae) and is most
evident on the stalk, especially in dry weather. Geoglossum species are very similar, but
are not as velvety and do not normally have such a well-defined “head.” Other species of
Trichoglossum can only be differentiated microscopically. They include: T. velutipes,
with 4-spored asci and mostly 7-11-septate spores; and T. farlowii, with 8-spored asci
and 0-5 (usually 3)-septate spores.

MICROGLOSSUM, SPATHULARIA, & Allies

(Colorful Earth Tongues)
Small fungi found on ground, moss, or rotten wood. FRUITING BODY upright, usually clublike
to spatula- or tongue-shaped, or with an enlarged fertile “head”; variously colored but not dark
brown to black. Flesh often rather tough, not gelatinous. STALK usually present, often slender;
smooth, mealy, minutely scaly, or velvety. SPORES round to elliptical, spindle-shaped, or needle¬
like, smooth, hyaline (colorless) under the microscope, sometimes septate. Asci lining upper part
of fruiting body or confined to the “head” if one is present; typically 8-spored, not amyloid, with
a pore at the tip but not operculate.

THESE earth tongues are more cheerfully colored than Geoglossum and Trichoglossum,
and their spores are colorless when viewed under the microscope. They may superficially
resemble the unbranched coral fungi or fairy clubs (Clavariaceae), but bear their spores
in asci rather than on basidia and usually have a swollen or flattened, fertile “head.” The
“head,” when present, lacks the abrupt edge and sterile undersurface of a Leotia, Cudonia,
or Helvetia.
Four common genera, all members of the Geoglossaceae, are treated here. Among
these, Spathularia stands out because of its peculiar flattened, fanlike “head” that runs
down opposite sides of the stalk. Neolecta is also distinctive because of its highly irregular
shape, while Microglossum and Mitrula have more or less clublike fruiting bodies, the
latter with a clearly differentiated “head.”

868
MICROGLOSSUM, SPATHULARIA, & ALLIES 869

These earth tongues, like Geoglossum and Trichoglossum, inhabit humus, soil, moss,
and rotten wood, but do not grow on insects or truffles. None are large enough or tasty
enough to collect for the table, but several are quite beautiful, and are worth getting to
know for this reason if no other. One species from each of thefour genera isdescribed here;
several others are keyed out.

Key to Microglossum, Spathularia, & Allies

1. Fruiting body green or greenish. 2
1. Not as above . 3
2. Fruiting body olive-green to green to dark green or blue-green; stalk often minutely scurfy
or scaly .M. viride, p. 870
2. Fruiting body with only a slight greenish tinge; stalk smooth .
.Microglossum olivaceum (see M. viride, p. 870)
3. Fruiting body with a flattened (paddle-like or fanlike) fertile “head” that extends down the
the stalk on opposite sides (i.e., stalk appears to be wedged into the cap) . 4
3. Not as above; “head” absent, or if present and flattened then not runningdown stalk on opposite
sides . 5
4. Stalk reddish-brown to dark brown and velvety; found mainly in eastern North America
.Spathularia velutipes (see S.flavida, p. 871)
4. Not as above; widespread .Spathulariaflavida & others, p. 871
5. Fruiting body with a small oval or rounded “head” that is sharply differentiated (and often
differently colored) from stalk; sometimes growing in water . 6
5. Fruiting body lacking a sharply differentiated “head” (but apex may be swollen, flattened, or
broadened); not growing in water . 8
6. Typically growing on submerged sticks in running water (often in cold mountain streams); stalk
usually brownish .Vibressea truncorum (see Mitrula abietis, p. 870)
6. Not as above; growing in still water or not in water at all . 7
7. “Head” pinkish-buff to pale flesh-colored to light brown; stalk light to dark brown; found in
duff under conifers . Mitrula abietis, p. 870
7. “Head” differently colored (often yellow or orange) and stalk often white or pinkish-tinged
and/ or growing in shallow pools or on very wet soil or leaves .
.Mitrula elegans & others (see M. abietis, p. 870)
8. Fruiting body pallid to pale ochre or dingy yellowish, always growing on wood; exterior usually
roughed or minutely pimpled by the projecting tips of perithecia (flasklike nests of asci); rare
..'.(see Pyrenomycetes, p. 878)
8. Fruiting body differently colored, or if similar in color then lacking perithecia; usually terrestrial
or in moss, sometimes on rotten wood . 9
9. Fruiting body pale yellow to bright yellow to orange . 10
9. Fruiting body differently colored . 12
10. Fruiting body clublike and slender, without a broadened fertile “head”; spores borne on basidia
.(see Clavariaceae, p. 630)
10. Fruiting body usually with a broadened, often flattened fertile “head” or if not, then extremely
variable in shape (often lobed, forked, flattened, twisted, etc., and usually at least 0.5-2.5 cm
broad at the top); spores borne inside asci . 11
11. Fruiting body very irregular in shape (often lobed, forked, flattened, twisted, etc.), usually
yellow and usually found under conifers; spores usually less than 10 microns long, elliptical
to nearly round; widely distributed .Neolecta irregularis & others, p. 871
11. Fruiting body regularly clublike (but usually with a broad or flattened “head”), yellow to orange;
found mainly in eastern North America; spores long and thin .
. Microglossum rufum (see Neolecta irregularis, p. 871)
12. Fruiting body yellow-brown to dark brown, purplish, or even black; usually slender; spores
borne in asci . Microglossum atropurpureum & others (see Geoglossum glutinosum, p. 866)
12. Fruiting body differently colored (usually paler or brighter than above) and/or thick; spores
borne on basidia .(see Clavariaceae, p. 630)
870 HELOTIALES

Microglossum viride (Green Earth Tongue) Color Plate 214

FRUITING BODY 1-5.5 cm tall, clublike to tongue- or spatula-shaped (i.e., with an
enlarged “head” at maturity). Fertile “head” 4-12 mm broad, dark green to pea-green,
green, olive-green, or even bluish-green, smooth or furrowed, often flattened or com¬
pressed in older specimens. Flesh greenish, rather tough. STALK 2-5 mm thick, colored
like the head or paler green, usually thinner; surface minutely scurfy or scaly, but some¬
times smooth in age. SPORES 14-22 * 4-6 microns, sausage-shaped to spindle-shaped,
smooth, hyaline (colorless) under the microscope, not septate or septate only when old.
HABITAT: Solitary, scattered, or in groups or tufts in soil, moss, and duff under both
hardwoods and conifers; widely distributed. It is not uncommon in coastal California in
the winter and early spring, but is often overlooked because of its small size and green
color. I find it most often under redwood and tanoak.
EDIBILITY: Too small to be worthwhile.
COMMENTS: The striking green color sets apart this petite, farflung earth tongue. The
specimens in the color plate are rather young and club-shaped, but as they grow older their
“heads” will become flatter (laterally) and more distinct. The minutely scaly or scurfy stalk
is also distinctive. M. olivaceum is a somewhat similar but smoother species with an olive-
buff to greenish-brown to dark brown (not truly green) fruiting body and shorter spores.
I have found it twice in our area, but it is more widely distributed. Other species of Micro¬
glossum are either yellow to orange (see M. rufum under Neolecta irregularis) or brown
to blackish (see M. atropurpureum and M.fumosum under Geoglossum glutinosum).

Mitrula abietis (Miniature Earth Tongue)

FRUITING BODY 0.5-4 (5) cm tall, with a stalkand sharply differentiated “head.” Fertile
“head’ 1-7 (10) mm broad and high, roundish to cylindrical (elongated), with a smooth
surface; pinkish-buff to pale flesh-colored to light brown. Flesh thin. STALK 0.5-3 (4) cm
long, 1 -4 (7) mm thick, equal or tapered slightly, thin, light to dark brown (usually darker
than cap), smooth or slightly powdered above, often with brown hairs at base. SPORES
10-14 x 2-2.5 microns, elongated, smooth, not septate, hyaline (colorless) under the
microscope.
HABITAT: Scattered to densely gregarious in needle duff under northern and mountain
conifers; common in western North America in the spring, summer, and fall, but absent
in our area.
EDIBILITY: Fleshless and probably flavorless.
COMMENTS: This little mushroom sometimes grows in large carpets on the forest floor.
The sharply differentiated “head” distinguishes it from most earth tonguesand fairy clubs,
and it is usually smaller than Cudonia and lacks the abrupt cap margin and sterile under¬
side of that genus. M. elegans, sometimes called the “Swamp Beacon,” is a similar, widely
distributed and common species that fruits in very wet humus or soil or on leaves in shallow
pools. It has a creamy to bright yellow or pale orange “head” and a white or pinkish-tinged,
sometimes viscid stalk and can often be found under mountain conifers in the spring and
early summer. It has long passed under the nameM. paludosa, a similar European species
that differs microscopically. Other species: M. gracilis is a northern and montane species
with an ochraceous to orange-buff “head”; it usually grows in moss. M. borealis, also
northern, has a golden-yellow “head” like that of M. elegans, but has elliptical or crescent-
shaped spores. M. lunulatospora is an eastern springtime, water-loving species with a
flesh-colored to yellowish “head” and crescent-shaped spores. Vibressea truncorum looks
like a Mitrula or a miniature Leotia, but usually grows on sticks in running water (often
in cold mountain streams). It is even smaller than Mitrula and somewhat gelatinous.
NEOLECTA 871

Neolecta irregularis (Irregular Earth Tongue) Color Plate 200

FRUITING BODY 1 -7 cm tall, clublike to very irregular (lobed, sparingly branched,
grooved and twisted, etc.), usually flattened or compressed, 0.4-2.5 cm broad at apex.
Fertile surface pale yellow to bright yellow or orange-yellow. Flesh white or yellowish,
rather tough. STALK sometimes absent but usually present as a sterile, pale yellow to
white base beneath the fertile portion; 1-6(10) mm thick (usually thinner than fertile area).
SPORES 5.5-10 * 3.5-5 microns, elliptical to nearly round, smooth, not septate, hyaline
(colorless) under the microscope. Asci lining at least the upper part of (and sometimes
the entire) fruiting body.
HABITAT: Widely scattered to gregarious or occasionally tufted on ground, moss, or
duff, usually under conifers; widely distributed. Along with N. vitellina (see comments),
it occurs throughout much of the West, but is apparently absent in our area. I have seen
large fruitings under spruce and fir in New Mexico and Oregon in the summer and fall.

EDIBILITY: In my opinion, not worth collecting. Mcllvaine, as usual, dissents: “Those

fortunate enough to find this species will hunt for it again assiduously. Even raw, when cut
in strips, it makes a picturesque and delicious salad.”
COMMENTS: The bright yellow color and highly irregular shape are usually enough to
identify this species, which is also known as Spragueola (or Mitrula) irregularis. Club-
shaped specimens can be mistaken for fairy clubs (Clavulinopsis), but bear their spores
in asci rather than on basidia, are usually broader, and often have irregularly-shaped
fruiting bodies growing nearby. Microglossum rufum is a farflung earth tongue with a
bright yellow to orange fruiting body. It is much more uniform in shape (usually with a
slender stalk 2-4 mm thick and a wider, flattened “head”) and has much longer spores
(20-40 microns). It is especially common in eastern North America in the summer and
early fall. Other species: N. vitellina closely resembles N. irregularis, but is slightly smaller
and paler, and differs microscopically.

Spathularia flavida (Fairy Fan)

FRUITING BODY 1-10 cm tall, with a stalk and fertile “head.” Fertile “head” very com¬
pressed or flattened laterally, spatula- or fanlike, 1 -3 cm broad, decurrent (running down)
on opposite sides of the stalk; surface (sides) smooth or wrinkled, sometimes lobed or
contorted or with a notched apex, pallid when young becoming pale yellow to yellow, buff,
or cinnamon-buff to brownish (or occasionally pale orangish). Flesh white, not gelatinous.
STALK (1) 2-8 cm long, 2-10 mm thick, variable in shape but often thicker or swollen at
base, usually hollow; surface smooth to finely mealy but not velvety, white to yellowish or
colored like the “head” but usually paler, with white to pale yellow mycelium at base.
SPORES 30-75 (95) * 1.5-3 microns, very long and narrow (needle-like), smooth, with
one to several septa (partitions) or none at all; hyaline (colorless) under the microscope
but often yellow-brown in mass, especially when dry.
HABITAT: Scattered to gregarious or even clustered, sometimes in lines or circles, on
humus or rotten wood under conifers (especially pine) or sometimes hardwoods; wide¬
spread. It is common in the summer and fall in the Pacific Northwest and Southwest and
occurs in northern California in the fall, winter, and spring, but I have yet to find it locally.

EDIBILITY: Said to be edible, but rather tough. Captain Charles Mcllvaine describes
it as “tenacious but tender.”
COMMENTS: The peculiar flattened, paddle- or fanlike “head” that extends down oppo¬
site sides of the stalk is unique to this little mushroom and its close relatives (see next page).
It might possibly be confused with Neolecta irregularis or a Microglossum, but is not as
Left: Spathularia flavida is easily told by its flattened, paddle-like “head.” Note how stalk of central
specimen appears to be wedged into the “head,” and how fertile surface can be wrinkled or smooth.
Right: Cudonia circinans (see p. 873). Note incurved cap margin and clustered growth habit.

brightly colored and has a more consistently compressed “head.” S. clavata is a synonym.
Other species: S. spathulata is said to be similar but has smaller spores and a somewhat
darker (yellow-brown to reddish-brown) fruiting body. Its cap ranges from flattened to
rounded (as in Cudonia) but is fertile over its entire surface rather than just at the top. It
was originally collected in Big Basin State Park, California, but I have not seen it there.
S. (-Spathulariopsis) velutipes of eastern North America is a common and distinctive
“fairy fan” with orange mycelium and a velvety dark brown to reddish-brown stalk that
may be thicker at the bottom or top, plus a yellow to yellow-brown flattened “head” that
is covered by a “veil” when very young and often retains “veil” remnants at maturity.

LEOTIA & CUDONIA

Small to medium-sized fungi found on ground or rotten wood. FRUITING BODY with a cap and
stalk. CAP rounded to convex or lobed, the margin often inrolled; surface smooth or wrinkled,
viscid when moist in Leotia, not viscid in Cudonia; variously colored. Flesh gelatinous in Leotia,
not gelatinous in Cudonia. STALK smooth or scurfy or minutely scaly, well-developed, variously
colored. SPORES spindle-shaped or needle-like, hyaline (colorless) under the microscope; some¬
times septate, smooth. Asci lining the upper surface of the cap, typically 8-spored, not amyloid,
inoperculate, with a pore at the tip.

THESE two genera are unique among the earth tongues in possessing a well-developed
and clearly differentiated cap and stalk. The cap is not just a swollen “head” as in Mitrula
or other earth tongues; instead, it has a sterile underside and an abrupt margin that sets
it off from the stalk. In addition, Leotia is easily distinguished by its gelatinous to semi-
gelatinous tissue. Cudonia, in contrast, is fleshy or tough but not gelatinous. It is apt to be
mistaken for a small elfin saddle (Helvetia), but its cap is usually convex or rounded and
not as dramatically lobed as in that genus.
Leotia and Cudonia used to be classified with other earth tongues in the Geoglossaceae,
but are now placed alongside a number of genera (e.g., Bulgaria) in a larger family, the
Leotiaceae. Both are widely distributed, but only Leotia is common in our area. They
grow on the ground or on rotten wood, often in groups or clusters. Neither genus is worth
eating, although Leotia might be useful as a lubricant! Two species of Leotia and one
Cudonia are described here.

872
LEOTIA & CUDONIA 873

Key to Leotia & Cudonia

1. Flesh gelatinous or semi-gelatinous; surface of fruiting body viscid or slimy when wet .... 2
1. Flesh not gelatinous; surface not normally viscid . 6
2. Growing on submerged sticks in running water; small .
. (see Vibressea truncorum under Mitrula abietis, p. 870)
2. Not as above . 3
3. Fruiting body small (0.5-3 cm tall), whitish to pinkish or brownish, sometimes with an ochra-
ceous or lavender tinge; found on rotting hardwoods in eastern North America; rare ......
.Neocudoniella (-Leotia) albiceps
3. Not as above; usually larger; often with yellow, ochre, or greenish shades; widespread .... 4
4. Cap consistently and distinctly green or greenish . 5
4. Cap some shade of yellow, buff, ochre, or cinnamon, at times with a greenish tinge (olive-brown
or olive-ochre, etc.) .L. lubrica, p. 874
5. Stalk white to yellow or orange .L. viscosa, p. 874
5. Stalk pale green to green .L. atrovirens (see L. viscosa, p. 874)
6. Cap yellowish to olive-buff, sometimes with small “veil” fragments on margin; found in eastern
North America, usually under hardwoods .C. lutea (see C. circinans, below)
6. Not as above; differently colored; widespread . 7
7. Cap creamy to pinkish-buff, cinnamon-buff, vinaceous-buff, pale brown, or occasionally
darker; found mainly in the late summer and fall; widespread .C. circinans, below
7. Cap pinkish-buff to pinkish-cinnamon to grayish-brown or dark grayish-brown; found in
western North America under conifers, usually in the spring and summer .
. C. monticola & C. grisea (see C. circinans, below)

Cudonia circinans (Common Cudonia)

FRUITING BODY with a cap and stalk. CAP 0.5-2 cm broad, usually rounded or convex,
sometimes with a central depression and sometimes convoluted; surface wrinkled or
smooth, creamy to pinkish-buff, cinnamon-buff, vinaceous-buff, pale brown, or occa¬
sionally darker; margin usually curved down and in toward the stalk. Underside sterile,
often with radiating veins that extend up from the stalk. Flesh thin but firm, not gelati¬
nous, rather tough or leathery when dry. STALK 1.5-7 cm long, 2-12 mm thick (but
usually less than 6 mm at apex), equal or more often thicker below, stuffed or sometimes
hollow in age; drab to dark brown (usually darker than cap), usually minutely scurfy, often
longitudinally striate or ridged, especially above. SPORES (28) 32-40 (46) * 2 microns,
very long and thin (needle-like), smooth, sometimes septate (partitioned), but usually not;
hyaline (colorless) under the microscope.
HABITAT: Scattered to gregarious or often in dense clusters in humus, soil, and on rotting
wood; particularly common under conifers, but also found with hardwoods; widely
distributed. It is said to be the most common member of its genus, but I have yet to find it
in our area. In the Pacific Northwest it is fairly common in the late summer and fall.
EDIBILITY: Poisonous, at least raw. It is said to contain high concentrations of MMH
(see pp. 799 and 893 for details).
COMMENTS: The small size, convex to somewhat convoluted cap, and non-gelatinous
flesh are the hallmarks of this species. It is reminiscent of a dry Leotia, but is not as brightly
colored. It can also be mistaken for an elfin saddle (Helvella), but the shape is different
and it often forms dense groups or clusters untypical of Helvella (see photo on p. 872).
C. monticola is a similar, pinkish-cinnamon to pinkish-buff to grayish-brown westerner.
It is the largest Cudonia (up to 10 cm high, cap 1-3 cm broad), but has much smaller spores
than C. circinans {only 18-25 microns long) and a frequently compressed or even somewhat
saddle-shaped cap. It is common under conifers in northern California and the Pacific
Northwest, usually in the spring and summer (whereas C. circinans is more frequent in
the fall). C. grisea also occurs under conifers in the Pacific Northwest, but has a gray to
874 HELOTIALES

dark grayish-brown or fuscous fruiting body and usually fruits in the spring. C. lutea is a
yellowish to olive-buff eastern species that sometimes shows “veil” fragments on the
margin of the cap. It grows scattered to gregarious, usually under hardwoods.

Leotia lubrica (Jelly Babies) Color Plate 215

FRUITING BODY with a cap and stalk. CAP 0.5-4 cm broad, round to convexto slightly
lobed or knobby; surface smooth or wrinkled, viscid to slimy when moist (but sometimes
drying out), buff to yellow, ochre, olive-ochre, or sometimes cinnamon or greenish-brown;
margin usually curved in toward stalk, often lobed or wavy. Underside sterile, paler. Flesh
gelatinous (at least the central core), often translucent. STALK 2-8 cm long, 0.3-1 cm
thick, equal or somewhat thicker below, smooth or scurfy (with minute granules), hollow
or more often filled with a gel; surface viscid when moist, colored like cap or sometimes
yellower. SPORES 16-25 * 4-6 microns, spindle-shaped and sometimes curved, smooth,
hyaline (colorless) under the microscope, septate (partitioned) at maturity.
HABITAT: Solitary to gregarious or clustered in duff, soil, or very rotten wood under
both hardwoods and conifers; widely distributed. It is the most common Leotia in North
America, but is not as frequent in our area as L. viscosa. I usually find it in the winter and
spring, but in other regions it fruits in the summer and fall. Alexander Smith describes
finding massive clusters buried in sand dunes!
EDIBILITY: Harmless but glutinous. It might be more useful as a lubricant than a
condiment!
COMMENTS: This farflung fungus is unlikely to be mistaken for any other. The com¬
bination of rounded or wrinkled cap, viscid fruiting body with gelatinous flesh, and overall
yellowish to ochre-buff color set it apart. Greenish-tinged forms approach L. viscosa,
and the two species may intergrade.

Leotia viscosa (Chicken Lips)

FRUITING BODY with a cap and stalk. CAP 0.5-3 cm broad, round to convex to slightly
lobed or knobby; surface smooth or slightly wrinkled, viscid or slimy when moist, dark
green to olive-green; margin usually incurved toward the stalk, often lobed or wavy.
Underside sterile, usually pallid or paler. Flesh (at least the central core) gelatinous,
often translucent. STALK 2-9 cm long, 0.3-1 cm thick, equal or tapered slightly upward,

Leotia viscosa, young specimens. Note the viscid-gelatinous stalk and flesh. As they mature the caps
will grow larger. The greenish cap distinguishes this species from L. lubrica (shown in color plate).
LEOTIA 875

smooth, hollow or filled with a gel, viscid to slimy when moist; white to yellow or orange,
sometimes with minute green dots or particles, especially above. SPORES 16-28 * 4-6
microns, spindle-shaped and often slightly curved, smooth, usually septate (partitioned)
at maturity, hyaline (colorless) under the microscope.
HABITAT: Solitary, scattered, or in groups or clusters in humus or on rotten wood;
widely distributed. This is the most common Leotia in our area. It fruits in the winter and
early spring under oak and various other trees.
EDIBILITY: Harmless but gelatinous.
COMMENTS: This gelatinous Ascomycete with the green head and white to orange stalk
can hardly be confused with any other. In dry weather the surface of the cap and stalk may
not be obviously viscid, but slicing open the fruiting body will usually reveal gelatinous
tissue within. The nickname “Chicken Lips” was obviously given to it by the same person
who dubbed Tricholoma flavovirens the “Man On Horseback!” Other species: L. atro-
virens (~L. chlorocephala) is a similar but smaller eastern species with a greenish to dark
green cap and green to pale green stalk. L. lubrica (see description) is usually yellower.

BULGARIA & Allies

Small to medium-sized fungi usually growing on wood or bark of hardwoods. FRUITING BODY
top-shaped to cup-shaped to nearly round to irregularly lobed; pinkish to reddish, purplish, dark
brown, or black. Flesh gelatinous or rubbery-gelatinous. ST ALK absent or present only as a short
narrowed base. SPORES elliptical to elongated, smooth or ribbed, septate or not septate; brown
(or many of them brown) or black in Bulgaria, otherwise hyaline (colorless) under the microscope.
Asci lining upper surface of fruiting body, typically 8-spored, inoperculate, with amyloid pore in
Bulgaria, otherwise not amyloid.

THESE are gelatinous or rubbery-gelatinous, hardwood-inhabiting fungi with little or

no stalk and a top-shaped to cup-shaped to irregular fruiting body. The gelatinous texture
can lead to confusion with the jelly fungi, which bear their spores on basidia, and with
Sarcosoma and allies, which have operculate asci. Since these differences are microscopic,
the species treated here are also keyed out under those groups. One species, Bulgaria
inquinans, is described here, and three others are discussed. They are placed with Leotia
in the Leotiaceae, but lack the well-developed stalk of that genus. Because of their texture
they are not worth eating except in dire emergencies.

Key to Bulgaria & Allies

1. Fruiting body usually irregularly lobed or brainlike, often forming confluent masses; found on
beech in eastern North America . . Ascotremella faginacea (see Bulgaria inquinans, p. 876)
1. Not as above . 2
2. Fruiting body shallowly cup-shaped to earlike or seaweed-like, reddish-brown to dark brown
but not usually black (unless dried out), typically 2 cm broad or more; spores borne on basidia
which usually line the lower (but sometimes the upper) surface of fruiting body; sterile sur¬
face minutely hairy or downy. (sec Auricularia auricula, p. 675)
2. Not as above; spores borne in asci which line the upper surface of fruiting body. 3
3. Upper (fertile) surface of fruiting body dark brown to black . 4
3. Upper surface of fruiting body pinkish to reddish, reddish-brown, purplish, or tan . 5
4. Found on dead hardwoods; asci not operculate .Bulgaria inquinans, p. 876
4. Found on ground or on dead conifers (rarely hardwoods); asci operculate (i.e., with “lids”)
.(see Sarcosoma & Allies, p. 826)
5. Fruiting body cup-shaped at maturity; fertile surface reddish-brown to brown; exterior with
minute dark hairs; asci operculate (with “lids”) .(see Sarcosoma & Allies, p. 826)
5. Fruiting body usually more or less top-shaped, pinkish to reddish to purplish; asci not oper¬
culate .Ascocoryne sarcoides& Neobulgariapura (see Bulgaria inquinans, p. 876)
Bulgaria inquinans. These gelatinous specimens look like licorice drops. (If only they tasted like
them!) The three on left are being viewed from the top, the one on the upper right, from the side.

Bulgaria inquinans (Poor Man’s Licorice; Black Jelly Drops)

FRUITING BODY 1 -4 cm broad and/or high, rubbery; at first rounded to somewhat
cylindrical to shallow cup-shaped or top-shaped, the top then broadening into a flattened
or broadly convex, flabby “cap.” Fertile (upper) surface blackish and often shiny when
wet. Exterior or underside brown to blackish and roughened. Flesh gelatinous or at least
very rubbery; pliant and tough, not brittle. STALK absent or present as a narrowed base,
continuous with and colored like the exterior or underside of the “cap.” SPORES 11-14
x 6-7 microns, more or less kidney-shaped, smooth, not septate; brown or black in mass,
but only the upper four in each ascus brown under the microscope, the other four hyaline
(colorless); asci typically 8-spored, their tips blueing in iodine.
HABITAT: Solitary, scattered, in rows, or densely gregarious or clustered on dead hard¬
wood logs and branches, especially of oak; widely distributed and common. In our area
it fruits throughout the mushroom season, especially on live oak and tanoak, and can be
seen on almost any wintertime trek through the woods.
EDIBILITY: Unknown, but as the common name implies, not worth eating except in
desperation!
COMMENTS: Also known as Phaeobulgaria inquinans, the flabby, funky fruiting
bodies of this fungus look something like licorice drops, or in the concise words of Judith
Scott Mattoon, “They remind me of rubber parts that fit into things by squeezing through
and then popping back into shape—you know what I mean?” The texture is reminiscent
of India rubber, or can be gelatinous, leading to confusion with the jelly fungi. However, it
bears its spores in asci rather than on basidia, and can be told in the field by its distinctive
shape and color. Its growth on dead hardwoods and smaller size distinguish it from the

Bulgaria inquinans. These specimens were collected in dry weather and consequently are rubbery
rather than gelatinous. Note the narrowed base beneath the fertile portion.
BULGARIA 877

conifer-loving “Starving Man’s Licorice” (Sarcosoma mexicana and S. latahensis). Des¬

pite its uncanny resemblance to Sarcosoma, it is classified with the earth tongues because
of its inoperculate (lidless) asci. Other gelatinous species: Ascocoryne (-Coryne) sar-
coides is a widespread, more or less top-shaped, gelatinous, flesh-colored to dark purplish
or reddish-brown species. It is smaller (up to 1 cm broad), has septate spores, and grows
on dead wood. Neobulgaria pur a (-Ascotremella turbinata) is a flesh-colored to reddish,
top-shaped eastern species with non-septate spores. Ascotremella faginacea is a larger,
raisin-colored, gelatinous species that is lobed or brainlike and often forms continuous
masses on beech trunks in eastern North America. All of these look like jelly fungi and
are keyed out under that group.

CIBORIA & Allies

Small to minute fungi found on rotten wood, soil, or more often on plant stems, leaves, nuts, and
other vegetable matter. FRUITING BODY usually cup-shaped to disclike or cushion-shaped,
with or without a stalk. Fertile (upper) surface variously colored, usually smooth. Underside sterile,
with or without hairs. STALK if present long or short, usually very thin, sometimes arising from
a swollen “tuber.” SPORES variously shaped, usually but not always smooth and hyaline (color¬
less) under the microscope. Asci lining the upper surface of the fruiting body, usually 8-spored,
inoperculate, with a pore at the tip which may or may not be amyloid.

THE above synopsis covers a vast group of minute but cute cuplike and disclike fungi that
are seldom collected except by professional specialists. They differ from the many cup
fungi in the Pezizales in having inoperculate (lidless) asci. Some, such as Ciboria, have
long stalks attached to the substrate. Others, such as Sclerotinia, Whetzelinia, and Myrio-
sclerotinia, have long stalks that arise from a tuberlike mass of tissue (sclerotium). Still
others, like Dasyscyphus, have shorter, sometimes hairy stalks. Finally, there are many
(e.g., Mollisia) that have no stalk at all. A large number of these fungi are parasitic on
plant parts, but some are saprophytes. Without exception they are too small to interest the
average mushroom hunter and are certainly too small to eat. The two genera described
here, Ciboria and Chlorociboria, belong to different families (the Sclerotiniaceae and
Leotiaceae, respectively). Several other related families are not treated here.

Key to Ciboria & Allies

1. Fruiting body blue to green or pallid with a blue or greenish tinge; growing on wood .
.Chlorociboria aeruginascens & others, p. 878
1. Not as above . 2
2. Fruiting body minute, bright yellow to orange-yellow; found on rotten wood, usually in swarms;
widely distributed .Bisporella citrina
2. Not as above .. 3
3. Stalk arising from a swollen “tuber” (sclerotium) Sclerotinia, Whetzelinia, & Myriosclerotinia
3. Not as above .4
4. Fruiting body yellow-brown to brown, with a stalk; growing on willow and alder catkins ....
.Ciboria amentacea, below
4. Not as above .Ciboria, Dasyscyphus, Mollisia, & many others

Ciboria amentacea (Catkin Cup)

FRUITING BODY with a stalk and cuplike cap. “Cap”0.5-1.2 cm broad, at first shallowly
cup-shaped, expanding to nearly flat in age. Fertile (upper or inner) surface light brown to
yellow-brown, smooth. Exterior of cup similar in color, also smooth. STALK 1-5 cm
long, 1 -2 mm thick, light brown to yellow-brown, equal, smooth, often curved, not arising
from a sclerotium. SPORES 7.5-13 * 4-6 microns, elliptical, smooth.
878 HELOTIALES

HABITAT: Solitary or in small groups on old, fallen alder and willow catkins; widely
distributed but seldom collected, usually fruiting in the spring.
EDIBILITY: Who knows?
COMMENTS: The growth on alder or willow catkins rescues this little brown cup fungus
from the anonymity it so richly deserves.

Chlorociboria aeruginascens (Blue Stain)

FRUITING BODY 3-7(10) mm broad, at first cup-shaped, then becoming flat or disclike
or with a slightly elevated margin. Fertile (upper) surface bright to pale blue-green or
turquoise, sometimes with a yellowish or orange-yellow tint developing in age; smooth or
slightly wrinkled. Exterior (underside) similarly colored (but not yellowish). Flesh thin,
also bluish-green. STALK usually present as a short, narrowed, typically off-center base;
up to 3 (6) mm long and 1-2 mm thick, same color as rest of fruiting body. SPORES 6-10 x
1.5-2 microns, spindle-shaped or elongated, smooth, with an oil droplet at each end.
HABITAT: Gregarious (several often arising from a common base) on dead or barkless
wood (usually oak); widely distributed and quite common, but easily overlooked because
of its diminutive dimensions. In our area it fruits mainly in the winter and early spring.
EDIBILITY: Indisputably inconsequential.
COMMENTS: This petite Ascomycete merits mention in this book because of its unusual
color. C. aeruginosa is a very similar, widely distributed species that is smaller (less than
5 mm broad), has a shorter, more or less central stalk, and orange-yellow flesh. Both
species can be detected when they are not fruiting because their mycelium stains the host
blue-green. The stained wood was once used in the manufacture of inlaid wooden objects
known as “Tunbridge Ware.” Both species have also been placed in the genus Chloro-
splenium.

Flask F ungi
spores

PYRENOMYCETES
THE Pyrenomycetes differ fundamentally from the Discomycetes (morels, cup fungi,
earth tongues, etc.) because they bear their asci in flask-shaped “nests” called perithecia.
The perithecia are usually imbedded in the fruiting body, but their necks or mouths often
protrude like small pimples.
The flask fungi are a varied lot, but only a few of them are conspicuous enough to be
considered in this tome. The most common types, Xylaria and Daldinia, are tough,
usually black, and grow on wood. Another distinctive group, Cordyceps, is parasitic on
insects and truffles. Still another, Hypomyces, engulfs other mushrooms in a pimpled or
powdery weft of tissue. The flask fungi treated here belong to a single order (Sphaeriales)
within the Pyrenomycetes. They are unpalatable except for one species of Hypomyces.
They fruit in moist weather, but the tougher types persist year-round.

Key to the Pyrenomycetes

1. Growing on wood (but wood sometimes buried) . 2
1. Growing on insects (including pupae or larvae), spiders, truffles, or other mushrooms (but the
host sometimes buried inside rotten wood!) . 3
PYRENOMYCETES 879

2. Fruiting body whitish to yellowish to pale ochre; rare . Podostroma, below

2. Fruiting body gray to dark brown or black, but sometimes covered with a white powder;
common .Xylaria & Daldinia, p.885
3. Fruiting body clublike, threadlike, or with a cap and stalk; growing on insects, spiders, or
certain truffles .Cordyceps, below
3. Fruiting body a pimpled or powdery layer of tissue that covers or partially covers its host;
growing on other mushrooms .Hypomyces, p. 882

PODOSTROMA
THIS rare, wood-inhabiting genus is represented by a single boring species in our area,
described below.

Podostroma alutaceum
FRUITING BODY 1-5 cm tall and 0.5-1 cm thick, cylindrical to club-shaped, without a
well-defined cap. Surface dry, minutely roughened by the slightly protruding perithecia
(flasklike nests of asci), whitish to yellowish to pale ochre, usually paler (white) at the
base. SPORES elongated, hyaline (colorless) under the microscope, finely warted and
septate (with one partition), breaking up into one-celled, round to elliptical segments
averaging 4-4.5 * 3^1 microns. Asci 8-spored, but each spore breaking in two to make 16.
HABITAT: Solitary or in small groups on rotting wood; widely distributed but rare.
I have found it only once in our area, on dead oak in the late winter.
EDIBILITY: Who cares?
COMMENTS: This forgettable clublike fungus can be recognized by its yellowish color,
growth on wood, and presence of perithecia (flasklike “nests” of asci) on the upper
portion of the fruiting body. It is most likely to be mistaken for a fairy club (Clavaria or
Clavulinopsis), but the above-mentioned features distinguish it.

CORDYCEPS
Small fungi found on insects (pupae, larvae, and adults), spiders, and certain truffles. FRUITING
BODY threadlike to club-shaped or with a differentiated “head" and stalk; often brightly colored
but sometimes dull or dark; surface often roughened or minutely pimpled by projecting perithecia.
STALK present, usually slender or very thin, arising from the host insect or truffle (which is often
buried). SPORES threadlike, typically hyaline (colorless) under the microscope and smooth,
multiseptate, but usually breaking up quickly into one-celled, barrel-shaped or elongated segments.
Asci borne in perithecia (flasklike structures) imbedded in or projecting from the“head”(if present)
or upper portion of the fruiting body.

THESE small but fascinating clublike fungi are obligate parasites of insects, spiders, and
certain truffles. As such they are easy to recognize, providing you dig them up carefully
so they can be traced to their host, which is often buried in humus, soil, or rotten wood.
When the host is overlooked or left behind, many species of Cordyceps can still be distin¬
guished from other clublike fungi by their minutely roughened or pimpled fertile surface.
Cordyceps is a fairly large genus and only a few species can be treated here. Most of them
parasitize insect larvae, pupae, and adults. The mycelium develops inside the insect,
killing it and devouring it. After the insect is completely mummified and emptied of
nutrients, the mycelium fruits and then dies. As the insect is the sole source of food for the
fungus, the size of the fruiting body is often dependent on the size of the host. Since insects
880 PYRENOMYCETES

are more abundant in warm, humid weather, it’s not surprising that Cordyceps is particu¬
larly prominent and diverse in eastern N orth America (where there are summer rains) and
the tropics. In California and the Pacific Northwest, where the summers are drier, insect¬
eating species are comparatively rare, but those that parasitize truffles are more common.
It should be emphasized, however, that even the “common” species of Cordyceps are rare
in relation to other mushrooms. Most mycologists consider one or two fruiting bodies of
Cordyceps a real find!
Cordyceps are worthless as food because of their small size and infrequent occurrence.
Their unique diet, however, makes them a fascinating group to study. Perhaps some day
we will find a practical use for them in the control of certain insect pests. Three species of
Cordyceps are described here, and several others are keyed out. Also worth mentioning
is the closely related genus, Claviceps, which parasitizes plants rather than insects or
truffles. The most potent hallucinogenic compound known, LSD, was derived from
Claviceps purpurea, better known as wheat ergot..

Key to Cordyceps
1. Growing on truffles (species of Elaphomyces) . 2
1. Growing on insects (adults, pupae, or larvae) or spiders. 3
2. Fruiting body with a cap and stalk .C. capitata & others, below
2. Fruiting body clublike (lacking a distinct cap), usually with yellow mycelial threads at base
or permeating the host .C. ophioglossoides (see C. capitata, below)
3. Fruiting body with a cap or “head” which is usually clearly delimited from the stalk or sterile
portion of fruiting body . 4
3. Fruiting body lacking a differentiated “head” or cap, but often thicker toward the top .... 7
4. Growing on beetles, moths, or butterflies (or their larvae or pupae) . 5
4. Growing on ants or wasps . 6
5. Stalk typically yellow; cap ochre to mahogany .... C. gracilis (see C. myrmecophila, p. 881)
5. Not as above; fruiting body brownish or tinged vinaceous; “head” warty .
. C. entomorrhiza (see C. myrmecophila, p. 881)
6. Growing on ants . C. myrmecophila, p. 881
6. Growing on wasps .C. sphecocephala (see C. myrmecophila, p. 881)
7. Fruiting body threadlike (less than 2 mm thick) .
. C. unilateralis, C. clavulata, & others (see C. militaris, p. 882)
7. Fruiting body not threadlike . 8
8. Fruiting body brown to purple-brown or blackish .C. ravenelii (see C. militaris, p. 882)
8. Fruiting body white to yellow, orange, or orange-red . 9
9. Growing on beetles (usually the larvae or pupae); fruiting body with sterile tip.
.C. melolanthae (see C. militaris, p. 882)
9. Growing on butterflies and moths (usually larvae or pupae); tip not sterile . 10
10. Fruiting body orange-buff to orange to orange-red .C. militaris, p. 882
10. Fruiting body whitish to yellow . C. washingtonensis (see C. militaris, p. 882)

Cordyceps capitata (Truffle Eater)

FRUITING BODY arising from certain underground truffles (Elaphomyces), 2-8 (12)
cm tall, with a well-defined cap or “head” and stalk. Fertile “head” 0.5-2 cm broad and
high, nearly round to convex or slightly conical; surface dark reddish-brown to brown,
dark olive-brown, or even blackish, roughened or minutely pimpled by protruding peri-
thecia (flasklike nests of asci). Flesh white. STALK 1.5-8 cm long, (0.2) 0.4-1.5 cm thick,
more or less equal, sometimes slightly flattened, often bent or curved, occasionally forked
(with two “heads”), rather tough; surface usually fibrillose of fibrillose-scaly, yellow to
yellow-ochre to yellow-olive, sometimes darker (olive to olive-black) in age; base often
CORDYCEPS 881

whitish. SPORES threadlike, hyaline (colorless) and smooth under the microscope,
usually breaking up into one-celled segments averaging (8) 12-27 (32) * 1.5-3 microns.
HABITAT: Solitary, tufted, or gregarious on ground, but arising from underground deer
truffles (Elaphomyces species); widely distributed and one of the more common members
of the genus. Scattered fruiting bodies are the norm, but sometimes it fruits prolifically.
I have not found it in our area, but it may well occur {Elaphomyces certainly does). In the
mixed coastal forests of northern California it can be found in the fall and winter.
EDIBILITY: Possibly worth trying since it is larger than most species of Cordyceps, but
I can find no information on it.
COMMENTS: This is one of several Cordyceps species that grow only on Elaphomyces.
The latter can occur several inches deep in the soil, but specimens close to the surface are
more apt to be parasitized. The reddish-brown or darker cap which is sharply differen¬
tiated from the yellow to olive stalk are the principal fieldmarks. C. canadensis is a very
similar truffle-eater with much larger spore segments; it is known from eastern North
America and Europe. C. ophioglossoides is another species that parasitizes Elapho¬
myces, but it has a clublike fruiting body that lacks a sharply defined “head.” The club is
sometimes yellow when very young but soon becomes reddish-brown to olive-brown to
nearly black except for a yellow base and yellow mycelial threads thatextend into the host.
It is said to be the most common Cordyceps in eastern North America, but is rather rare
in California.

Cordyceps myrmecophila (Ant Fungus; Ant Eater)

FRUITING BODY arising from an ant (often buried), 0.8-5 (10) cm tall, with a thin stalk
and small “head.” Fertile “head” 2-8 mm broad, usually oval; surface ochre to ochraceous-
salmon, sometimes with short longitudinal ridges or furrows in dry weather, minutely
pimpled from the slightly projecting perithecia (flasklike nests of asci). STALK 0.8-9.5
cm long, 0.5-1 (2) mm thick, very thin and more or less equal, colored like the “head” or
often paler (pale yellow, sometimes shading to white near base, or entirely white if not
exposed to light). SPORES threadlike and multiseptate, smooth, hyaline (colorless)
under the microscope, breaking into one-celled segments averaging 8-10 x 1.5 microns.
HABITAT: Scattered to gregarious on the mummified, often buried carcasses of ants
(usually one per ant); widely distributed, fruiting in damp weather, usually around ant
nests in the woods. Although rare, it is sometimes prolific when it fruits. It has been found
in British Columbia, Washington, and Oregon (as well as Europe, China, and Brazil), and
may well occur in California; there is certainly no shortage of potential hosts!
EDIBILITY: Unknown.
COMMENTS: This is one of several Cordyceps species that are capitate (i.e., that have
a differentiated “head”) and parasitize insects. The ochre to yellow color and growth on
ants distinguish it. As in other species, the length of the stalk depends largely on whether
or not the host is buried, and if so, how deep. Other capitate species include: C. spheco-
cephala, a very similar southeastern species that grows on wasps and has a very thin,
creamy to yellow or yellow-brown fruiting body; C. entomorrhiza, brownish or vinaceous-
tinged with a very warty “head,” growing on beetle larvae in the Pacific Northwest, rare;
and C. gracilis, growing on larvae of beetles, moths, and butterflies in eastern North
America, with a yellow stalk and ochre to mahogany-colored “head,” also rare. For
capitate species that grow on truffles, see C. capitata, and for the clublike (non-capitate)
types that grow on insects, see C. militaris.
882 PYRENOMYCETES

Cordyceps militaris (Caterpillar Fungus)

FRUITING BODY arising from buried moth and butterfly larvae or pupae, 2-8 cm tall,
cylindrical to spindle- or club-shaped (i.e., with a slightly swollen upper fertile region, but
lacking a well-defined “head”); often with a longitudinal furrow. Upper (fertile) portion
of club 2-6 mm broad, orange to orange-buff to orange-red, finely roughened or pimpled
by the slightly protruding perithecia(flasklike nests of asci). STALK (sterile lower region)
smooth, usually paler, often curved or wavy. SPORES threadlike and multiseptate,
smooth, hyaline (colorless) under the microscope, breaking up into one-celled, barrel¬
shaped segments averaging 2-6 * 1-1.5 microns.
HABITAT: Solitary to gregarious or clustered on buried pupae or less commonly larvae
(caterpillars) of moths and butterflies; widely distributed. It is one of the more common
species in the genus, but is rare in California. In eastern North America I have found it
several times in the summer and fall.
EDIBILITY: Unknown.
COMMENTS: The orange to orange-buff, club-shaped fruiting body that arises from
mummified pupae or larvae is most distinctive. It might be confused with other clublike
fungi (e.g., Clavulinopsis, Microglossum) if it is dug up carelessly or the host is overlooked,
but the pimpled upper portion (see photo on p. 883) will still identify it. Other species that
parasitize insects and do not have a clearly defined “head” or cap include: C. wash-
ingtonensis of the Pacific Northwest, also growing on moth and butterfly pupae or larvae,
and very similar, but with a whitish to yellow fruiting body; C. melolanthae, growing on
beetles (usually buried grubs), sometimes in huge numbers, in eastern North America,
often larger than C. militaris, with a whitish to yellow to orange fruiting body that has a
sterile tip; C. ravenelii, a rare eastern species with a brown to chocolate-brown, purple-
brown, or blackish, clublike fruiting body, also growing on beetle grubs; C. unilateralis, a
minute (up to 3 cm tall and less than 1 mm thick) brown eastern and southern species that
feeds on ants, bees, and wasps; and C. clavulata, growing on scale insects. There are many
other species, particularly in eastern North America and the tropics; some can only be
identified with a microscope. For species that parasitize insects and have a well-defined
“head,” see C. myrmecophila.

HYPOMYCES
U biquitous fungi that parasitize other mushrooms, partially or completely covering them in a layer
of pimpled or powdery tissue. FRUITING BODY taking on the shape of its host, but disfiguring
it; composed of a layer of tissue in which numerous perithecia are imbedded (but asexual stages
lack perithecia). SPORES (sexual) elongated, smooth or warted, often 1-septate, hyaline (color¬
less) under the microscope. Asexual spores of various shapes and sizes also produced by some
species. Asci borne in perithecia (flasklike structures).

THIS distinctive genus is parasitic on other mushrooms (mostly agarics and boletes),
engulfing them in a pimpled or powdery layer of tissue. The actual fruiting bodies (peri¬
thecia) of Hypomyces are not large enough to qualify as mushrooms, but are given shape
and substance by the mushrooms they grow on. Hypomyces can be found wherever
suitable hosts occur, sometimes in epidemic proportions. Apparently the mycelium lives
with or on the mycelium of its host and fruits at the same time. H. lactifluorum, sometimes
called the “Lobster Mushroom” because of its bright red to orange color, is eaten by many
people. Other species of Hypomyces are unpalatable and potentially dangerous, parti¬
cularly if the host is a poisonous mushroom disfigured beyond recognition. Two common
species of Hypomyces are described here and several others are keyed out.
Left: Cordyceps militaris (see p. 882). Note the pimpled surface caused by protruding perithecia, and
the insect on which it is growing (partly visible on left). (Alan Bessette) Right: Hypomyces chryso-
spermum{ see below) engulfs boletes in a white to bright yellow mass of tissue.

Key to Hypomyces
1. Growing on boletes, covering them in a white to bright yellow mass of tissue .
.H. chrysospermum, below
1. Not as above . 2
2. Growing on fruiting bodies of Russula and Lactarius, covering them in a minutely pimpled
layer of bright orange to red or magenta tissue .H. lactifluorum, p. 884
2. Not as above . 3
3. Growing on gilled mushrooms (mostly Amanita, Lactarius, and Russula) . 4
3. Growing on elfin saddles or coral fungi . 5
4. Growing on fruiting bodies of Amanita, covering them in a white to flesh-colored or pinkish
layer of tissue .H. hyalinus(see H. lactifluorum, p. 884)
4. Growing mostly on fruiting bodies of Lactarius and Russula (usually on the gills and upper
stalk), covering them in yellow to greenish tissue H. luteovirens (see H. lactifluorum, p. 884)
5. Growing on coral fungi (e.g., Ramaria) ...//. transformans(see H. chrysospermum, below)
5. Growing on elfin saddles (e.g., Helvella) ...//. cervinigenus(see H. chrysospermum, below)

Hypomyces chrysospermum (Bolete Eater)

FRUITING BODY beginning as a white moldy-looking growth that attacks and quickly
engulfs boletes, then turns bright yellow and powdery, then finally becomes reddish-
brown (but this last stage rarely seen) and becomes pimpled. Flesh of the host often soft
or mushy. SPORES in white stage 10-30 x 5-12 microns, elliptical, smooth; in yellow
stage, 10-25 microns, round, thick-walled, and warted; in final (sexual) stage, 25-30 x 5-6
microns, spindle-shaped, hyaline (colorless) under the microscope and usually 1-septate
(partitioned). Perithecia only present in final stage.
HABITAT: Solitary, scattered, or gregarious on boletes; widely distributed and very
common in our area whenever boletes are out; also reported on Paxillus and Rhizopogort.
EDIBILITY: Not edible, possibly poisonous. It is often associated with bacterial decay.
COMMENTS: This is the white to bright yellow fungus you see so often on boletes, and
which you’ve probably cursed a hundred times for depriving you of your meal. The spores
produced in the white and yellow stages are asexual; only those in the final stage are per¬
fect spores (sexually produced inside asci). This final stage is seldom seen, however, be¬
cause it occurs only after the host is decayed beyond recognition and is very unpleasant

883
Hypomyces hyalinus (specimens on right) disfigures various species of Amanita, in this castA.rubes-
cens (shown on left). See comments under H. lactifluorum for more details.

to handle. The different stages of the fungus have been given different names, e.g., Sepe-
donium chrysospermum was originally applied to the yellow stage. Other species: In our
area, H. cervinigenus commonly attacks the black fluted elfin saddle (Helvetia lacunosa),
covering it with white or pinkish tissue. H. transformans performs a similar transfor¬
mation on several species of coral fungi (particularly Ramaria). Neither of these should be
eaten. For species that attack gilled mushrooms, see H. lactifluorum.

Hypomyces lactifluorum (Lobster Mushroom) Color Plate 216

FRUITING BODY growing on and engulfing gilled mushrooms (species of Russula and
Lactarius) in a layer of roughened or pimpled, bright orange to orange-red to purple-red
or occasionally yellow-orange tissue which is firm to the touch. Overall shape of the host
mushroom and parasite often like an inverted pyramid. Flesh (of the host) crisp, white.
SPORES 30-50 x 4.5-8 microns, spindle-shaped or shaped like caraway seeds, hyaline
(colorless) under the microscope, septate (with one partition), warted. Perithecia im¬
bedded in the tissue that covers the host, but protruding as small pimples.
HABIT AT: S olitary, scattered, or gregarious in woods, often partially buried in the duff,
usually on the fruiting bodies of Lactarius and Russula (especially the large white species
like R. brevipes)', widely distributed. It is common in some regions, but rare in our area.
The largest fruitings I’ve seen were under ponderosa pine in the Southwest.
EDIBILITY: Rated highly by many people and sold in markets in Mexico. There is no
absolute assurance that the host species is edible, but I can find no mention of poisonings
by this species. Perhaps it only attacks edible species! Material I sampled was fairly good.
COMMENTS: This fungus is best recognized by its bright orange to reddish color and
minutely pimpled surface. The gills of the host are often reduced to blunt, chanterelle¬
like ridges, but the pimpled appearance caused by the numerous perithecia and crisper
texture distinguish it. The bright fluorescent color of H. lactifluorum make it the most
spectacular member of its clan. H. luteovirens is a somewhat similar yellow-green to
greenish species that covers the gills and upper stalk of various Lactarius and Russula
species. It is more common in our area than L. lactifluorum, but not nearly as conspicuous.
H. hyalinus is a white to pinkish or flesh-colored species that attacks species of Amanita
(particularly A. rubescens), turning them into pimpled or warty upright clubs (see photo
above). It should not be eaten because its host might be deadly poisonous! It is common
in eastern North America, but I have yet to find it on the west coast.

884
 885

XYLARIA& DALDINIA
Very tough to hard or charcoal-like, wood-inhabiting fungi. FRUITING BODY erect and clublike
or branched in Xylaria, or stalkless and hemispherical in Daldinia; usually black when mature,
but sometimes covered with a white or grayish or brown powdery coating of asexual spores (coni-
dia). Flesh tough or charcoal-like, usually white in Xylaria, concentrically zoned in Daldinia.
STALK present in Xylaria, absent in Daldinia. SPORES dark brown to black, usually spindle-
shaped or elliptical, smooth; asexual spores (conidia) hyaline (colorless) under the microscope,
smooth. Asci borne in flasklike structures (perithecia) imbedded in the fruiting body (usually
upper portion).

THESE tough, mostly black, wood-inhabiting fungi are very distinctive. The erect, club¬
like forms (Xylaria) might be confused with the black earth tongues (Geoglossum and
Trichoglossum), but are much tougher or harder and have paler flesh. Daldinia, on the
other hand, has a sessile (stalkless) fruiting body. It might be mistaken for a charred
polypore or crust fungus, but its concentrically zoned interior, pimpled exterior, and
brittle, charcoal-like consistency are distinctive. Both Xylaria and Daldinia
frequently produce spores asexually. These asexual spores (conidia) take the form of a
white to gray or brown powder that coats the surface of the young fruiting body. The
powder may disguise the black undersurface, but is easily rubbed or licked off. In mature
specimens, numerous asci-containing flasks (perithecia) can be seen if the fruiting body
is sliced open.
Xylaria and Daldinia are very common, but much too tough to be edible. Identification
of species is based largely on microscopic characteristics, but the genera are easily learned.
Two widespread Xylarias and one Daldinia are described here.

Key to Xylaria & Daldinia

1. Fruiting body round to hemispherical or lumpy, with little or no stalk (not growing erect);
flesh charcoal-like (hard but usually brittle).Daldinia grandis & others, p. 887
1. Not as above; fruiting body erect, clublike (unbranched) or antlerlike (branched) . 2
2. Fruiting body slender, often branched and/or covered with a whitish powder (at least over
upper portion) .Xylaria hypoxylon, below
2. Not as above . 3
3. Fruiting body very tough or hard, up to 3 cm thick; flesh inside usually white or pallid;
surface often minutely warted or cracked . .'.Xylaria polymorpha & others, p. 886
3. Not with above features .(see Geoglossum & Trichoglossum, p. 866)

Xylaria hypoxylon (Candlesnuff Fungus)

FRUITING BODY 2 -8 cm high, very tough, erect, slender, cylindrical or narrowly club¬
like when young but usually becoming antlerlike (branched sparsely or forked at the tip)
in age. Upper portion or tip (or occasionally entire surface) white and powdery when
young, eventually becoming black and minutely roughened (use hand lens!). Flesh very
tough, white or pallid. STALK (lower sterile portion of fruiting body) thin, usually 1-3 (5)
mm thick, black, minutely hairy, very tough or wiry. SPORES (sexual) 10-14 * 4-6
microns, bean-shaped, smooth, black in mass but brown under the microscope; asexual
spores (conidia) smooth and elliptical or elongated, white in mass but hyaline (colorless)
under the microscope. Perithecia imbedded in the upper part of fully mature fruiting body.
HABITAT: Scattered to densely gregarious or clustered on rotting logs, stumps, buried
sticks, etc.; very widely distributed and common. In our area it occurs year-round in many
habitats, but especially on oak and tanoak in the fall and winter.
EDIBILITY: Much too tough to be of value.
Left: Xylaria hypoxylon. Antlerlike specimens such as these are typical, but unbranched ones also
occur. At least the upper portion of black fruiting body is usually covered with white spore powder
until fully mature. Right: “Dead Man’s Fingers,” Xylariapolymorpha. Note thick fruiting body with
blunt tip and sterile base or stalk. These two specimens are black, but the picture is overexposed.

COMMENTS: Also known as Xylosphaera hypoxylon, the candlesnuff fungus is easily

told by its very tough, slender, antlerlike fruiting body that is black below and dusted with
white powder above. The powdered appearance of young specimens is caused by masses
of asexual spores (conidia) that form directly on hyphae instead of in asci. Later on, the
asci form inside “flasks” (perithecia) at the top of the fruiting body. There are many
Xylarias that more or less resemble this species (e.g., X. cornu-damae), but they are best
differentiated microscopically. Some are short and cylindrical, others are branched or
clustered. All grow on wood (sometimes buried) and are very tough. For thicker club-
shaped species, see X. polymorpha.

Xylaria polymorpha (Dead Man’s Fingers)

FRUITING BODY 2-8 cm tall, 0.5-3 cm thick, very tough and hard or carbonaceous;
erect, club- or finger-shaped to somewhat irregular or twisted, the tip usually blunt or
rounded and occasionally lobed. Outer surface hard and crustlike, usually wrinkled,
roughened, and/ or cracked, black when mature but often covered with a whitish to grayish
or brownish powder when very young. Flesh (interior) hard or corky, white or pallid.
ST ALK present as a short sterile base, usually well-defined and narrower than fertile part;
black. SPORES (sexual) 20-32 * 5-10 (12) microns, spindle-shaped, smooth, dark brown
to black; asexual spores (conidia) when present smaller, elongated or elliptical, smooth,
hyaline under the microscope. Perithecia imbedded in upper portion of fruiting body.

HABITAT: In groups or clusters on hardwood stumps, logs, etc., but often appearing
terrestrial if the wood is buried; widely distributed and common, but apparently absent
or very rare in our area. In eastern North America, where it favors beech and maple, the
fruiting bodies usually appear in the spring and mature (blacken) by the summer. They
last for months without decaying.
EDIBILITY: Much too tough and rough to be edible.
COMMENTS: Also known &s Xylosphaera polymorpha, this fungus is easily recognized
by its hard, dark fingerlike fruiting bodies. They are much thicker than those of X. hy¬
poxylon, Geoglossum, and Trichoglossum, and the white or pallid interior is also dis¬
tinctive. There are several very similar temperate and tropical species that are collectively
called “Dead Man’s Fingers.” These are differentiated primarily on microscopic charac¬
teristics. One, X. longipes, is similar but consistently slimmer (0.3-1 cm thick) than X.
polymorpha. It occurs across the continent, usually on hardwoods.

886
Daldinia grandis. Note the concentrically zoned interior (shown in sectioned specimen at upper right)
and minutely pimpled exterior. Charcoal-like consistency and growth on wood are also distinctive.

Daldinia grandis (Crampballs; Carbon Balls; King Alfred’s Cakes)

FRUITING BODY very tough and woody or charcoal-like, 1-6 cm broad or sometimes
larger, hemispherical to nearly round to somewhat lumpy or irregular, stalkless. Exterior
black (or sometimes dark brown when young), roughened or pimpled by the perithecia,
often cracked in age. Flesh (interior) brown to grayish-black, somewhat lustrous, with
lighter and darker concentric zones; very brittle and charcoal-like. STALK absent.
SPORES (sexual) 14-17 (27) * 6.5-11 microns, elliptical or elongated, smooth, dark brown
to black; asexual spores (conidia) minute, smooth, and hyaline (colorless) when present.
HABIT AT: Scattered to gregarious or in masses on dead logs, branches, or bark of hard¬
woods; widely distributed. It is abundant year-round in our area, especially on oak.

EDIBILITY: Unequivocally inedible—but perhaps useful as a substitute for charcoal!

COMMENTS: The distinctive charcoal-like fruiting bodies of this fungus can be found
on almost any walk through the woods. The pimpled surface of mature specimens (see
photo) is caused by the protruding tips of the perithecia (flasks of asci). As in Xylaria,
younger specimens are sometimes coated with pale, asexual spores (conidia). The nick¬
name “Crampballs,” incidentally, can be attributed to the old folk belief that carrying
one around under your armpits would cure cramps! Other species: D. concentrica is the
common crampball of eastern North America. It is very similar to D. grandis, but has
slightly smaller spores and is more apt to be dark brown to bronze-black when young
(and black in age). D. vernicosa is also similar, but usually has a narrowed base beneath
the fertile portion and an interior zoned with dark brown and white or gray. Other species
can only be differentiated microscopically.

“King Alfred’s Cakes,” Daldinia grandis, sharing a juicy log with Phellinus gilvus (see p. 582).
888

MUSHROOM COOKERY
The abundance boneless
Without husk or scale or thorn,
Granting us this festival of all-embracing freshness

Pablo Neruda’s tribute to the tomato is also a tribute to immediacy and

vitality, that incomparable freshness that distinguishes a homegrown tomato or
cucumber or wild mushroom from its flavorless, mass-produced counterpart.
The challenge in cooking wild mushrooms is to maximize their freshness and
earthy essence while highlighting their individuality. After all, they are not one
vegetable, but many—a pleasant surprise to people who are conditioned to
mushrooms that smell and taste “mushroomy.” The major constraint is that you
must make do with what you have. Obviously, you can’t cook boletus broth when
you have a basketful of blewits, but you can make blewit burgers, or blewit
biscuits, or a three-bean blewit salad.
The most important thing to remember is that you can’t expect wild mush¬
rooms to be special unless you take the time to make them special. They are
ephemeral, temperamental, delicate. It is a relatively simple task to render the
most marvelous mushroom tasteless. Likewise, many “mediocre” mushrooms
are delightful when cooked with care and imagination. If you make a concerted
effort to seek out, gather, identify, and eat wild mushrooms, it only makes sense
to do them justice in the kitchen. Don’t just throw them into the pot with a bunch
of other vegetables—unless, of course, you want them to taste like a bunch of
other vegetables. Different mushrooms call for different treatment; only then
will they respond with their full measure of flavor.
With each type you will go through a period of discovery and experimentation
—succulent successes and unforgettable failures—followed by a process of
adjustment and subtle refinement. Each kind of mushroom will gradually acquire
its own culinary identity and cease to be a mushroom except in the botanical
sense that broccoli is a plant. After all, when you’re having broccoli, you d on’t say,
“We’re having steamed plants for dinner.” Similarly, it will no longer be “mush¬
rooms” for dinner, but “chanterelles,” or “poor people’s truffle,” or in the case
of the cultivated mushroom, “Agaricus bisporus” (to be pronounced with a
subtle insinuation of distaste).
Strive for a marriage (but not a compromise!) between elegance and simplicity.
Successful mushroom cookery doesn’t require exotic foods, or a bottomless bank
account, or idle afternoons, or a degree in gastronomical mechanics. It does
require patience, sensitivity, enthusiasm, and imagination. There are no rigorous
rules, but some basic do’s and don’ts are summarized below.

HELPFUL HINTS

1. Don’t eat a mushroom unless you’re absolutely sure it’s edible. In other words:
“When in doubt, throw it out.”
2. You wouldn’t eat a rotten egg, so don’t eat a rotten mushroom. Food poisoning is
a frequent cause of so-called “mushroom poisoning.”
3. You wouldn’t eat five pounds of asparagus, so don’t eat five pounds of mushrooms,
no matter how delicious they are. Overindulgence (COLOR PLATE 217) is another
common cause of so-called “mushroom poisoning,” particularly for those who
“What was desire in the hills becomes fulfillment in the kitchen,” says Angelo Pellegrini in The Savory
Wild Mushroom. At left is fresh Agaricus campestris. At right, a delicious cream sauce with white
onions and sliced A. campestris.

don’t eat mushrooms regularly. On the other hand, don’t be stingy—most wild
mushrooms cook down more than the commercial variety, so use them generously.
4. When trying a species for the first time, eat only a small amount. Then wait for a few
hours to see if you have an adverse reaction to it. Just as some people are allergic to
eggs or chocolate or scallops or strawberries, some are adversely affected by certain
kinds of mushrooms. Species to which many people are “allergic” (e.g., Laetiporus
sulphureus, Lepiota species) should not be served to large groups.
5. In the event of an “allergy,” you’ll want to pinpoint the culprit, so don’t mix two or
more species together unless you’ve eaten them before.
6. As a rule, maggot-riddled mushrooms (see photo at bottom of p. 277) should not be
eaten, especially when uninfested specimens are available. However, in the case of
certain choice species (e.g., Agaricus augustus), you may wish to remove the maggots
with a knife (if there are only a few) or even leave them in (they’re just a little extra
protein). Use your own judgment on this matter.
7. Use as little water as possible when cleaning mushrooms. They absorb it so readily
that it dilutes their flavor and causes them to cook up slimy. On the other hand,
there’s nothing worse than gritty wild mushrooms (unless it’s gritty domesticated
mushrooms), so don’t hesitate to use water if nothing else works. If you wash them,
drain them on a paper towel before cooking. The best place to clean mushrooms is in
the field (providing you already know their identity). Trim away all dirt with a knife
or small brush, and don’t mix dirty specimens with clean ones. Also, check for
maggots and remove any that are present so they won’t multiply and spread (even
though maggots are the larvae of gnats or flies, they are able to reproduce partheno-
genetically).
8. Use mushrooms as soon as possible after picking them. Prolonged refrigeration
deprives all vegetables (including mushrooms) of their freshness and flavor. Species
of Coprinus should be eaten the day they’re picked, or they will digest themselves.
There’s an old saying to the effect that you should boil the water before harvesting
the corn. Well, it’s not a bad idea to melt the butter before picking the shaggy manes!

889
Left: One day’s catch: Boletus edulis, Amanita calyptrata, Ramaria spp., and others. Right: Enjoying
the day’s catch (in this case, shaggy manes fried in egg batter and bread crumbs).

9. During periods of heavy rainfall, most mushrooms will be waterlogged. These are
apt to cook up insipid and slimy, but can be sliced and dried for later use (thereby
concentrating their flavor).
10. Don’t steam or pressure-cook mushrooms. You want to get rid of excess moisture,
not add to it. Pressure-cooked mushrooms bear an uncanny resemblance to slugs.

11. Don’t drown mushrooms in spices, butter, salt, garlic, or olive oil. All of these com¬
plement mushrooms nicely when used in moderation. Mushrooms and onions, for
instance, are practically made for each other, but the mushrooms must always
dominate in quantity because their flavor is more delicate.
12. If you don’t like a “choice” mushroom, give it a second and third chance. After all,
a lot depends on how you cook it and in what condition it was found. The environ¬
ment can also have an influence. For instance, blewits that grow under cypress are
often bitter-tasting. No one agrees on which kinds are best, but a species does not
acquire a widespread reputation unless it has something special to offer. On the other
hand, some relatively unknown mushrooms (e.g., Chroogomphus) are quite good.
Improvise!

PRESERVING MUSHROOMS FOR CONSUMPTION

Most mushrooms have fickle fruiting habits, appearing in large numbers for one or two
weeks, then disappearing for the rest of the year. To take advantage of their fleeting
abundance, you have to harvest them while you can and then preserve them for later use.

DRYING

Drying mushrooms is the easiest and most satisfactory way to preserve them. Fleshy
types like Boletus must be cut in thin slices; smaller species like Marasmius oreades can be
dried whole. Don’t use an oven. Circulation is more important than heat—you want
moisture to be carried away. Spread out the mushrooms on screens and use a light bulb
or hot plate as a heat source (unless it’s arid enough to sun-dry them). Or string them up

890
Dried mushrooms make a marvelous addition to sauces, soups, and gravies. They should be stored
in airtight jars to protect them from insects. This array of dehydrated delicacies includes Agaricus,
Boletus, Chroogomphus, Craterellus, Clitocybe, and Marasmius.

on thread and hang them in a warm, dry place. Remove all maggots before drying mush¬
rooms, and try to get them as clean as possible without washing them. If necessary, they
can be cleaned before use by placing them in a strainer and scalding them with water.
When thoroughly dried (brittle), the mushrooms should be stored in an airtight jar to
protect them from insects and mold. They will keep for months in this state. They can also
be pulverized or powdered and used as a condiment. Certain mushrooms, such as Can-
tharellus cibarius and Laetiporus sulphureus do not dry well (they become too tough and
leathery). Others, like Marasmius, Boletus, Leccinum, Craterellus, Chroogomphus,
Morchella, and Agaricus are excellent.
There are several methods for reconstituting dried mushrooms, depending on the type
of mushroom and kind of dish. They can be crumbled directly into soups or sauces, but
should be soaked first if they’re to be put in drier foods. An excellent method is to place
dried pieces between wet paper towels overnight. This allows them to absorb moisture
gradually while retaining their flavor.

FREEZING

Freezing is another excellent and easy way to preserve mushrooms, pro viding you saute
them briefly beforehand. (If you freeze raw mushrooms, they will decompose as soon as
they thaw out.) Practically all mushrooms freeze well—I have stored Amanita calyptrata
for over one year without any noticeable change in flavor! The mushrooms should be
stored in an airtight container (e.g., a ziplock bag). Mushroom sauces and stocks can also
be frozen for later use.

CANNING

Canning is a big undertaking, and is only worthwhile when you have an enormous
amount of mushrooms. A pressure canner, mason jars, and lids are required. Wash the
mushrooms thoroughly and let them cook in their own juices for a while. Then pack them
in sterilized pint jars, cover with boiling water, seal, and process at 10-15 pounds pressure
for 30 minutes (a very necessary step because they can easily become tainted with botulism).
A little ascorbic acid (vitamin C) can be added to retain color. Mushrooms will keep for
years if canned properly, but failure to observe sterile procedures can be disastrous. In
my experience, Cantharellus, Tricholoma, Dentinum, Boletus, Hericium, and Agaricus
can well.
891
892 MUSHROOM COOKERY

PICKLING

Pickling is just marinating on a larger and more elaborate scale. In Europe, mushrooms
are often preserved under oil and vinegar. It is a fairly simple procedure, but obviously,
the mushroom flavor is masked. Jars should be sterilized, but pressure cooking isn’t
necessary because of the acidic medium. I have successfully pickled Cantharellus, Clito-
cybe nuda, and Amanita calyptrata.

SALTING

This is another technique that is popular in some parts of Europe. The mushrooms
are cooked briefly and then packed in rock salt and stored in airtight jars in the refrigerator.
Naturally they are salty this way, but the salt is easily dissipated by adding the mushrooms
to soups or other watery dishes.

MUSHROOM TOXINS
MOST CASES of “mushroom poisoning” are the result of allergies* (idiosyn¬
cratic reactions or hypersensitivity), overindulgence (especially of raw mush¬
rooms), or food poisoning (ingestion of rotten mushrooms). All three usually
result in nausea, vomiting, and/or diarrhea. Another common type of “mush¬
room poisoning” is imaginary—people who have lingering doubts about the
safety of their meal are apt to experience discomfort whether or not there is a
physiological basis for it. All the more reason not to eat a mushroom unless
you’re absolutely certain it’s edible!
The two most common causes of mushroom poisoning are carelessness and
ignorance. Despite what people say, mushroom experts do not die from mush¬
room poisoning. But of course, it is much more sensational for newspapers to say
they do. Relatively few species are poisonous, but some of the most dangerous
ones are exceedingly common, and almost any poisonous mushroom can be fatal
to a small child or a person in poor health. Therefore it is useful to learn about the
different kinds of mushroom poisoning, should you or a friend experience
discomfort. A brief rundown of the major groups of mushroom toxins is pre¬
sented here. “Mushroom toxin,” as defined here, is a compound which produces
an abnormal effect on the human body. This definition encompasses mind-
altering drugs such as psilocybin, whether or not they are ingested deliberately.
As with any kind of poisoning, the two most important things to do are to seek
immediate medical attention and identify the agent responsible. Idiosyncratic
reactions to edible mushrooms are generally not serious enough to warrant a trip
to the hospital, but if there is any doubt, consult a physician.

AMANITA-TOXINS (AMATOXINS)

Mushrooms: Amanita phalloides, A. ocreata, A. verna, A. virosa, A. bisporigera;

Conocybe filaris; Galerina autumnalis, G. marginata, G. venenata;
Lepiota castanea, L. helveola, L. josserandii & close relatives.

Poisoning by amatoxins is extremely serious, with a fatality rate of about 50%. It is

doubly dangerous because the symptoms are delayed for 6-24 hours after ingestion of the
mushroom, by which time the toxins have been absorbed by the body.
There are several groups of amatoxins, at least in the Amanitas. Phallolysin was the
first toxin discovered. It destroys red blood cells when injected into rats and has a very
*In this book the term “allergy” is used loosely to describe an adverse reaction to a normally harmless mushroom
even though it may not be a genuine hypersensitivity.
MUSHROOM TOXINS 893

high mortality rate. However, it is unstable and apparently destroyed by cooking and/or
the human digestive tract. A group of complex cyclic polypeptides called phallotoxins
comprise the second group. They are also fatal when injected intravenously (they attack
the liver), but are apparently destroyed by the digestive tract. It is another group of poly¬
peptides, the amanitins, that are the culprits. They are twenty times more lethal than the
phallotoxins. Their concentration varies tremendously from individual mushroom to
individual mushroom, but an average fatal dose is about 2 ounces (fresh weight) of
Amanita phalloides.
All recent mushroom-induced fatalities in California have been caused by Amanita
phalloides and A. ocreata. Both are large, handsome, tempting mushrooms, whereas
the other deadly types (such as Galerina) are small and nondescript and therefore unlikely
to be eaten. Of course, there is no excuse for eating any of these mushrooms if you take
the time to learn about them.

Symptoms and Treatment

Amanitin poisoning usually manifests itself in four stages: (1) a latency period of 6-24
hours after ingestion, during which time the toxin is actively working on the liver and
kidneys, but the victim experiences no discomfort; (2) a period of about one day charac¬
terized by violent vomiting, bloody diarrhea, and severe abdominal cramps; (3) a period
of about one day during which the victim appears to be recovering (if hospitalized, the
patient is sometimes released!); (4) a relapse, during which liver and kidney failure often
leads to death. There is sometimes more than one relapse.
The effects of the toxin are centered on the liver and kidneys, but amanitin damages
tissue throughout the body by inhibiting RN A synthesis within each cell. To make matters
worse, the kidneys are apparently unable to eliminate amanitin from the body. The
pancreas, adrenal glands, heart, lungs, muscles, intestines, and brain may be damaged,
not only by the amanitin, but indirectly because of liver and kidney failure. It is a slow
and painful way to die.
There is no known antidote to amatoxin poisoning. Treatment is largely supportive and
symptomatic—maintaining blood sugar and salts, eliminating urea by dialysis, and
helping the body to get rid of the toxins. If you have any reason to think someone has eaten
a deadly Amanita (or an amanitin-containing mushroom), don’t wait for the symptoms
to appear! If the person is taken to the hospital soon after ingesting the mushrooms, at
least some of the toxins can be removed before they are absorbed.

GYROMITRIN (MONOMETHYLHYDRAZINE)

Mushrooms: Several Gyromitra species (especially G. esculenta & G. infula), also

many related Ascomycetes (e.g., Verpa, Cudonia, Helvetia spp.)

Gyromitrin’s product of hydrolysis, monomethylhydrazine (MMH), is a very toxic

carcinogenic compound used in the manufacture of rocket fuel. Gyromitrin poisoning has
puzzled scientists for many years because of the very narrow threshold between complete
absence of discomfort and severe poisoning or even death. This is due to the volatile nature
of gyromitrin, which is removed by the process of cooking or drying providing it has a
chance to escape. Gyromitras cooked in a closed pan can cause severe poisoning, and
inhalation of the vapors is dangerous. Raw Gyromitras, of course, pose the greatest threat.
The situation is complicated by differences in the toxicity of different geographical
strains. Gyromitra esculenta has caused numerous fatalities in Europe, but not a single
one in California.

Symptoms and Treatment

The symptoms, which appear 2-24 hours after ingestion, include headaches, abdominal
distress, severe diarrhea, and vomiting. In severe cases the liver, kidney, and red blood
cells are damaged (much as in poisoning by amatoxins), which may result in death.
Treatment is largely supportive; a physician should be consulted.
894 MUSHROOM TOXINS

MUSCARINE
Mushrooms: Inocybe species; Clitocybe dealbata and seveal relatives; Omphalotus
species, and certain red-pored species of Boletus.

Muscarine was originally isolated in Amanita muscaria, but occurs in that mushroom
in insignificant amounts. However, many Inocybes contain large amounts of muscarine—
enough so that they can be fatal in large quantities.

Symptoms and Treatment

The effects, which manifest themselves 15-30 minutes after ingestion, are focused on
the parasympathetic (involuntary) nervous system. They include excessive salivation,
perspiration, tears, and lactation (in pregnant women), plus severe vomiting and diarrhea.
These symptoms may be accompanied by visual disturbances, irregular pulse rate, de¬
creased blood pressure, and difficulty in breathing. The victim normally recovers within
24 hours, but in severe cases, death may result from respiratory failure. Atropine is a
specific antidote, but must be administered by a physician.

IBOTENIC ACID/MUSCIMOL
Mushrooms: Amanita muscaria, A. pantherina, A. gemmata.

There are many contradictions in the literature regarding the principal toxins of
Amanita muscaria and A. pantherina. Muscarine was originally believed to be the toxin,
and then bufotenine was put forth as a candidate. It turns out, however, that the main
active principle is ibotenic acid, which is converted by the human body into muscimol, a
more powerful form that passes out in the urine.
Amanita muscaria is apparently one of the oldest intoxicants known. Its use by certain
Siberian peoples has been extensively documented and R. Gordon Wasson, in his book
SOMA: The Divine Mushroom of Immortality, makes a convincing case for it being
the mystical Soma plant of the RgVedas (sacred Hindu texts). It may have been used
throughout Eurasia in ancient times, but if so, its use has been suppressed. Curiously,
many Europeans fear A. muscaria more than its deadly cousin, A. phalloides. John
Allegro’s attempt to link it to the origins of Christianity (The Sacred Mushroom and the
Cross) is far-fetched and abstruse.
Amanita muscaria is erroneously listed in older books as deadly poisonous. It can be
fatal in large doses, but so can practically any poisonous mushroom. According to most
sources, A. pantherina is somewhat more dangerous, while A. gemmata is less so. Deli¬
berate ingestion of these mushrooms has increased now that it has been shown that they
are not as dangerous as once believed. However, their effects vary greatly from person to
person. As people’s metabolisms are different and the concentrations of the toxins vary
from mushroom to mushroom, there is no way of predicting what one’s reaction will be.
Some people experience extreme discomfort, others have vivid dreams, still others ex¬
perience no effects whatsoever. The ingestion of these mushrooms is definitely not
recommended here. Incidentally, not all of the toxins have been identified. For instance,
neither pure ibotenic acid nor muscimol produce the nausea and vomiting that frequently
occurs after eating A. muscaria.

Symptoms and Treatment

Symptoms normally appear 30 minutes to 2 hours after ingestion, and last for several
hours. Nausea and vomiting are common, but the principal effects are on the central
nervous system: confusion, mild euphoria, loss of muscular coordination, profuse
sweating, chills, visual distortions, a feeling of greater strength, and sometimes halluci¬
nations, delusions, or convulsions. (An inordinate number of “trippers” mistake them¬
selves for Christ). Drowsiness is also a common phenomenon. In fact, those who ingest
MUSHROOM TOXINS 895

A. muscaria frequently fall asleep (“swoon”), to awaken hours later with little or
no memory of their experiences. There is no “hangover” effect as with alcohol, but most
people who try it (including myself) do not wish to repeat the experience. Since muscimol
passes out through the urine, Siberian users “recycled” their A. muscaria by drinking their
own urine. I know of no one in the United States who has tested this approach.
Treatment of muscimol poisoning is largely supportive—reassuring the victim that the
effects are temporary. In the mistaken belief that muscarine was the principal toxin, older
texts prescribe atropine as an antidote. Atropine, however, is likely to exacerbate the
effects of ibotenic acid/ muscimol.

PSILOCYBIN/PSILOCIN
Mushrooms: Psilocybe baeocystis, P. caerulescens, P. cubensis, P. cyanescens,
P. semilanceata and many others; also certain Panaeolus species (e.g.,
P. cyanescens, P. subbalteatus)\ certain Conocybe and Gymnopilus
species; possibly Pluteus salicinus, P. cyanopus, and others.

These indole derivatives are well known for their psychedelic properties, and “psilo¬
cybin mushrooms” are often consumed for recreational purposes. Several hallucinogenic
mushrooms played an important role in the religious and medicinal rites of Native
Americans in Mexico and Central America. But the Spaniards suppressed their use to
such an extent that their existence was seriously doubted by early 20th century botanists.
The mushrooms were “rediscovered” in Oaxaca in the 1930’s, and 20 years later they were
identified as belonging principally to the genus Psilocybe. Since then their properties
have been so publicized that Oaxaca has been inundated by pleasure-seeking gringos. It
has subsequently been discovered that many psilocybin-containing mushrooms grow in
the United States as well.
Psilocin (a dephosphorylated version of psilocybin) is about ten times as active as psilo¬
cybin. Most psilocybin-containing mushrooms have only a trace of psilocin, but the
human body coverts most of the psilocybin into psilocin. A blueing reaction associated
with the presence of psilocybin and psilocin is caused by an enzyme that oxidizes psilocin.
However, not all mushrooms that stain blue contain psilocybin or psilocin, and not all
“psilocybin mushrooms” stain blue.

Symptoms and Treatment

Symptoms are similar to those of LSD. Shortly after ingestion, and for a duration of
several hours, the “victim” experiences heightened color perception, visual distortions,
rapdily shifting shapes and images, a “kaleidoscope effect” with eyes closed, elation or
hilarity, and hallucinations or delusions. Nausea and vomiting are rare. Some people
report the sensation of leaving their bodies, or of traveling into the future or past, or other
highly subjective experiences. Others experience profound anxiety.
In case of accidental ingestion or a “bad trip,” the victim should be repeatedly assured
that the effects are temporary. A factor to bear in mind is that transferring the person to
an unfamiliar environment can be frightening, and that sedatives may worsen the effects,
especially if administered forcibly. LSD, incidentally, was derived from another fungus,
Clavicepspurpurea (wheat ergot).

GASTROINTESTINAL IRRITANTS
Mushrooms: Many, including: Agaricus californicus, A. hondensis, A.placomyces, A.
praeclaresquamosus, A. xanthodermus; Boletus sat anas, B. erythropus,
B. haematinus, B. pulcherrimus, B. subvelutipes; Chlorophyllum molyb-
dites; Entoloma species; Gomphus floccosus; Hebeloma species; many
acrid and/or purple- or yellow-staining Lactarius species; Laetiporus
896 MUSHROOM TOXINS

sulphureus and Lepiota rachodes & L. naucina (sometimes); Naema-

toloma fasciculare; Omphalotus species; Ramariaformosa and relatives;
many acrid Russula species; Scleroderma species; and several Tricholoma
species (notably T. pardinum and T. pessundatum).

As evidenced by the list, this is by far the largest and most prevalent group of mushroom
toxins. Very few of the active principles have been identified, however, perhaps because
they are rarely fatal. The most frequent culprits in gastrointestinal poisoning are the
phenol-smelling Agaricus species and Chlorophyllum molybdites, undoubtedly because
they closely resemble edible types. The most dangerous are Entoloma species, Hebeloma
species, Tricholoma pardinum, Boletus satanas and close relatives (raw), Naematoloma
fasciculare, and Chlorophyllum molybdites. “Allergic” reactions to edible mushrooms
normally take the form of gastrointestinal upset.
Symptoms and Treatment
Symptoms usually appears shortly after ingestion (20 minutes-4 hours). They include
nausea, vomiting, cramps, and diarrhea,* which normally pass after the irritant is expelled.
Severe cases, however, may require hospitalization. Treatment is largely supportive—
helping the body to eliminate that which it is not equipped to handle. Though not as serious
as other types of mushroom poisoning, gastrointestinal upsets are not to be taken lightly,
as evidenced by the fact that many people acquire a lingering distaste for mushrooms after
an all-night bout with nausea and diarrhea.

MISCELLANEOUS TOXINS

Coprinus atramentarius contains a disulfram-like compound (coprine) that reacts

with alcohol in the body to produce acetaldehyde, which in turn produces a peculiar but
transitory set of symptoms; reddening of the ears and nose, a metallic taste in the mouth,
lightheadedness, rapid heart beat, a throbbing sensation, and sometimes nausea and
vomiting. Recovery is normally spontaneous and complete. The alcohol needn’t be con¬
sumed simultaneously with the mushrooms to have a synergistic effect—therefore anyone
who indulges in alcohol regularly should not eat C. atramentarius. Individual reactions
vary, suggesting that some people may be more sensitive or the mushrooms themselves
may differ in coprine content. It has also been suggested that raw C. atramentarius does
not actually contain coprine, but that it is formed in the process of cooking. Similar
alcohol-related effects have been reported for Clitocybe clavipes.
Paxillus involutus is said to be toxic raw, but is eaten by many people (especially Euro¬
peans) after being pickled or parboiled. However, the human body apparently develops
a sensitivity to it, manufacturing antibodies that destroy red blood cells. Thus, someone
who has eaten it for years can suddenly be poisoned (even fatally!).
Cortinarius orellanus, C. gentilis, and some close relatives are commonly fatal. They
contain toxins which, like the amatoxins, attack the liver and kidneys. Symptoms do not
appear for up to three weeks after ingestion of the mushrooms, making diagnosis and
treatment much more difficult. Some of these Cortinarii occur in the United States. As
they are difficult to identify, all dry-capped Cortinarius species are best avoided.
Naematoloma fasciculare, a common wood-loving mushroom, can also cause liver and
kidney damage. Fortunately, it is rarely eaten because of its bitter taste. There is also
evidence to suggest that the substrate on which a mushroom is growing can affect its
edibility. The edible honey mushroom (Armillariella mellea), for instance, will often
cause digestive upsets when growing on buckeye.
Last but not least, there is the danger of contamination by pesticides and other environ¬
mental poisons. Always be aware of this possibility, especially when picking mushrooms
in towns, along well-traveled roads, and in forests, fields, or range land where herbicides
and pesticides are used.
*The literal translation of the Japanese name for certain poisonous Tricholomas is“Unable to Cross the Valley”—
presumably because one is unable to cross the valley after eating them!
Russula cyanoxantha, a robust mushroom with brittle flesh and white gills. The cap color is ex¬
tremely variable (see description on p. 94). Maggot-free specimens such as these are unusual.

Lactarius uvidus (see p. 75) is one of several purple-staining milk caps. The cap is usually sticky or
slimy, but in one mountain-loving variety it is practically dry. Note how the stalk snaps open cleanly
like a piece of chalk—a characteristic feature of Lactarius and Russula.

Melanoleuca evenosa group (see p. 171) is reminiscent of a Russula or Tricholoma but has a fibrous
stalk and amyloid white spores. Look for it in the spring under mountain conifers soon after the
snow melts. Note how the gills are crowded and whitish.
Two agarics that typically grow in dense clusters. Top: Clitocybula familia (see comments under
Collybia acervata, p. 215) grows on rotting wood. These are young specimens; as they mature the
caps will broaden somewhat. Below: Lyophyllum decastes group (see p. 174) is usually but not
always terrestrial. This cluster is rather unusual because-the caps have watery spots.

Abnormalities are common in mushrooms. They are the result of environmental influence or im¬
proper development; they are not inherited or passed on. One widespread abnormality is the develop¬
ment of a small rosette of gills (“rose gill”) on the cap surface. In extreme cases, an entire mushroom
may ride “piggy-back” on another, like these Russulas (left). Albino individuals also occur. In this
cluster of Craterellus cornucopioides (right), half is normally colored and half is whitish! Other ab¬
normalities include aborted, parasitized, or sterile specimens and failure of the cap or gills to form.
 899

WHAT IT ALL MEANS

(A Short Dictionary of Scientific Names)
Otidea, Ramaria, Tricholomopsis, Volvariel/a
Exidia, Stropharia, Hygrophoropsis, Arcangeliella

Y ou may groan at the sight or sound of scientific names, but the paucity of common names
for mushrooms forces us to use them. As explained in the chapter on classification (p. 8),
many people are intimidated by scientific names because they are derived from Latin and
Greek. “How do you remember all those names?” I am repeatedly asked. Y et posers of this
question have usually mastered many scientific names without realizing it:

Asparagus, Magnolia, Sassafras, Eucalyptus

Rhinoceros, Hippopotamus, Chrysanthemum, Citrus

Since that which is understood is more likely to be remembered than that which is not,
learning the meanings of scientific names can increase your ability to remember them.
By refusing to be intimidated, you can demystify the names of mushrooms while you are
demystifying the mushrooms themselves.
Scientific names can be divided into three categories: descriptive, honorary, and
geographical. (A fourth possible category, nonsensical, needs no elaboration.)
Descriptive names are the most numerous as well as the most helpful. They can aid in
the identification process because they usually tell us something significant about the
mushroom, its habitat, or time of growth. For example:

Flav- means yellow, and it is a safe bet that Hygrocybeflavescens, Tricholoma

flavovirens, Boletus flaviporus, and Russula claroflava are at least partially
yellow. (They are.) Similarly, Macrocystidia cucumis(macro-large, cucumis
^cucumber) has giant cystidia (sterile cells) on its gills and smells like cucum¬
ber, and Hypsizygus tessulatus (hyps-hig\\ up, zygus-yoked or joined,
tessw/tf/ws-mosaic-like) grows high up on elms and develops mosaic-like
scales on its cap as it matures.

You will soon discover that many of the longest and most “difficult” names are actually
composed of shorter, more familiar elements—familiar because the English language
is a thicket with many Latin roots. For instance:

Climacodon is derived from climac- (ladder, as in the English word climax)

and odon (tooth or teeth, as in the English word orthodontist), and the Greek
element derm (skin) appears in many mushroom names (xanthodermus,
calyptroderma, Dermocybe, etc.) as well as English words like dermatolo¬
gist, epidermis, and pachyderm.

A few descriptive names, unfortunately, are apparent misnomers, forthey bear no obvious
relation to the mushroom and may even contradict it:

Appendiculat- means with a fringe or small appendage, but Boletus appendi-

culatus has neither. Although alnicol- means alder-dweller, Lactarius al-
nicola favors conifers and oaks (but may have originally been collected in a
mixed forest containing alder).

Honorary names usually commemorate a mycologist, mycologist’s best friend, or

mushroom collector. In most cases an -ii, -i, or -ae is added to the person’s name if it is a
species(e.g., kauffmanii, barrowsii, smithii), oran-a, -ea, or-/'aifitisagenus(e.g., Barssia).
Other suffixes, particularly diminutives or those meaning “pertaining to,” can also be
added (e.g., booniana, Longula, Lenzites).
900 SCIENTIFIC NAMES

Honorary names are usually recognizable as such because they don’t sound Latin.
Some, however, may not be obvious, particularly those that honor Europeans. Examples:

Hohenbuehelia is named after the Austrian botanist Hohenbuhel, Arcan-

geliella and Battarrea after the Italian mycologists Arcangeli and Battarra,
and Galiella and Rozites after the French mycologists Le Gal and Roze.

One major drawback of an honorary name is that it tells us nothing about the mushroom
itself. On the other hand, precisely because it tells us nothing it cannot be misleading!
Geographical names are formed by adding an adjective suffix to the name of a particular
region or locality. Most are self-evident (e.g., californicus, marylandensis, cubensis,
mexicana, olympiana). Some are Latinized, however (e.g., novaboracensis, of New York,
suecica, of Sweden) and others are obscure (e.g., hondensis, named after bustling La
Honda, California, and pitkinensis, which immortalizes populous Pitkin, Colorado).
Geographical names can be deceptive because they usually indicate where a species was
discovered rather than where it occurs. For instance, Longula texensis, originally col¬
lected in Texas, and Suillus sibiricus, first found in Siberia, are both widely distributed
in western North America.

SUFFIXES
Suffixes can have specific meanings, vague meanings, multiple meanings, or no
meaning at all. Some suffixes simply transform nouns into adjectives. Others indicate
possession, likeness, size, action, or degree. Still others designate gender—a concept
familiar to anyone who has studied French or Spanish. Thus a single root word can have
several forms depending on the gender and spelling of the word(s) with which it is used.
Example:

Brunne-, the root word for brown, becomes brunneus (masculine form) in
Boletus olivaceobrunneus, brunnea (feminine form) in Leptonia vinaceo-
brunnea, brunneum (neuter form) in Tricholoma albobrunneunm, and
brunneo-, brunnea-, or brunnei (combining forms), as in Clitocybe brunneo-
cephala. Suffixes with specific meanings can also be attached, as in brun¬
ne scens (“becoming brown”) or brunneola (“less than brown, brownish”).

As you can see, it is not necessary to know all the quirks and nuances of Latin and Greek
grammar in order to recognize root words such as brunne in their various manifestations.
Since suffixes are sundry and sometimes difficult to recognize, the more common ones
are listed separately. However, common prefixes as well as most suffixes with very specific
meanings (e.g., odon-iooi\\, pes-foot or stem, osm-=odor) can be found in the dictionary
of word elements.
The dictionary is for descriptive names only. It does not include people or places, self-
evident words such as giganteus (gigantic), caninus (canine), parasitica (parasitic), or
fragilis (fragile). Nor does it include names whose meanings are obscure, debatable, or
unknown. A hyphen indicates that the root word can have various endings (see discussion
of suffixes).
Space does not permit listing all the root words used in naming mushrooms, but with
the help of the dictionary and the material preceding it, you should be able to make sense
out of most of the scientific names in this book—and enrich your English vocabulary in
the process! Examples:

(1) cortin-curtain, arius-with or pertaining to, thus Cortinarius means

“with a curtain.”
(2) cybe^head, con-cone, derm-skin, mo^fiber, c/zV^sloping, psil-smooth,
thus Conocybe means “cone head,” Dermocybe means “skin head,” Ino-
cybe means “fiber head,” Clitocybe means “sloping head,” and Psilo-
cybe means “smooth head.”
LATIN AND GREEK SUFFIXES 901

(3) /yc=wolf, p^r<io/7=flatulence, thus Lycoperdon means “wolf fart.”

(4) vermicul-little worm, osw^full of, thus vermiculosus means “full of little
worms” (i.e., maggoty).
(5) canthar-drinking cup, ellus is a diminutive suffix, thus Cantharellus
means “little drinking cup.”
(6) paxill-smstll stick or stake and -us is a masculine gender ending, thus
Paxillus means “small stick” or “small stake.”
(7) leuc-white, thus Leucopaxillus means “white Paxillus.”
(8) a as a prefix means without, p/iy//=leaves or gills, phor-bearing or bearer,
ales denotes an order of fungi, thus Aphyllophorales means “order of fungi
not bearing gills.”
(9) orth-straight, odont-teeth, ist is an English suffix meaning “one who,”
thus orthodontist means “one who straightens teeth.”
(10) succ-juice or sap, ulent-full of, thus succulent means “full of juice.”

Common Suffixes
Note: Some suffixes, particularly those with feminine endings, may not be in alpha¬
betical order because they are listed under their masculine form. For instance, -ata
is listed under -atus, and -aria is listed under -arius.

-a: most common feminine gender ending -ator: see -tor

-abilis: see -bilis -atus, -ata, -atum: (1) possessing, furnished
-abulum, -aculum: see -bulum with (e.g., armillatus, ringed or with a
-aceae: denotes a family of fungi or plants ring) (2) resembling, similar to (e.g.,
-aceus, -acea, -aceum: color of, made of, with ovatus, egglike)
quality of, closely resembling, relating to -ax: apt or tending to
(e.g., olivaceus, olive-colored) -bilis: capable of, able to, tending to (e.g.,
-acius, -acia, -acium: pertaining to, pos¬ mutabilis, able to change)
sessing -bulum: instrument, container, agent, means
-ae: named after, pertaining to (e.g., annae) -bundus, -bunda, -bundum: implies action
-aeus, -aea, -aeum: belonging to -ceae: see -aceae
-ago: (1) resembling, similar to (2) made of, -cellus, -cella, -cellum: diminutive
color of -cillus, -cilia, -cillum: diminutive
-ales: denotes an order of fungi or plants -colus, -cola, -colum: inhabitant, dweller(e.g.,
-alis, -ale: belonging to, pertaining to, relating corticola, bark dweller)
to, characteristic of -cuius, -cula, -culum: diminutive (e.g., auri¬
-aneus, -anea, -aneum: made of, color of, cula, small ear)
resembling -cundus, -cunda, -cundum: able to, tending to
-ans: implies action (like -ing in English, e.g., (e.g., rubicundus, tending to redden)
radicans, rooting) -e: (1) gender ending (usually feminine) (2) see
-anus, -ana, -anum: pertaining to, charac¬ -ae
teristic of (often connotes position, e.g., -ea: see -eus
montanus, of the mountains) -ellus, -ella, -ellum: diminutive (e.g., Can¬
-arion: diminutive tharellus, small cup)
-aris, -are: belonging to, pertaining to -ens: see -ans
-arius, -aria, -arium, -arum: pertaining to, -ensis, -ense: indicates place of growth or
possessing, furnished with, relating to, origin (e.g., arvensis, growing in fields)
(e.g., Cortinarius, with a curtain) -enus: belonging to
-ascens: becoming, almost, somewhat (e.g., -er, -eres: provided with
purpurascens, becoming purple) -escens: becoming, almost, somewhat (e.g.,
-aticus, -atica, -aticum: denotes place of rubescens, becoming red or almost red)
growth or origin (e.g., silvaticus, growing -estris, -ester, -estre: indicates place of growth
in the woods) or origin (e.g., campestris, growing in
-atilis, -atile: denotes place of growth (e.g., fields)
saxatilis, growing among rocks)
902 LATIN AND GREEK SUFFIXES

-etes: (1) denotes class or larger grouping of -ius, -ia, -ium: (1) pertaining to, characteristic
fungi (2) one who is of, resembling (e.g., regius, pertaining to
-ettus, -etta, -ettum: diminutive kings) (2) occasionally used as diminu¬
-etum: denotes collective place of growth tive or comparative
-eus, -ea, -eum: (1) pertaining or belonging -ivus, -iva, -ivum: (1) possessing, furnished
to (2) color of, made of, similar to (e.g., with (2) pertaining to(3) capable of, able to
melleus, honey-colored) -izans: becoming like, resembling
-formis, -forme: resembling, especially in -limus: see -rimus
shape or form (e.g., strobiliformis, shaped -ma, mus: indicates action or result
like a pine cone) -nellus, -nella, -nellum: diminutive
-genus, -gena, -genum: born of, originating -odes: see -oides
from -oides, -oideus, -oidea, -oideum: resembling,
-i: see -ii similar to (e.gphalloides, like a phallus)
-ia: (1) see -ius(2) see -a (3) see -ae -olentus, -olenta, -olentum: see -ulentus
-iacus, -iaca, -iacum: pertaining to -olus, -ola, -ole, -olum: (1) diminutive (2) less
-iae: see -ae than, somewhat (e.g., luteolus, yellowish)
-ianus, -iana, -ianum: see -anus -on: neuter gender ending
-ibilis: see -bilis -onius, -onia, -onium: pertaining or related to
-icons: becoming, almost, closely resembling -opsis: resembling, similar to (e.g., Tricho-
(e.g., nigricans, becoming black) lomopsis, like Tricholoma)
-icius, -icia, -icium: (1) made of, pertaining to -orius, -oria, -orium: capable of, able to (e.g.,
(2) implies or denotes action tine tor ius, able to dye)
-icos: pertaining to -orum: pertaining to
-icus, -ica, -icum: belonging to, pertaining to -os: masculine or feminine gender ending
(e.g., californicus, of California) -osus, -osa, -osum: denotes fullness, abun¬
-ides: resembling, similar to dance, marked degree of development
-idion, -idius, -idia, -idium: diminutive (e.g., (e.g., succosa, full of juice)
Gomphidius, little peg) -otus, -ota, -otium, -otum: (1) resembling,
-idus, -ida, -idum: (1) resembling, similar to similar to, possessing, furnished with
(2) becoming, a\most(e.g.,a lb ida, whitish) (2) see ot- in dictionary
-iellus, -iella, -iellum: diminutive -rimus, -rima, -rimum: superlative (e.g., pul-
-iensis, -iense: see -ensis cherrimus, most beautiful)
-ii: named after (e.g., kauffmanii) -ros: pertaining to
-ilis, -He: able to, capable of, with property of -tatos, -tate, -taton: superlative
(e.g.,fragilis, able to be broken) -ter: agent or means
-illus, -ilia, -ilium: diminutive -teros, -tera, -teron: comparative
-imos: pertaining or relating to -tes, -tis, -tor, -tra, -tria, -tron, -tros, -trum,
-imus, -ima, -imum: superlative -trus: denotes agent, means, tool, or object
-inans: becoming, almost (e.g., necator, killer)
-ineus, -inea, -ineum: color of, made of, -ua: see -uus
similar to (e.g.,ferruginea, color of rust) -ugo: (1) made of (2) able to, capable of (3)
-inius, -inia, -inium: named after, relating to disease or rust
-inos: (1) made of (2) see -inius -ulentus, -ulenta, -ulentum: denotes full¬
-inus, -ina, -inum: (1) pertaining or belonging ness, abundance, marked degree of de¬
to, possessing, resembling (2) diminutive velopment
-ioides: see -oides -ulus, -ula, -ulum, -ullus: (1) pertaining to
-ion: (1) diminutive (2) denotes occurrence (2) diminutive (3) denotes lesser degree of
-ior: (1) comparative (e.g., strictior, very development (e.g., dulcidulus, slightly
straight) (2) see -ios sweet)
-ios: pertaining or relating to -um: most common neuter gender ending
-is: pertaining or relating to -unculus, -uneula, -unculum: diminutive
-isce, -iseus, -iscos: diminutive -urus, -ura: (1) implies result or action (2)
-issimus, -issima, -issimum: superlative (e.g., pertaining to
speciosissimus, showiest) -us: most common masculine gender ending
-istos: comparative or superlative -uscu/us, -uscula, -usculum: diminutive
-itas: see -itius -ustris, -uster, -ustre: see -estris
-ites, -itis: pertaining to, named after -utus, -uta, -utum: possessing, furnished with
-iticus, -itica, -iticum: (1) possessing, fur¬ -uus, -ua, -uum: implies result or possibility
nished with (2) capable of, able to of action
-itius, -itia, -itium: (1) with quality or color of, -ydrion, -yllion: diminutive
similar to (2) implies action or result -ys: comparative or superlative
 903

Dictionary of Selected
Latin and Greek Word Elements
A
areolat-: areolate (cracked like dry mud)
a-, ab-, ad-, etc.: prefix meaning without,
argent-: silver
absent, or away from argill-: clay
abies, abie-, abiet-: fir, firs argillace-: clay-colored (dull yellow-brown)
acanth-: thorn, prickle argyr-: silver
acer-: (1) maple (2) acrid, sharp armen-: apricot, apricot-color
acerb-: bitter, sour armill-: bracelet, ring
acerv-: heap arquat-: (1) bowed, arched (2) rainbow-
acet-: vinegar colored
acetabulum: vinegar cruet arv-: field(s)
acicul-: bristle, needle, splinter asc-: bladder, bag, wineskin
acr-: acrid asper-, aspr-: rough
acumin-, acumen-: sharp point aster-, astr-: star
acut-: sharp, pointed, acute astragalin-: pertaining to goldfinch
adust-: scorched ater-, atr-: black, very dark
aegerit-: pertaining to poplar atrament-: ink
aereus: copper, bronze atrat-: clothed in black
aerug-: blue-green, green, or deep green attenuat-: narrowed, tapered, reduced
aest-: summer augean-: pertaining to Augeos, who main¬
affin-: related; adjacent tained an uncleaned stable for 30 years
agaric-: ancient term for mushroom august-: majestic, august
agath-: pleasant, good, excellent aur-: see aure- and auri-
agglutin-: glued auranti-: orange
agr-: field aurat-: gilded, ornamented with gold
alb-: white aure-: gold, golden
aleur-: wheat flour auri-, auric-: ear
alii-: garlic auriscalpium: ear scrape
aln-: alder austr-, austral-: southern
alutace-: leather-colored (yellowish-tan) avellan-: avellaneous (pale gray tinged with
amab-: pretty pink)
amanita-: ancient term for mushroom azur-: azure, blue
amar-: bitter
B
amentace-: pertaining to catkins
amethyst-: violet (amethyst) baccat-: (1) set with pearls or berries (2) soft,
amianth-: unpolluted, spotless pulpy
ammo-: sand badi-: reddish-brown
amoen-: pleasant, lovely bae-: slim, small
amygdal-: almond balsam-: balsam, fir
amylo-: amyloid (containing starch) balte-: belt, girdle
ananas: pineapple basidi-: basidium or basidia (derived from
andros-: a tiny herb or plant word for little base or pedestal)
angi-: vessel (derived from little base
angust-: small, narrow basilar-: fixed at the base of something, basal
annul-: ring, annulus bell-: pretty, handsome
anth-: flower benzoin-: resinous
anthrac-: coal, charcoal betul-: birch
apothec-: storehouse bi-: prefix meaning two, double, twice
appendiculat-: with a small appendage, biennis: lasting two years
appendiculate bol-: throw, thrower
apt-: fastened, bound bolaris: with little lumps of paint
aqu-: watery bolbit: cow dung
arachn-: spider or spider web bolet-, bolites: ancient words for a superior
arai-, arae-: thin, narrow kind of mushroom
arane-: spider or spider web bombyc-: silky
arcularius: maker of small chests boreal-: northern
aren-: sand botry-, botryt-: bunch of grapes
areol-: a small open space bovista: ancient term for puffball (probably
derived from word for ox)
904 LATIN AND GREEK DICTIONARY

brev-: short, brief cerrusat-: as if painted with white lead

brum-: winter cervin-: pertaining to deer; fawn-colored
brunne-: brown, dark brown chaet-: long hair or bristle
buf-: toad chamal-: false
bulb-: bulb chamonixia-: on the ground (?)
bulbos-: with a bulb, bulbous cheil-: lip, brim, margin
bulg-: wine skin, leather bag cheim-: winter
butyrace-: buttery chelidon-: russet (colored like a swallow’s
byssised-: seated in or pertaining to a mass throat)
of fine threads or filaments chelis: (1) hoof or claw (2) cloven
chil-: lip, brim, margin
C
chion-r: snowy
caelat-: chiselled, carved chlamyd-: mantle, cloak
caerul-: blue, deep blue chlor-: green, greenish, greenish-yellow
caes-: glaucous, light gray, blue-gray choir-: sow, pig
caesar-: caesarean, regal chondr-: (1) cartilaginous (2) granular
caespitos-: cespitose (tufted or clustered) chro-: (1) color (2) superficial appearance,
calamistrat-: curled skin
calig-: boot chrom-: bright color, e.g., chrome-yellow
callist-: beautiful chrys-: golden, gold
calo-: beautiful cibar-: food
calv-, calvat-: bald, hairless cibor-: drinking vessel or cup
calyc-: cup, bud, calyx ciliat-: ciliate (fringed with hairs)
calyptr-: protective cap or hood cilic-: cloth made of goat’s hair
camar-: arched, curved, vaulted cincinnat-: curly, with curled hairs
camp-: fields, plains, countryside cinere-: ash-colored, smoky, of cinders
campan-: bell cingul-: girdle, belt, collar
campanul-: little bell cinnabarin-: red, cinnabar-colored
can-: hoary, pale gray, whitish cinnamom-: cinnamon-colored
cancell-: lattice-work circell-: small ring
candid-: shing, bright, white, clear circin-: circular, or bent like the head of a
canthar-, canth-: drinking cup crosier
caperat-: wrinkled, folded cirrhat-: frayed, fringed, curled
capill-: hair citrin-: lemon-yellow
capn-: smoke clar-: clear, light, brilliant
capr-: goat clathr-: iron grating, latticework
capreol-: (1) wild goat (2) tendril claud-: lame, closed
carbon-: coal, charcoal clav-: club
carchar-: rough, jagged clavat-: clavate (club-shaped)
cam-, came-: flesh, flesh-colored clavicul-: small key
carp-: fruit, fruiting body climac-: ladder
cary-, caryc-: nut tree or nut (especially walnut clit-: sloping
or hickory) clype-: round shield
castan-: chestnut, chestnut-colored coccin-: scarlet, red
catathelasma: running down cochleat-: like a snail shell
catin-: small bowl, basin, hollow cocos-: coconut
caud-: tail cognat-: related, kindred
caul-: stalk, stem, stipe col-: dweller, inhabitant (usually a suffix)
cav-, cavi-: hollow, cavernous cole-: sheath
cedr-: cedar (Old World cedar, Cedrus) coli-: sieve, colander
cedret-: cedar woods, cedars collinit-: smeared or covered with slime
celer-: swift collyb-: small coin or coin-shaped pastry
centuncul-: a patchwork cloth columb-: dove
cep-: (1) onion (usually cepa) (2) head (see com-: hair
ceps) comat-: covered with hair
cephal-: head con-: (1) cone (2) prefix meaning with
ceps: heads conchat-: shell-shaped, shell-like
cer-: horn concinn-: neatly arranged or joined
cerace-: waxy concrescens: congealed
cerea-: wax confluens: confluent; running together
cerebr-: brain confragos-: rough, scaly
LATIN AND GREEK DICTIONARY 905

conigen-: growing on or born from cones deceptiv-, decipiens: deceiving

connat-: born together decolorans: fading
connex-: connected decor-: beautiful, befitting, elegant
connivens-: blended together, converging decurrens: decurrent (running down)
consobrin-: cousin delibut-: grease, greasy
controvers-: turned against, in opposite delica: weaned, without milk
directions delicat-: dainty, pleasing
copr-: dung delph-: womb
cor-: see cori- delphus: brother
cordy-: club, cudgel, swelling dendr-, dendron: tree
cori-: leather dens-: crowded, thick, compact
corn-: horn dent-: tooth, teeth
corniculat-: with little horns derm-: skin
cornucopi-: horn of plenty destruens: destroying
coron-: crown detonsus: sheared, clipped
coronill-: little crown, garland di-: prefix meaning two, twice
corrug-: wrinkle, wrinkles dibaphus: dyed twice
cor tic-: (1) bark or hard crust (2) cork dicty-: net
cortin-: curtain difform-: deformed
corvin-: shiny black (like a raven) dilatat-: dilated, enlarged
coss-: wood worm dipl-: prefix meaning double
cost-: rib disc-: disc
cotone-: color of wild olives or quince discised-: seated in a disc
cotyl-: cup, cup-shaped discoid-: (1) disc-shaped (2) descriptive term
crani-: skull, cranium for one color at the center of another color
crass-: fat, thick, big, heavy dissimulans: one who is disguised
crater-: cup, crater don-, dont-: tooth, teeth
cren-: notch dry-: oak
crenulat-: notched, scalloped dryad-: (1) wood nymph (2) oak
crep-, crepid-: shoe, slipper dulc-: sweet
cretace-: chalky dur-: hard, resistant, durable
crin-: hair duracin-: hard-fruited
crinitus: long-haired, very hairy E
crisp-: curly, crisped
eburne-: ivory
crist-: crest
eccilia: rolled up
croc-: saffron-colored
echin-: hedgehog, sea-urchin (hence spiny,
crocolit-: wearing saffron
prickly)
cruent-: gory, blood-colored
edodes: food
crust-: crust, crusty
edulis: edible
crustulin-: thin bread crust
elaph-: deer
crypt-: hidden, secret
elasm-: thin plate or gill
cucull-: hood, cap
elat-: raised, tall
cucum-: cucumber
elegan-, elegans: elegant, choice
cudonia: pertaining to a leather helmet
encephal-: brain
cumatil-: colored like a wave or sea water
end-, endo-: within, inner, inside (prefix)
cuspidat-: with a sharp tip, pointed
ent-: rolled inward
cyan-: blue, deep blue
enteron-: internal
cyath-: cup
ep-: see epi-
cyb-, cybe: head
cycl-: circle, wheel ephippi-: mounted as on a horse
cypt-: stooped forward epi-: prefix meaning on or upon
cyst-: blister, bladder, cyst epipterygia: surmounted by a small wing
ereb-: dark
D eric-: heath
dacry-: a tear (as in weeping) ericet-: heaths
daedal-: pertaining to maker of labyrinth, erinace-: pertaining to hedgehog
beautifully wrought erub-: red, reddish
dal-: fire-brand, charred wood erythr-: red, reddish
dasy-: shaggy, hairy esculent-: edible, good to eat
dauc-: carrot eury-: large, broad, wide
dealbat-: whitewashed evanescens: evanescent, withering away
decastes: by tens evenos-: without veins
906 LATIN AND GREEK DICTIONARY

ex-: prefix meaning without, beyond, on G

the outside, or possessed of
gal-, galact-: milk
excels-: lofty gale-, galer-: helmet or fur cap
excor-: peeled, flayed
gambos-: large-hocked, with a swollen base
exid-: staining, exuding, perspiring
gan-: shiny, lustrous
exim-: extraordinary, distinguished
gaster-, gastr-: stomach, belly
F gausapat-: woollen cloth
fag-: beech ge-: see geo-
fall-,fallax: deceptive gemmat-: gemmed
farin-: flour, meal gen-: (1) born of, originating from (2) race
farinace-: farinaceous (like fresh meal) genea: derived from gen-, or from ancient
fascicul-: bundle word for compact, knobby bodies
fastigiat-: pointed, gabled gentil-: of the same race
fav-: honeycomb geo-: earth
fell-: very bitter (bile) geotrop-: positively geotropic, i.e., erect
fenn-: Finland ger-: bearer, carrier, bearing (often a suffix)
fer-: bearer, carrier, bearing (usually a suffix) gibb-: humped, rounded
ferr-: iron gigas-: giant
ferrug-: rust, rusty gilv-: pale yellow, dull yellowish
fibr-: fiber glaber, glabr-: hairless, smooth
fibril-: fibril (fine fiber or hair) glandul-: gland
fibros-: fibrous glauc-: glaucous (pale grayish-bluish-green)
fibul-: pin or clasp or silvery
fid-: divided, forked gleb-: gleba (derived from word for clod)
fil-: thread gli-: glue
fim-: dung glischr-: glutinous, sticky
fimbriat-: fringed gloeo-: glutinous, sticky
fimet-: dung heap gloss-: tongue
firm-: firm, strong glyc-, glycy-: sugar, sweet
fistul-: tube or pipe gomph-: bolt, peg, stake, nail
flabell-: small fan gon-: joint, angle
flamm-: flame gracil-: gracile (slender, thin)
flav-: yellow gramin-: grass, grain
flocc-: woolly gramm-: sign, mark, line
flu-: secreting fluid, letting flow (usually a gran-: seed, grain
suffix) granul-: granule (fine grain)
foe-: fire grav-: heavy, strong, pregnant with
foenisecii-: pertaining to hay-making or grifola: braided fungus
lawn-mowing gris-, grise-: gray
foetens, foetid-: fetid, stinking gumm-: gum
foli-: leaf or leaves (gills) gutt-: drop, droplet
fomes,foment-, fomit-: tinder gymn-: naked
form-: shape, form, appearance gyr-: round; a circle
formos-: lovely, shapely, beautifully formed H
fornicat-: arched or vaulted
haem-, haemat-: blood, blood-red
fr act-,frag-: break
haemorrhoid-: bleeding
frond-: leaf, frond
hal-: salt, sea
fruct-: fruit hapal-: soft-boiled, tender
fulg-: shine hebe-: blunt, obtuse
fulig-: soot helv-: see helveol-
fulmin-: lightning helvella: ancient term for an aromatic herb
fulv-: fulvous (fox-colored, reddish-yellow,
helveol-, helvol-: light bay, pale reddish,
tawny) blonde, honey-colored, “the dingy color
fum-: smoke, smoke-color (gray, usually of oxen”
mixed with brown) hemi-: half
funalia: made of rope hepat-: liver
furc-: fork, forked hericium-: pertaining to hedgehog
furfur-: bran, scurf hetero-: prefix meaning irregular, dif¬
fus-: spindle ferent, other
fusc-,fusco-: dark, dusky, fuscous hiem-: winter
him-: jug
LATIN AND GREEK DICTIONARY 907

hirsut-: hirsute (hairy) jub-: mane, crest

hiri-: bristly or shaggy with weak hairs junonius: pertaining to Juno, wife of Jupiter
hispid-: hispid (roughened by stiff hairs), L
shaggy, spiny
hoi-, holo-: entire, whole lace- paint
lacer-: tear, lacerate
hort-: garden, gardens
humil-: low, stunted, humble, dwarfish lachn-: wool, woolly
hyal-: clear, transparent, colorless lachrim-: tears (as in weeping)
hydn-: ancient term for an edible mush¬ laciniat-: torn, frayed
room or tuber (truffle) lacrim-, lacrym-: tears (as in weeping)
hydr-: water lact-: milk
hygr-: moisture, humidity lacun-: cavity, pit
hymen-: membrane, hymenium laet-: bright, gay, pleasing, abundant
hyp-: see hypo- laev-: smooth
hyper-: prefix meaning above, beyond, over laevigat-: polished, smooth
hyph-: fringed with tissue, webbed, woven lag-: hare, rabbit
hypn-: tree-moss lamell-: plate or plates (gills)
hypo-: prefix meaning below, under, lamin-, lamn-: thin plate
beneath, not quite, less than normal lan-: wool
hyps-: high up land-: (1) wool (2) to tear into pieces
hysgin-: red vegetable dye lanuginos-: downy
hyster-: womb larg-: large, wide
hystrix, hystric-: porcupine laric-: larch
lat-: broad, large
IJK later-: brick
ianth-: violet laterit-: brick-colored, bricklike
icterin-: jaundiced (pale yellow or greenish- laurocerasi: pertaining to English laurel
yellow) leccinum: Italian term for fungus
ign-: fire lei-: smooth
illin-: smeared lent-: (1) pliable, tenacious (2) sticky, viscid
illot-: dirty, unwashed lenticul-: (1) lentil (2) small lens
illudens: (1) deceiving (2) emitting light lentig-: freckle, speck
im-: see in- leo-\ lion
imbricat-: covered with tiles or scales, leot-: smooth
shingled lepi-: scale
impatiens: quickly ripening lepid-: agreeable, pretty, neat
impolit-: unpolished (matt), rough lepide-: scaly
impudic-: shameless, immodest lepis-: scale
in-: prefix meaning not, lacking, on, toward, lepist-: wine pitcher or goblet
intensely, to cause to become lepor-: rabbit or hare
inaurat-: gilded, covered with gold lepr-: rough, scurfy, scaly
incarnat-: flesh-colored lept-: thin, fine, delicate
indie-: pertaining to India or indigo leptonia: fine like a small coin
indigo: dark blue or purple-blue leuc-: white, light
indusi-: garment, overall levi-, levigat-: see laev-, laevigat-
infract-: humble, subdued, uniform liber-, libr-: free
inful-: priest’s cap or hood lign-: wood
infundibul-: funnel ligny-: ash, soot
ino-: fiber ligul-: little tongue
inquinans: polluting, staining limac-, Umax: slug, slime
insign-: illustrious, distinctive, well-marked limb-: border, edge
insuls-: tasteless, unattractive liquiriti-: licorice
integr-: entire, whole, renewed, lith-: stone
intyb-: chicory, endive livid-: livid, lead-colored (an indefinite gray
invers-: inverted, upside down or bluish-gray color)
involut-: rolled up, inrolled loma-: margin, edge
io-, iod-: violet loricat-: armored
irin-: pertaining to iris lubric-: lubricous, slippery, smooth
irpex: harrow, wolfs teeth lucid-: glossy, polished, clear
irradians: radiant, radiating luculent-: clear, beautiful
ischn-: slender, thin, weak lup-: wolf
ithy-, ithys-: straight, erect
908 LATIN AND GREEK DICTIONARY

lurid-: lurid, unclean, dingy, or an indefinite my-: see myo-

color “between purple, yellow, and gray” myc-: fungus, mushroom
— Mcllvaine mycena: ancient term for mushroom
lute-: yellow myo-, myos-: mouse
lyc-, lyco-: wolf myri-: myriad, countless, many
lyo-, lyso-: prefix meaning loose, free, myrm-, myrmec-: ant
dispersed, or dissolving myx-: mucus, slime
M N
macr-: large, long naemat-: with threads
macul-: spot, stain, blotch nan-: dwarf
madid-: moist, soaked nap-: turnip
magn-: great nasc-, nascens: arising, beginning
magnat-: of magnates (dignitaries) nauc-: nut shell
mamm-: nipple, teat, breast nebul-: vapor, cloud, fog, mist, smoke
mappa: napkin necator: killer
marasm-: withered, emaciated nemor-: of the woods, sylvan
marg-, margin-: margin, edge, border neo-: prefix meaning new
marzuol-: pertaining to March neolecta: new selection (?)
mastruc-: sheepskin nid-: nest
maur-: dark, obscure nidor-: fume
maxim-: largest, greatest nidoros-: reeking, with a bad or burnt smell
med-: middle, medium nidul-: little nest
meg-: prefix meaning large or great nidulans: nesting
mela-, melan-, melaen-, melas: black nidus: nest
meleagr-: speckled nigell-: blackish, dark
melle-: pertaining to honey, honey-colored niger, nigr-: black
merd-: dung nit-, nitel-: shining, bright, trim
mer-, meri-: a part nitid-: glossy, spotless
mesenter-: middle intestine niv-: snow
met-, meta-: (1) prefix denoting change (2) nola-: little bell
prefix meaning between, next to, etc. not-: back, buttock
metric-: measure, measuring nub-: cloud, clouds
micr-: small nud-: naked, bare
militar-: soldierlike, occurring in troops
O
miniat-: painted with red lead
minut-: minute oblectabil-: beguiling, delightful
mir-, mirab-: admirable, marvelous obscur-: dark, dusky
mit-, mitis: mild, tender, harmless occidental-: western
mitr-: miter, headdress, cap ochr-, ochrac-: ochre, ochraceous
mokusin: a region in China ocrea-: sheath, boot, greave
moll-: soft, tender ocul-: eye
molyb-: lead or pertaining to lead odon-, odont-: tooth, teeth
monach-: monk or nun officinal-: with pharmaceutical use
mont-: mountain(s) olea-: the olive tree
morb-: disease olens-: smelling (usually a suffix)
morchell-: German term for edible fungus olid-: fragrant, smelly
or morel olig-: little, small, few
morph-: form, shape olivace-: olive-colored
muc-: slime, mucus oil-: pot or jar
mucid-: slimy omphal-: navel (umbilicus)
mucronat-: with a sharp tip, pointed ono-: donkey
multi-: prefix meaning many onust-: full, loaden down
multiplex: many pieces onych-: onyx
mund, mundul-: neat, clean ophi-: snake, serpent
mur-: mouse orbicul-: flat and round (disclike), circular
muricat-: (1) roughened by many short orcell-: small vase
spines (2) pointed oreades: pertaining to mountain nymphs
musca-: a fly or fairies
musci-, muscu-, musco-: moss orellan-: pertaining to mountains
mut-, muta-: change ori-: of the mountains
mutinus-: a phallic deity worshipped by orichalc-: copper-colored
Roman brides, hence a penis oriental-: eastern
LATIN AND GREEK DICTIONARY 909

orth-: straight petal-: petal or leaf

os-: see oss- petasat-: with a wide-brimmed hat, having
osecanus-: whitened like bone a cap on
osm-: smell, scent, perfume, odor peziza: ancient term for a mushroom with
oss-: bone little or no stalk
ostre-: oyster phae-: dark, dusky
ot-: (1) ear (used as suffix or prefix) (2) see phall-: swollen, puffed up, phallic
list of suffixes phallus: rod, phallus
ov-: egg phan-, phaner: appearing, visible
ovat-: egg-shaped, oval phell-: cork
ovin-: ovine (pertaining to sheep) phil-: loving, fond of, friend of
ox-, oxy-: sharp phleb-, phlep-: vein(s)
oxyd-: oxide, oxidizing phlegm-: mucus
oz-: branch phloe-: rind, bark
P phlog-: flame, flame-colored, reddish
phoenic-: red, purple-red, crimson
pachy-: thick
phol-, pholid-: scale
paleace-: chaffy (with small weak scales)
phor-: bearer or carrier, bearing (usually
pallens, pallid-: pale, pallid
a suffix)
palm-: palm (of a hand)
phragm-: palisade, hedge, fence
palud-: swamp, marsh, bog
phyc-: algae, seaweed
palustr-: marshy, pertaining to swamps
phyl-: tribe, race
panaeolus: variegated, all-variegated
phyll-: leaf or leaves (gills)
pann-: rag
phys-: (1) small bladder, bubble (2) growth
panus: tumor
pic-: magpie
papilio-: butterfly
picea-: spruce
papill-: nipple, papilla, pimple pici-: pitch-black (pertaining to pitch)
par-, para-: prefix meaning near, beside, picoa: coated with pitch or tar (black)
related to pier-: bitter, pungent
pare-, parci-: stingy, spare, sparse pict-: painted, streaked, embroidered
pard-: leopard pil-, pile-: (1) usually means pileus or cap
pargamen-: pertaining to parchment (2) pile- can also mean hair (3) pila can
parv-: small, little, pretty, dainty mean ball or bullet
patel-: dish pin-: pine
pauc-: few pinet-: pine wood
pauper-: poor pingu-: (1) fat or grease (2) fat or stout
paxill-: small stake, small stick, peg piperat-: peppery
pectin-: comb pipt-: easily detachable, falling off
ped-, pedes: foot or base piri-: see pyr-
pediades: of the plains, of the soil pis-: pea
pelargon-: pertaining to geranium pistil-, pistill-: pestle
peli-: livid, dark pithy-: wide-mouthed jar
pell-, pelli-: skin plac-: a flat round plate, i.e., flat
pelliculos-: with a well-developed pellicle placid-: mild, gentle
pellucid-: shining through, pellucid, glossy placit-: pleasing
penetrans: penetrating platy-: broad, wide, flat
penio-: fabric
pled-: plaited, twined, twisted
pepl-: (1) skirt, robe, coat (2) gilded pleur-: side, beside, on the side
per-: prefix meaning intensely or very plex: (1) pieces (2) a knitting or interweaving
percomis: very courteous or elegant plicat-: folded, pleated
perd-, perdon: flatulence plinth-: brick, block
perenn-: perennial, lasting, year-round plum-: feather or long hair
perfor-: pierce plumb-: lead or lead-color
peri-: prefix meaning around, all around, plute-: bracket, shed, penthouse, parapet
enclosing pocillator: cup-bearer
perlat-: widespread, enduring (may also be pod-, podi-: foot, base (stalk)
a corruption of word for pearl) pog-, pogon: beard
peronat-: sheathed, booted, rough-booted poli-: gray, hoary
persic-: peach poly-: prefix meaning several, many, much
personal-: masked pom-: fruit tree
pes: foot or base (stalk) ponderos-: heavy, ponderous
pessundat-: ruined
910 LATIN AND GREEK DICTIONARY

popul-: poplar, cottonwood, aspen racem-: branched like a grape stalk

por-: pore, pores rack-: misspelling of rhac-
porphyr-: (1) purple (2) reddish-brown or radi-: ray, spoke, plate
russet radians: (1) with rays or spokes (2) radiant
porrigens: extending forward, projecting radio-: root
horizontally, spreading, stretching out radul-: scraper
portentos-: portentous, prodigious ram-: branch
prae-: prefix meaning before or very raphan-: radish
praeclare-: very clearly rasil-: shaved, scraped, worn smooth
praecox: early, premature recis-: cut off, cut back
pras-: leek, leek-green reg-: king, royalty
prat-: meadow, meadows renidens: shiny
pre-: see prae- repand-: folded backward, turned up
privignus: step son resim-: turned up or bent back
pro-: prefix meaning before ret-: net
procer-: lofty reticulat-: reticulate (netted)
prolix-: stretched, elongated rhac-: rag
prote-: a sea god who could change shape at rhach-: spine or backbone
will, i.e., protean rhac odes: ragged, shaggy
proto-: prefix meaning primary or giving rhe-: flow, let flow, exuding, flowing
rise to rhen-: Rhine
proxim-: nearest, next rhiz-: root
pruinos-: with a fine bloom, frosted rhod-: rose, rose-colored (pink or red)
prun-: plum rhopal-: club, stick
psall-: ring or collar, fringed or curb rim-: crack, fissure
psamm-: sand ringens: gaping
psathyr-: fragile, brittle ripa-: stream bank
pseud-: prefix meaning deceptive, similar rivul-: channel, groove, stream
to, easily confused with, false (often used robinia: locust tree
before the name of a similar mushroom) robust-: robust, stout, strong
psil-: naked, bare, hairless, smooth ros-: rose
psitt-: parrot rostr-: beak, snout
pter-: wing, membrane rot-: wheel
p ter id-: fern rotul-: little wheel, pinwheel
pteryg-: wing, fin, membrane rub-, ruber: red
ptych-: fold or layer, leaf rube/-: almost red, reddish
pub-: hair rubig-: rusty, reddish-brown
pubescens: downy, pubescent rubr-: red
pudic-, pudor-: modest, rosy rud-: uncombed, made of untreated wool,
puellaris: pertaining to girl, i.e., pretty rough
pulch-, pulcher-, pulchr-: beautiful ruf-: rufous (reddish)
pulver-: powder, dust rug-: wrinkle
punct-: fine spot, dot, or puncture russ-: (1) reddish (2) Russian
punctat-: punctate (finely spotted or dotted) rutil-: ruddy or warm red
punic-: red or purple-red, pomegranate-red, S
garnet-red
pur-: clean, pure sacc-: sac, sack
purpur-: purple sacchar-: sugar, sugary, sweet
pus-: (1) foot, base, or stalk (2) also seepusil- saep-: hedge
pusil-, pusill-: very small, little, weak saev-: savage
pustul-: pustule, pimple salic-, salix: willow
pycn-: thick, dense, compact, strong salmon-: salmon-colored
pyr-: (1) pear (especially pyri-) (2) fire sandarac-: flesh-color
(especially pyro-) sangui-, sanguin-: bloody, blood-red
pyram-, pyramid-: pyramid, pyramidal sapine-: fir or pine
pyren-: kernel, pit (of a fruit) sapon-: soap
pyrr-, pyrrh-: flame-colored, reddish sarc-: flesh, fleshy
pyx-, pyxid-: a small box sardoni-: very bitter, acrid
sarx: flesh
QR sax-: stone, rock
quadr-: four, fourfold (usually a prefix) scaber, scabr-: rough, scurfy, pertaining to
querc-: oak scaber(s)
LATIN AND GREEK DICTIONARY 911

scamb-: bow-legged (i.e., curved or bent) spiss-: thick, massive, compact

scaur-: with swollen ankle or foot splendens: polished, glittering, resplendent
schiz-: split spong-: spongy
sciss-: cut spor-: spore, seed
scler-: hard spret-: despised, scorned, rejected
sclerotoid-: pertaining to a sclerotium(tuber spurn-: foam, froth
or knot of tissue) squam-: a scale
scolec-: worm squamul-: small scale
scord-, scorodon-: garlic squarros-: squarrose (with upright scales,
scrobiculat-: scrobiculate (pitted) rough, scurfy)
scut-: shield st ell-: star
scutel-: dish ster-: hard
scyph-: cup sterquil-: dung heap, dung pit
seb-, sebac-: tallow, grease stict-: marked, lined, dappled, dotted
sec-: an enclosure or nest stip-: stalk or stem (stipe)
secund-: conforming, following stiptic-: close, dense (also see styptic-)
sejunct-: disjoined, separated, severed stom-: mouth
semi-: prefix meaning half or less than stramin-: straw-colored
semital-: of footpaths, byways striat-: striate (finely furrowed or lined)
semot-: remote strict-: very straight, erect (usually narrow)
sep-: see saep- strigos-: strigose (with coarse flattened, rigid
separans: distinctive hairs or bristles)
separat-: separated, separate, apart strobil-: pertaining to pine cone, or to
septentrional-: northern (from a northern something twisted
constellation) strom-: bed, mattress
sepulchr-: tomb stroph-: (1) belt or pectoral band (2) a turning
sepult: buried or twisting
sequoia-: redwood styptic-: astringent (also see stiptic-)
ser-: (1) whey, serum (2) to fasten or bind su-: pig, swine
seri-: silk suav-: sweet, agreeable
serial-: in rows sub-: common suffix meaning (1) somewhat
serotin-: late or almost (2) below or under
serp-: snake subtil-: fine, delicate, minute
ser rub: finely serrate (toothed like a saw) succ-: juice, sap
sibir-: Siberia sudor-: sweat
sicc-: dry sudorific-: sweat-producing
silv-: woodland, forest suec-: Sweden
sim-: flattened or concave suillus: an ancient term for fungus (derived
simbl-: beehive from word for swine)
sin-: curve, bay sulc-: furrow or groove
sinap-: mustard summ-: summit, top
sindon-: muslin surrect-: rising, erect
sinopic-: reddish-ochre sylv-: woodland, forest
sinu-, sinus: curve, bend, bay synaps-, synapt-: joined together
sinuat-: sinuate (notched gills)
T
sistr-: rattle
smaragd-: emerald, green tabac-: tobacco (color, odor, etc.)
som-: body tabescens: decomposing
sordid-: sordid (dingy, filthy, foul) tabid-: decomposed
soror-: pertaining to sister tard-: slow
spad-, spadic-: (1) bright brown (the color of tect-: roof, cover
a fresh date) (2) shaped like a palm frond telamon-: belt or supporting band; web
or spade tenac-, tenax: tenacious
sparassis: torn to pieces, lacerated tener-: tender, delicate
spathul-: little spade or blade, spatula tenu-, tenui-: thin
specios-: beautiful, showy tephr-: ashes, ash-colored, gray
spectabil-: notable, remarkable, visible terfez-: Arabic word for truffle
terr-: earth
sperm-: seed (spore)
sphaer-: sphere, round tessulat-: mosaic-like, checkered
sphagn-: sphagnum, moss testace-: testaceous (brick-colored)
sphec-: (1) wasp (2) lichen thee-: case or box
sphinct-: a tightening, tightly bound thej-, thei-: sulfur, brimstone
spin-, spinul-: spine, thorn
912 LATIN AND GREEK DICTIONARY

thraust-: easily crumbled, fragile, brittle v

thrix: hair
vacc-: cow
thuj-, thy-: cedar (arbor-vitae, not Cedrus)
vaccini-: huckleberry
tigr-: tiger
vagin-: sheath, scabbard
tinct-: tinted, dyed
valens: powerful, strong, robust
toga-: toga (robe)
valid-: powerful, robust
toment-: wool or hair
vaporari-: pertaining to humid places
torminos-: causing dysentery
vari-: change, changing
toros-: muscular, i.e., bulging or swollen
variat-: variable
torqu-: necklace, collar
variegat-: variegated, multicolored
tortil-: twisting, winding
vas-: vessel, vase
torul-: (1) tuft of hairs (2) bulge or swelling
vel-: .veil
torv-: wild, savage
veil-: fleece, wool
tost-: toasted
vellere-: covered with fleece, woolly
trabe-: beam, timber
velut-: velvet, velvety
trachy-: rough
ven-: vein
trag-: goat
venen-: poison
tram-, trama: (1) the woof (as in weaving)
venet-: colored like sea water
(2) something thin
ventr-: belly
trametes: one who is thin
ventricos-: ventricose(swollen, bulging)
tremell-: trembling
venust-: handsome, elegant
tri-: prefix meaning three, thrice
verm-: worm
trich-: hair(s), fiber(s)
vermicul-: little worm
trivial-: common, vulgar
vern-: spring, of spring
trullisat-: plastered
vernicos-: varnished
trunc-: trunk (as of a tree) verp-: rod, penis
truncat-: truncate (appearing chopped off) verruc-: wart
tsuga-: hemlock vers-: changing
tub-, tuba-: trumpet or tube versicolor: multicolored
tuber: (1) a tuber (2) a bump or knob vesc-: (1) edible (2) feeble
tul-, tulo-: see tyl- vesicul-: small bladder, blister
tumid-: swollen vestit-: dress, attire
tunic-: tunic, garment
veternos-: weak, lethargic
turbinat-: shaped like a top
via-: way, biway
turmal-: a troop
vibec-, vibic-: bruise
turp-: vile, ugly
viet-: wrinkled, wilted, shrunken
tyl-, tylo-: (1) swelling, bump, cushion,
vill-: shaggy hair
pillow (2) knob or callus on a club
villatic-: rustic, rural
typh-: (1) smoke, cloud (2) a plant used for
vin-, vini-: wine (especially red wine)
stuffing beds
viol-: violet
tyr-: cheese
vir-, virens, virid-: green
U virg-: stripe, streak
ud-: wet, moist, damp virgin-: virgin (i.e., white or pure)
uligi-: moisture virid-: greenish
uliginos-: full of moisture; boggy, swampy viros-: poisonous
ulm-: elm vise-, viscid-: viscid (sticky)
umbell-: umbrella or sunshade vitell-: egg yolk
umbilic-: umbilicus (navel) vitil-: coiling, twining
umbon-: knob (derived from a word for volemus: filling the palm of the hand, or
shield) flowing enough to fill the hand
umbr-: dark, shade volva-: wrapper, womb (volva)
ungu-: grease vor-: devour, devourer, eater
ungul-: hoof vulgar-: common, known, usual
uni-: prefix meaning one, single vulp-: fox
urbic-: urban, of the city WXYZ
und-, undul-: wave
xanth-: yellow
urens-: burning
xer-: dry
urs-: bear
xerampelin-: colored like a dry vine leaf
ustal-: russet (derived from word for burnt)
xyl-, xylon: wood, wooden, woody
uter-, utri-: wine skin, womb
zanth-: see xanth-
uvid-: damp, humid
zon-: band or zone (usually concentric)
zyg-: yoked (connected, joined, attached)
 913

GLOSSARY
acanthocytes needle-like crystalline de¬ boletivore one with an inordinate fond¬
posits on mycelium of certain mushrooms ness for eating boletes (p. 546)
acrid taste burning or peppery brown rot carbonizing decay
acute pointed or sharp buff an indefinite pale color: pale dull
adnate gills attached broadly to stalk (p. yellow or very pale tan
17) bulbous stalk with an enlarged base (p.
adnexed gills attached narrowly to stalk 17)
(P- 17) BUM Boring Ubiquitous Mushroom
agaric a mushroom with gills button a young fruiting body before it has
alveolate surface of spore or cap with opened up
broad pits (p. 842) butt rot a rot confined to the base or roots
amyloid staining blue, gray, or black in of the tree
iodine solution (e.g., Melzer’s reagent)
cap the caplike part of the fruiting body
anastomosing gills connected by cross¬
which supports the spore-bearing surface
veins
capillitium modified threadlike, often-
angular spores 4- to 6-sided, with corners
branched hyphae enmeshed in the spore
or angles
mass of many Gasteromycetes
annulus ring or collar of tissue on stalk
capitate furnished with a rudimentary
formed by ruptured veil
cap or head
apex top
carbonizing decay a cellulose-decom¬
apical at or near the top
posing decay
apical pore in certain Gasteromycetes,
carminophilous see siderophilous
the mouth at the top of the spore case
cartilaginous with the texture of carti¬
through which spores are released; in
lage; tough or rindlike, not fleshy
spores, a germ pore
cellular tissue composed of round or pear-
apiculate furnished with an apiculus
shaped cells (p. 19)
apiculus short projection on either end of
cellulose a compound composed of glu¬
a spore
cose units; it is a major constituent of
apothecium the fruiting body of an
wood and of plants’ cell walls
Ascomycete in which the asci are ar¬
cespitose (caespitose) tufted or clus¬
ranged in an exposed hymenium
tered
appendiculate margin of cap fringed or
chlamydospores thick-walled asexual
adorned with veil remnants or other tissue
spores formed by breaking up of hyphae
appressed flattened down or pressed
chrysocystidia cystidia with a highly re¬
against
fractive golden content as seen in KOH
areolate cap surface cracked into plaques
clamp connection a small bump, loop,
or blocks (like dried mud)
or swollen area formed at the cross-wall
ascocarp fruiting body of an Ascomycete
between hyphae in the process of cell
ascus (pi., asci) saclike mother cell in
division
which spores of Ascomycetes are formed
clamps mycological jargon for clamp con¬
(pp. 4, 782)
nections
autodigestion self-digestion
class a grouping of related orders
basal at or near the base clavate club-shaped
basidiocarp fruiting body of a Basidio- club-shaped stalk thickened noticeably
mycete toward base (p. 17); basidia thickened
basidium (pi., basidia) a cell, usually toward spore-bearing end
club-shaped, on which spores of Basidio- collarlike type of volva formed when the
mycetes are formed (p. 4) universal veil breaks around the circum¬
bolete a fleshy mushroom with tubes on ference of the cap, forming a collar or free
underside of cap rim near base of stalk (p. 264)
914 GLOSSARY

columella internal stalk; a sterile column divergent gill hyphae projecting down¬
of tissue projecting into the spore mass of ward (away from cap) as seen in cross-
certain Gasteromycetes section; diverging from the center (p. 19)
complex stalk folded or convoluted in division a group of related classes
cross-section; a group of closely related dry cap or stalk neither viscid nor hygro-
species or forms phanous
compound fruiting body with more than duplex flesh of two distinct textures
one cap and/or stalk arising from a
eccentric stalk off-center
common base
echinulate spores spiny like a sea urchin
conidia asexual spores formed by the
egg the button stage of a mushroom with
pinching off of hyphae
a universal veil (e.g., Amanita); also,
conifer a cone-bearing tree; one that has
spore-containing capsule in a bird’s nest
needles like a pine or redwood, or scales
fungus
like a cypress
elliptical spores rounded at both ends
conk a woody, usually perennial poly¬
with sides curved slightly
pore
endemic native to a region and generally
context the flesh of the cap or stalk
restricted to it
convergent gill hyphae projecting up¬
endoperidium the inner layer of the
ward toward the cap as seen in cross-
spore case in puffballs and their allies
section; converging toward the center (p.
entire gills with even edges; not serrated
19)
or toothed
convex cap rounded or domed (p. 17)
epigeous growing on or above the ground
convoluted wrinkled like a brain
epiphragm the thin membrane covering
coprophilous dung-loving; growing on
the nest of many young bird’s nest fungi
dung
equal stalk of more or less uniform thick¬
cortina a veil with a silky or cobwebby
ness throughout (p. 17)
texture
ericaceous pertaining to the heath family
cuticle the skin or surface layer of the cap,
erumpent fruiting body erupting through
more specifically one differentiated from
the ground but scarcely rising above it
the flesh
evanescent transitory; disappearing
cystidium (pi., cystidia) specialized ster¬
exoperidium the outer layer of the spore
ile cells projecting from the gills, tubes,
case in puffballs and their allies
cap, or stalk (p. 19)
expanded cap fully developed; spread
daedaloid pores elongated or meander¬ out
ing like a labyrinth; mazelike
fairy ring a circle or arc of mushrooms (p.
decurrent gills running down the stalk (p.
7)
17)
family a group of related genera
decurved margin of cap curved down¬
farinaceous odor like fresh meal or
ward
cucumber
delignifying decay a lignin and cellulose-
fetid (foetid) ill-smelling
decomposing rot
fibril a fine fiber or hair
deliquescing gills dissolving into a fluid;
fibrillose covered with or composed of
liquefying
fibrils
depressed cap concave, i.e., the center
fibrous composed of tough, stringy tissue
sunken below the level of the margin (p.
filamentous tissue composed of thread¬
17)
like cells or filaments (p. 19)
dextrinoid staining brown or red-brown
flaccid lacking firmness; flabby
in iodine solution (e.g., Melzer’s reagent)
flesh the tissue of the cap or stalk; the
dichotomous forking or dividing in pairs
meaty portion
differentiated different; not the same
fleshy having substance (e.g., a fleshy
throughout.
fungal fructification) or soft as opposed to
disc center of the cap
tough
distant gills widely spaced
GLOSSARY 915

floccose woolly or cottony; dry and habitat natural place of growth

loosely arranged hardwood used here in a broad sense to
fluted stalk ribbed; with longitudinal denote any tree that is not a conifer
ridges heart rot a rot of the heartwood
free gills not attached to stalk (p. 17) homogeneous the same throughout; not
friable easily crumbling differentiated
Friesian pertaining to Elias Fries (the humus decaying organic material in or on
“father” of mushroom taxonomy) and to soil
his classification system based on macro¬ hyaline spores colorless under the micro¬
scopic features scope
fructification fruiting body hygrophanous cap surface changing co¬
fruiting body the reproductive structure lor markedly as it loses moisture; usually
of a fungus; a mushroom watery-looking when wet and opaque
fulvous fox-colored when dry
fungophile one who loves fungi hygroscopic sensitive to moisture
fungophobe one who fears fungi hymenium a layer or palisade of spore¬
fungi (pi., fungi) any member of a large bearing cells
group of organisms that lack chlorophyll, hypha (pi., hyphae) a threadlike fungal
reproduce by means of spores, and have a cell, the basic structural unit of any
filamentous vegetative phase mushroom
funiculus the cord attaching peridiole to hypogeous fruiting underground
the “nest” of certain bird’s nest fungi
incurved cap margin curved inward to¬
furfuraceous scurfy; roughened by mi¬
ward stalk
nute particles
indeterminate growth that ceases and
fuscous an indefinite dark smoky color,
begins anew according to weather con¬
sometimes with a slight violet tinge
ditions, frequently creating a zoned or
fusiform spores spindle-shaped; elonga¬
lumpy cap
ted and tapering from the middle to both
indusium netlike skirt in certain stink-
ends
horns
gastroid like a Gasteromycete, i.e., not inferior annulus located near base of stalk
forcibly discharging its spores innate a part of; not superficial or easily
gelatinous jelly-like in consistency or ap¬ removed
pearance inner veil see partial veil
generic pertaining to genus inrolled cap margin curved in toward gills
genus (pi., genera) a group of closely and rolled up; tucked under
related species interwoven gill hyphae entwined or tan¬
germ pore a soft spot in wall of certain gled as seen in cross-section; not forming
spores, through which initial germination a regular pattern (p. 19)
takes place
JAR Just Another Russula
gills spore-producing blades on underside
of an agaric KOH chemical symbol for potassium
glabrous bald hydroxide (potash)
glandular dots resinous spots or smears
lamellae the gills
on stalk of certain boletes (Color Plate
lamellulae short gills that extend only
120)
part way to stalk
gleba the spore mass of a Gasteromycete
lateral stalk attached to side of cap
globose spherical
latex a juice or milk, as in Lactarius
glutinous slimy or very sticky
LBJ Little Brown Job (p. 33)
granulose covered with granules
LBM Little Brown Mushroom(pp. 32-33)
gregarious growing close together but
leathery tough, pliant, not easily break¬
not in clusters
ing
lignicolous wood-inhabiting
916 GLOSSARY

lignin a major constituent of wood oblong spores elongated, with approx¬

lubricous somewhat slippery or greasy to imately parallel sides
the touch but not viscid or slimy obtuse blunt; not pointed
lumper a taxonomist with a broad species ochraceous yellow or yellow-orange with
or genus concept a brownish tinge
lutescent becoming yellow order a group of related families
ornamentation any discontinuity (warts,
macroscopic discernible without a micr¬
ridges, etc.) on the surface of the cap,
oscope
stalk, or spore
margin the edge of the cap or gills
outer veil see universal veil
marginate gills with edges differently
oval spores or cap egg-shaped
colored than faces
marginate-depressed bulb with a raised pallid very pale; an indefinite whitish
rim color
mealy having a granular appearance or parallel gill hyphae arranged more or less
smelling like fresh meal parallel to each other as seen in cross-
median ring at or near middle of stalk section (p. 19)
Melzer’s reagent an iodine test solution: paraphyses unspecialized sterile cells (p.
44 parts water (by weight), 3 parts 782)
potassium iodide, 1 part iodine, and 40 parasitic feeding on another living or¬
parts chloral hydrate (poisonous!) ganism
membranous membranelike; skinlike partial veil protective tissue stretching
metagrobolizing puzzling, bewildering, from the stalk to the cap margin in many
confounding, confusing, perplexing young mushrooms
micrometer see micron pedicel a slender stalk
micron microscopic unit of measure equal¬ pellicle a viscid skin that usually peels
ing 0.001 mm easily
microscopic discernible only with a mi¬ pendant hanging down; also, annulus
croscope skirtlike
montane of the mountains percurrent columella or stalk extending
mushroom the fruiting body of a fungus, through the spore mass to the top of the
especially one which has gills; also, a fruiting body
fungus that produces a fleshy fruiting peridiole small spore capsule in certain
body Gasteromycetes
mushrump a hump in the humus caused peridium the wall of the spore case in
by a developing mushroom many Gasteromycetes
mycelium a complex network of hyphae; perithecium a flask-shaped “nest” of asci
the vegetative portion of a fungus (in the Pyrenomycetes)
mycologist one who studies fungi peronate sheathlike; also, a sheathing an¬
mycology the study of fungi nulus or veil
mycophagist one who eats mushrooms persistent persisting; not evanescent
mycophile one who loves mushrooms phenolic odor reminiscent of phenol
mycophobe one who despises or fears pileate with a pileus or cap
mushrooms pileus the cap of a mushroom
mycorrhiza a mutually beneficial rela¬ pip-shaped shaped like an apple seed
tionship between a fungus and the plane cap having a flat surface (p. 17)
rootlets of a plant (especially trees), in pocket rot a rot producing hollow poc¬
which nutrients are exchanged kets in a tree
polymorphic with many shapes or forms
naked without hairs or other tissue
polypore any of a large group of mostly
nodulose spores covered with small
tough, wood-inhabiting fungi which bear
bumps or nodules
their spores in pores
notched gills abruptly adnexed, as though
pores the mouths of the tubes in boletes
a small wedge-shaped piece of tissue had
and polypores
been removed at juncture to stalk (p. 17)
poroid resembling pores
GLOSSARY 917

potato chip conditions see footnote on septate partitioned; with one or more
p. 35 cross-walls
pruinose powdery or appearing finely serrated gill edges toothed like a saw
powdered; dandruffy sessile lacking a stalk
pseudocarp something that resembles a setae pointed, elongated, thick-walled
mushroom but isn’t sterile cells
pseudorhiza a rootlike extension of the sheathlike annulus or veil sheathing the
stalk; a “tap root” stalk (p. 312)
pubescent minutely hairy; downy siderophilous basidia with granules that
pulverulent appearing powdered darken when heated in acetocarmine
punctate dotted with minute scales or simple fruiting body unbranched, not
points compound; stalk not complex
putrescent readily decaying sinuate gills notched (p. 19)
sinuous crooked or curved
radially arranged pores or pits arranged
skirtlike annulus hanging down like a
in rows which normally radiate like the
skirt (p. 312)
spokes of a wheel
sordid dingy or dirty in appearance
recurved curved up and back
spawn the mycelium
resupinate lying flat on substrate; with¬
species (sp.) a particular kind of or¬
out a stalk or well-defined cap
ganism; the fundamental unit of tax¬
reticulate stalk marked with lines crossed
onomy
like the meshes of a net; netted (p. 489);
sphaerocyst a round or swollen cell in
spores with a network of ridges
certain mushrooms (e.g., Russula, Lac-
rhizomorph a rootlike or stringlike bun¬
tarius)
dle of mycelial hyphae
spines pendant spore-bearing “teeth” in
ring see annulus
the teeth fungi
rivulose marked by riverlike lines
splitter a taxonomist with a narrow genus
rufescent becoming reddish
or species concept
rufous brownish-red
spore the reproductive unit of a fungus,
rugose wrinkled
usually a single cell
saclike volva shaped like a sack, pouch, spore case the large chamber that holds
or cup (p. 264) the spores in puffballs and related fungi
saprophytic feeding on dead or decaying (Gasteromycetes)
matter squamose furnished with scales; scaly
sap rot decay of the sapwood squamulose furnished with small scales
scabers tufted hairs or short projecting stalk the stemlike structure that supports
scales on stalk (Color Plates 145-147) the cap in most mushrooms
scabrous roughened by scabers stellate star-shaped; with rays
scales pieces of differentiated tissue on sterigma (pi., sterigmata) prong on the
the cap or stalk, often of a different color basidium on which a spore forms
than background sterile not producing spores; infertile
scaly furnished with scales; volva with sterile base the sterile chambered base
concentric rings or scales at base of stalk beneath the spore mass in many puffballs
(P- 264) stipe the stalk
sclerotium a knot or tuber of hyphae, stipitate furnished with a stalk or stipe
usually underground, in certain mush¬ striate marked by lines; finely furrowed
rooms; a resting stage that fruits periodi¬ stuffed stalk stuffed with a pith
cally subdistant gills fairly well spaced
scrobiculate stalk pitted with conspicu¬ subperonate somewhat or slightly pero-
ous spots nate
seceding gills breaking away from stalk substrate the material to which a fruiting
as the cap expands body is attached
secotioid resembling Secotium, a genus subtomentose finely or somewhat to-
of gastroid agarics; reminiscent of an mentose
undeveloped or aborted agaric
918 GLOSSARY

Silicate conspicuously furrowed; deeply umber a deep dull brown

striate umbilicate cap with a navel-like central
superficial scales or warts easily remova¬ depression (p. 17)
ble; not innate umbo a knob or bump at center of cap (p.
superior ring located at or near top of 5)
stalk umbonate cap furnished with an umbo
(P- 17)
tacky slightly sticky but not truly viscid
universal veil a protective layer of tissue
tawny the color of a lion
that envelops all or most of the young
taxonomy the classification of organisms
fruiting body of certain mushrooms
to reflect their natural relationships
tenacious tough veil a protective tissue (see partial and
terrestrial growing on the ground universal veils)
tidepool conditions see footnote on p. ventricose stalk swollen at or near the
35 middle
toadstool a mushroom, especially one vinaceous the color of fine red wine or
that is poisonous slightly paler
tomentose covered with soft hairs viscid slimy or sticky to the touch, at least
top rot a rot confined to the top of a tree when moist
translucent-striate cap with the gills volva remnants of the universal veil at the
showing through to give a striate (lined) base of the stalk in certain mushrooms,
appearance usually in the form of a sack, collar, or
truncate appearing chopped off or flat¬ series of concentric rings or scales (p. 264)
tened at one end volval patch a large patch of universal
trunk rot decay found throughout the veil tissue on the cap
trunk of a tree
warts small pieces of universal veil tissue
tuberculate with low bumps
deposited on the cap; also, any wartlike
tuberculate-striate striate with small
scale or protuberance
bumps
white rot a delignifying decay
tubes the tubelike structures on the under¬
side of boletes and polypores, in which YAM Yet Another Mycena
spores are produced
zonate cap or flesh concentrically zoned
turbinate top-shaped

Leucopaxillus amarus, a common but bitter-tasting brown-capped agaric. N ote how some specimens
have a ribbed cap margin. Gills and flesh are white. See p. 168 for details.
 919

BIBLIOGRAPHY:
Suggested Readings
and Primary References
N on-Technical Literature
(Note: Publications dealing with specific groups of fungi are listed under
“Technical Literature,” even though they may be written for amateurs)

Field Guides
Biek, David (1984). The Mushrooms of Northern California. Redding, Calif: Spore Prints.
Bowers, Jeannette & David Arora (1984). Mushrooms of the World Coloring Book.
New York: Dover.
Guzman, Gaston (1978). Hongos. Mexico City: Limusa.
Kauffman, C.H.(1918). The Gilled Mushrooms (Agaricaceae) of Michigan and the Great
Lakes Region. New York: Dover (1971 reprint).
Krieger, Louis C. (1936). The Mushroom Handbook. New York: Dover (1967 reprint).
Lange, Morton& F.B. Hora (1963). A Guide to Mushrooms and Toadstools. NewYork:
E.P. Dutton.
Largent, David, Harry Thiers, Roy Watling, & Daniel Stuntz (1973-). How to Identify
Mushrooms to Genus (5 vols.). Eureka: Mad River Press.
Lincoff, Gary (1981). The Audubon Society Field Guide to North American Mushrooms.
New York: Alfred Knopf.
Mcllvaine, Charles & Robert Macadam (1902). One Thousand American Fungi.
NewYork: Dover(1973 reprint).
McKenny, Margaret & Daniel Stuntz (1971). The Savory Wild Mushroom. Seattle:
University of Washington Press.
Miller, Orson K. (1972). Mushrooms of North America. New Y ork: E.P. Dutton.
Moser, Meinhard (1983). Keys to Agarics and Boleti. England: Roger Phillips.
Phillips, Roger (1981). Mushrooms and Other Fungi of Great Britain and Europe.
London: Pan Books.
Ramsbottom, John (1953). Mushrooms and Toadstools. London: Collins.
Rinaldi, Augusto & Vassili Tyndalo (1974). The Complete Book of Mushrooms. New
York: Crown.
Smith, Alexander (1949). Mushrooms in Their Natural Habitats. New York: Hafner
(1973 reprint).
Smith, Alexander (1975). A Field Guide to Western Mushrooms. Ann Arbor: University
of Michigan Press.
Smith, Alexander, Helen V. Smith, & Nancy Smith Weber (1979). How to Know the
Gilled Mushrooms. Dubuque: William Brown.
Smith, Alexander, Helen V. Smith, & Nancy Smith Weber (1981). How to Know the
Non-Gilled Mushrooms, 2nd edition. Dubuque: William Brown.
Smith, Alexander & Nancy Smith Weber (1980). The Mushroom Hunter’s Field Guide
(revised). Ann Arbor: University of Michigan Press.
Weber, Nancy Smith & Alexander Smith(1985). A Field Guide to Southern Mushrooms.
Ann Arbor: University of Michigan Press.
Wright, Greg (1981). Key to the Gilled Mushrooms and Boletes of Southern California.
Published by the author.
920 BIBLIOGRAPHY

Cookbooks
Grigson, Jane (1975). The Mushroom Feast. New York: Alfred Knopf.
Puget Sound Mycological Society (1973). Oft Told Mushroom Recipes (republished as
Wild Mushroom Recipes). Seattle: Pacific Search.

Poisonous & Hallucinogenic Mushrooms

Duffy, Tom & Paul Vergeer (1977). California Toxic Fungi. Mycological Society of
San Francisco.
Lincoff, Gary & D.H. Mitchel (1977). Toxic and Hallucinogenic Mushroom Poisoning.
New York: Van Nostrand Reinhold.
Stamets, Paul(1978). Psilocybe Mushrooms and Their Allies. Seattle: Homestead Books.

Mushroom Cultivation
Harris, Bob (1976). Growing Wild Mushrooms. Berkeley: Wingbow Press.
Stamets, Paul & J.S. Chilton (1983). The Mushroom Cultivator: A Practical Guide to
Growing Mushrooms at Home. Olympia: Agarikon Press.

Miscellaneous
Bo, Liu & Bau Yun-sun (1980). Fungi Pharmacopoeia (Sinica). Oakland, Calif: Kinoko
Company.
Jaeger, Edmund C. (1955). A Sourcebook of Biological Names and Terms. Springfield,
Illinois: Charles C. Thomas.
Rice, Miriam & Dorothy Beebee (1980). Mushrooms for Color. Eureka: Mad River Press.
Wasson, R. Gordon (1968). Soma: The Divine Mushroom of Immortality. New York:
Harcourt Brace Jovanovich.
Wasson, Valentina & R. GordonWasson(1957). Mushrooms, Russia and History (2 vols).
New York: Pantheon.

Technical Literature

General
Bigelow, H. & H. Thiers, editors (1975). Studies on Higher Fungi. Germany: J. Cramer.
Miller, O.K. & D.F. Fa.rr(\915).AnIndexof the Common Fungi ofN orth America. Liech¬
tenstein: J. Cramer, Bib. Myco. 44.
Petersen, R., editor (1971). Evolution in the Higher Basidiomycetes: An International
Symposium. Knoxville: Univ. of Tennessee Press.
Richardson, M. & R. Watling (1968). Keys to Fungi on Dung. Bull, of the Brit. Myco.
Soc.2: 18-43.
Saylor, H., P. Vergeer, D. Desjardin, & T. Duffy. California Mushrooms 1970-1980:
Fungus Fair and Foray Collections. Myco. Soc. of San Francisco.
Shaffer, R.L. (1968). Keys to Genera of Higher Fungi, 2nd edition. Ann Arbor: Univ. of
Michigan Biological Station.
Singer, R. (1975). The Agaricalesin Modern Taxonomy, 3rd edition. Germany: J. Cramer.
Snell, W.H. & E.A. Dick (1957). A Glossary of Mycology. Cambridge, Mass: Harvard
Univ. Press.
Watling, R. & A.E. Watling (1980). A Literature Guide for Identifying Mushrooms.
Eureka: Mad River Press.

Russulaceae& Hygrophoraceae
Hesler, L.R. & A.H. Smith (1963). North American Species of Hygrophorus. Knoxville:
Univ. of Tennessee Press.
SUGGESTED READINGS AND PRIMARY REFERENCES 921

Hesler, L.R. & A.H. Smith (1979). North American Species of Lactarius. Ann Arbor:
Univ. of Michigan Press.
Largent, D. (1985). The Agaricales of California, 5: Hygrophoraceae. Eureka: Mad River.
Methven, A. (1985). New and Interesting Species of Lactarius from California. Mycologia
77: 472-182.
Shaffer, R. (1962). The Subsection Compactae of Russula. Brittonia 14: 254-284.
Shaffer, R. (1964). The Subsection Lactarioideae of Russula. Mycologia 56: 202-231.
Shaffer, R. (1970). Notes on Subsection Crassotunicatinae of Russula. Lloydia33:49-96.
Shaffer, R. (1972). North American Russulas of Subsection Foetentinae. Mycologia 64:
1008-1053.

T richolomataceae
Baroni, T. (1983). Tricholoma manzanitae—A New Species from California. Mycotaxon
18:299-302.
Bigelow, H.E. (1970). Omphalina in North America. Mycologia 62: 1-32.
Bigelow, H.E. (1982). North American Species of Clitocybe, Part 1. Liechtenstein:
J. Cramer.
Desjardin, D.E. (1985). New Marasmioid Fungi from California. Mycologia 77: 894-902.
Gilliam, M.S. (1975). New North American Species of Marasmius. Mycologia 67:817-844.
Gilman, L. & O.K. Miller (1977). A Study of the Boreal, Alpine, and Arctic Species of
Melanoleuca. Mycologia 69:927-951.
Hailing, R.E.(1983). The Genus Collybiain the Northeastern United States and Adjacent
Canada. New York Bot. Gar. Myco. Mem. 8 (Germany: J. Cramer).
Lennox, J.W. (1979). Coilybioid Genera in the Pacific Northwest. Mycotaxon9: 117-231.
Miller, O.K. & L. Stewart(1971). The Genus Lentinellus. Mycologia 63: 333-369.
Mueller, G. (1984). New North American Species of Laccaria. Mycotaxon 20: 101-116.
Singer, R. & A.H. Smith (1943). A Monograph of the Genus Leucopaxillus. Pap. Mich.
Acad. Sci. 28: 85-132.
Smith, A.H. (1947). North American Species ofMycena. Ann Arbor: Univ. of Mich. Press.
Smith, A.H. (1960). Tricholomopsis in the Western Hemisphere. Brittonia 12: 41-76.
Smith, A.H. (1979). The Stirps Caligata of Armillaria in North America. Sydowia 8:
368-377.
Smith, A.H. & R. Singer (1945). A Monograph of the Genus Cystoderma. Pap. Mich.
Acad. Sci. 30: 71-124.
Thiers, H.D. & W.J. Sundberg(1976). Armillaria in the Western United States. Madrono
23: 448-153.

Entolomataceae & Pluteaceae

Hesler, L.R. (1967). Entoloma in Southeastern North America. Germany: J. Cramer.
Homola, R.L. (1972). Section Celluloderma of the Genus Pluteus in North America.
Mycologia 64: 1211-1247.
Largent, D. (1970-1974). Studies in the Rhodophylloid Fungi I: Madrono 21: 32-39;
II: Mycologia 62: 437-452.
Largent, D. (1971-). Rhodophylloid Fungi of the Pacific Coast I: Brittonia 23: 238-245;
II: Northwest Sci. 46: 32-39; III: Mycologia 66: 987-1021.
Largent, D. (1977). The Genus Leptonia on the Pacific Coast of the United States.
Germany: J. Cramer, Bib. Myco. 55.
Shaffer, R. (1957). Volvariella in North America. Mycologia 49: 545-579.
922 BIBLIOGRAPHY

Amanitaceae, Lepiotaceae, & Agaricaceae

Bas, C. (1969). Morphology and Subdivision of Amanita and a Monograph on its Section
Lepidella. Persoonia5: 285-579.
Freeman,- A.E.H. (1979). Agaricus in Southeastern United States. Mycotaxon8: 50-118.
Isaacs, B.F.(1963). A Survey of Agaricus in Washington, Oregon, and California. Master’s
Thesis, University of Washington; unpublished.
Jenkins, D.(1977). A Taxonomic and Nomenclatural Monograph of the Genus Amanita,
Section Amanita for North America. Germany: J. Cramer, Bib. Myco. 57.
Kerrigan, R. (1979-1985). Studies in Agaricus I: Mycologia71: 612-620; II: Mycologia
77: 137-141; III: Mycotaxon 22: 419^34.
Kerrigan, R. (1982). The Genus Agaricus in Coastal California. Master’s Thesis, San
Francisco State University; unpublished.
Smith, H.V. (1944). The Genus Limacella in North America. Pap. Mich. Acad. Sci. 30:
125-147.
Sundberg, W. The Family Lepiotaceae in California. Master’s Thesis, San Francisco State
University; unpublished.
Thiers, H.D. (1982). The Agaricales (Gilled Fungi) of California: Amanitaceae. Eureka:
Mad River Press.

Coprinaceae& Strophariaceae
Guzman, G. (1983). The Genus Psilocybe. Liechtenstein: J. Cramer.
Smith, A.H. (1951). The North American Species of Naematoloma. Mycologia 43:
467-521.
Smith, A.H. (1972). The North American Species of Psathyrella. N.Y. Bot. Gar. Myco.
Mem. 24.
Smith, A.H. & L.R. Hesler (1968). The North American Species of Pholiota. New York:
Hafner.
Vande Bogart, F. (1976-1979). The Genus Coprinus in Western North America I: Myco¬
taxon 4: 233-275; II: Mycotaxon 8: 243-291; III: Mycotaxon 10: 155-174.

Cortinariaceae & Gomphidiaceae

Ammirati, J.F. & A.H. Smith (1977). Studies in the Genus Cortinarius II: Section
Dermocybe, New North American Species. Mycotaxon 5: 381-397.
Hesler, L.R. (1969). North American Species of Gymnopilus. New York: Hafner, Myco.
Mem. 3.
Hesler, L.R. & A.H. Smith (1965). North American Species of Crepidotus. New York:
Hafner.
Miller, O.K. (1964). Monograph of Chroogomphus. Mycologia 56: 526-549.
Miller, O.K. (1971). The Genus Gomphidius. Mycologia 63: 1129-1163.
Smith, A.H. (1939). Studies on the Genus Cortinarius I. Contr. Mich. Univ. Herb. 2:1-42.
Smith, A.H. (1957). A Contribution Towards a Monograph of Phaeocollybia. Brittonia9:
195-217.
Smith, A.H., V.S. Evenson,& D.H. Mitchel(1983). The Veiled Species of Hebe/oma in the
Western United States. Ann Arbor: Univ. of Michigan Press.
Smith, A.H. & J. Trappe (1972). The Higher Fungi of Oregon’s Cascade Head Experi¬
mental Forest and Vicinity I. Mycologia 64: 1138-1153.
Stuntz, D. Macroscopic Field Key to Some of the More Common Species of Cortinarius
Found in Washington. Pacific Northwest Key Council.
Stuntz, D. Interim Skeleton Key to Some Common Species of Inocybe. Pacific North¬
west Key Council.
Thiers, H.D. & A.H. Smith (1969). Hypogeous Cortinarii. Mycologia 61: 526-536.
SUGGESTED READINGS AND PRIMARY REFERENCES 923

Boletaceae
Singer, R. (1977). The Boletineae of Florida. Liechtenstein: J. Cramer, Bib. Myco. 58.
Smith, A.H. & H.D. Thiers (1964). A Contribution Towards a Monograph of North
American Species of Suillus. Published by the authors.
Smith, A.H. & H.D. Thiers (1971). The Boletes of Michigan. Ann Arbor: Univ. of
Michigan Press.
Smith, A.H., H.D.Thiers,&O.K. Miller(1965). The Species of Suillus and Fuscoboletinus
of Priest River Experimental Forest and Vicinity, Priest River, Idaho. Lloydia28:
120-138.
Snell, W.H. & E.A. Dick (1970). The Boleti of Northeastern North America. Germany:
J. Cramer.
Thiers, H.D. (1975). California Mushrooms: A Field Guide to the Boletes. New York:
Hafner.
Thiers, H.D. (1976). Boletes of the Southwestern United States. Mycotaxon 3: 261-273.
Thiers, H.D. (1979). The Genus Suillus in the Western United States. Mycotaxon 9: 285-296.
Wolfe, C.B. (1979). Austroboletus and Tylopilus Subgenus Porphyrellus with Emphasis
on North American Species. Liechtenstein: J. Cramer, Bib. Myco. 69.

Polyporaceae, Stereaceae, & Allies

Canfield, E.,R. & R,L. Gilbertson (1971). Notes on the Genus Albatrellus in Arizona.
Mycologia 63: 964-971.
Gilbertson, R.L. (1976). The Genus Inonotus in Arizona. N.Y. Bot. Gar. Mem. 28: 67-85.
Gilbertson, R.L. (1981). North American Wood-Rotting Fungi That Cause Brown Rots.
Mycotaxon 12: 372^4-16.
Overholts, L.O. (1953). The Polyporaceae of the United States, Alaska, and Canada. Ann
Arbor: Univ. of Michigan Press.
Stuntz, D. (1980). Trial Field Key to the Pacific Northwest Polypores. Pacific Northwest
Key Council.
Welden, A.L. (1971). An Essay on Stereum. Mycologia 63: 790-799.

Hydnaceae
Hall, D. & D.E. Stuntz (1971-72). Pileate Hydnaceae of the Puget Sound Area I: Myco¬
logia 63: 1099-1128; II: Mycologia 64: 15-37; III: Mycologia 64: 560-590.
Harrison, K. A. (1964). New or Little Known North AmericanStipitateHydnums. Canad.
Jour. Bot. 42: 1205-1234.

Clavariaceae& Cantharellaceae
Bigelow, H.E. (1978). The Cantharelloid Fungi of New England and Adjacent Areas.
Mycologia 70: 707-756.
Marr, C. & D. Stuntz(1973). Ramaria in Western Washington. Germany: J. Cramer, Bib.
Myco. 38.
Petersen, R.H. (1968). The Genus Clavulinopsis in North America. N.Y. Bot. Gar. Myco.
Mem. 2.
Petersen, R.H. (1971). Notes on Clavarioid Fungi IX. Persoonia6: 219-229.
Petersen, R.H. (1971). The Genera Gomphus and Gloeocantharellus in North
America. Nova Hedw. 21: 1-118.
Petersen, R.H. (1975). Ramaria subgenus Lentoramaria. Leichtenstein: J. Cramer, Bib.
Myco. 43.
Petersen, R.H. (1981). Ramaria subgenus Echinoramaria. Leichstenstein: J. Cramer,
Bib. Myco. 79.
Scates, C. (1982). Trial Key to the Species of Ramaria in the Pacific Northwest. Pacific
Northwest Key Council.
924 BIBLIOGRAPHY

Wells, V.L. & P.E. Kempton (1968). A Preliminary Study of Clavariadelphus in North
America. Mich. Botanist 7: 35-57.

Lycoperdales& Sclerodermatales
Coker, W.C. & J.N. Couch (1928). The Gasteromycetes of the Eastern United States and
Canada. Chapel Hill: Univ. of North Carolina Press.
Guzman, G. (1970). Monografia del Genero Scleroderma. Darwiniana 16: 233-407.
Long, W.H. & D.J. Stouffer(1948). Studies in the GasteromycetesXVI: The Geastraceae
of the Southwestern United States. Mycologia40: 547-586.
Ramsey, R. (1980). Lycoperdon nettyana, a New Puffball from Western Washington
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Smith, A. H. (1951). Puffballs and Their Allies in Michigan. Ann Arbor: Univ. ofMichigan
Press.
Zeller, S.M. (1947). More Notes on Gasteromycetes. Mycologia 39: 282-312.

Tulostomatales, Podaxales, & Allies

Long, W.H. (1940-1946). Studies in the Gasteromycetes I: Dictyocephalos (with O.A.
Plunkett): Mycologia 32: 696-709; VII: Schizostoma{with D.Stouffer): Mycologia
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Long, W.H. (1946). The Genus Phellorina. Lloydia9: 132-138.
McKnight, K. (1985). The Small-Spored Species of Podaxis. Mycologia 77: 24-35.
Rea, P.M. (1942). Fungi of California I: Battarrea. Mycologia 34: 563-573.
Thiers, H.D. & R. Watling (1971). Secotiaceous Fungi from the Western United States.
Madrono 21: 1-9.

Hymenogastrales& Allies
Fogel, R. (1985). Studies on Hymenogaster. Mycologia 77: 72-82.
Singer, R. & A.H. Smith(1960). Studies in Secotiaceous FungiIX: The Astrogastraceous
Series. Mem. of Torrey Bot. Club 21: 1-112.
Smith, A.H. & R. Singer(1959). Studies in Secotiaceous Fungi IV: Gastroboletus, Trunco-
columella, Chamonixia. Brittonia 11: 205-223.
Thiers, H.D. (1979). New and Interesting Hypogeous and Secotioid Fungi from Cali¬
fornia. Sydowia Ann. Myc. 2:8: 361-390.
Trappe, J. (1975). A Revision of the Genus Alpova with Notes on Rhizopogon and the
Melanogastraceae. Beih. Nova Hedw. 6: 279-309.
Zeller, S.M. (1939). New and Noteworthy Gasteromycetes. Mycologia 31: 1-31.
Zeller, S.M. (1941). Further Notes on Fungi. Mycologia 33: 196-214.
Zeller, S.M. (1944). Representatives of the Mesophelliaceae in North America. Myco¬
logia 36: 627-637.

Phallales& Nidulariales
Arora, D. & W. Burk (1982). Clathrus archeri, a Stinkhorn New to North America.
Mycologia 74: 501-504.
Blanton, R.L. & W. Burk (1980). Notes on Pseudocolusfusiformis. Mycotaxon 12: 225-234.
Brodie, H.J. (1975). The Bird’s Nest Fungi. Toronto: Univ. of Toronto Press.
Burk, W.R. (1979). Clathrus ruber in California and Worldwide Distributional Records.
Mycotaxon 8: 463^468.
SUGGESTED READINGS AND PRIMARY REFERENCES 925

Dring, D.M. (1980). Contributions Toward a Rational Arrangement of the Clathraceae.

Kew Bulletin 35: 1-96.
Rea, P.M. (1955). The Genus Lysurus. Pap. Mich. Acad. Sci. 40: 49-66.

Pezizales
Denison, W.C. (1961). Some Species of the Genus Scutellinia. Mycologia51: 605-635.
Dissing, H. (1966). The Genus Helvetia in Europe. Dansk. Bot. Ark. 25: 1-172.
Kanouse, B.B. (1948). Studies in the Genus Otidea. Mycologia41: 660-677.
Kempton, P.E. & V.L. Wells (1970). Studies on the Fleshy Fungi of Alaska IV: A Prelimi¬
nary Account of the Genus Helvetia. Mycologia 62: 940-959.
Korf, R. (1973). Discomycetes and Tuberales (Chapter 9, Vol. IVA of The Fungi: An
Advanced Treatise, edited by Ainsworth, Sparrow, and Sussman). New York:
Academic Press.
Larsen, H.J.& W.C. Denison(1978). A Checklist of the Operculate Cup Fungi(Pezizales)
of North America West of the Great Plains. Mycotaxon 7: 68-90.
McKnight, K.H.(1969). A Note on Discina. Mycologia 61: 614-630.
Seaver, F.J. (1928). The North American Cup Fungi: Operculates. New York: Hafner
(1961 reprint).
Seaver, F.J. (1942). The North American Cup Fungi: Inoperculates. New York: Hafner.
Tylutki, E.E. (1979). Mushrooms of Idaho and the Pacific Northwest: Discomycetes.
Moscow, Idaho: Univ. of Idaho Press.
Weber, N.S. (1972). The Genus Helvetia in Michigan. Mich. Botanist 11: 147-201.

Tuberales
Burdsall, H.H. (1968). A Revision of the Genus Hydnocystis (Tuberales) and of the
Hypogeous Species of Geopora (Pezizales). Mycologia 60: 496-525.
Gilkey, H.M. (1916). A Revision of the Tuberales of California. Univ. of Calif. Publ. Bot.
6: 275-356.
Gilkey, H.M. (1954). Tuberales. North American Flora 2, 1: 1-36.
Parks, H.E. (1921). California Hypogeous Fungi—Tuberaceae. Mycologia 13: 301-314.
States, J.S. (1983). New Records of Hypogeous Ascomycetes in Arizona. Mycotaxon 16:
396-402.
Trappe, J. M. (1975). The Genus Fischerula (Tuberales). Mycologia 67: 934-941.
Trappe, J.M. (1979). The Orders, Families, and Genera of Hypogeous Ascomycotina
(Truffles and Their Relatives). Mycotaxon 9: 297-340.

Helotiales
Mains, E.B. (1954). North American Species of Geoglossum and Trichoglossum.
Mycologia 46: 586-631.
Mains, E.B. (1955). North American Hyaline-Spored Species of the Geoglossaceae.
Mycologia47: 846-877.
Mains, E.B. (1956). North American Species of the Geoglossaceae, Tribe Cudonieae.
Mycologia 48: 694-710.

Pyrenomycetes
Child, M. (1932). The Genus Daldinia. Ann. Missouri Bot. Garden 16: 4114-86.
Mains, E.B. (1940). Species of Cordyceps. Mycologia 32: 310-320.
Mains, E.B. (1947). New and Interesting Species of Cordyceps. Mycologia 39: 535-545.
926

GENERAL INDEX
Note: This index includes topics, persons mentioned or quoted in the text, scientific
names other than those listed in the “Genus and Species Index,” and selected common
names—those which are best known or most likely to be remembered.
Bold face numbers indicated detailed treatment of the subject; if the treatmentcovers
several consecutive pages, only the first is given.
*—Indicates a photograph or illustration without accompanying text other than
the caption.
To avoid the confusion that results from using two sets of numbers, color plates are
not included; they are listed on the page where the subject is described.

Abnormalities 898* Asci 4*, 782

Acacia 40 Ascobolaceae 817,834
Admirable Bolete 521 Ascomycetes 4, 55*, 782
Agaricaceae 310 Ascomycotina—see Ascomycetes
Agaricales 57,58 Ascus—see Asci
Agarics 52*, 57, 58 Aspen 42
Agaricus (section) 313 Aspen Bolete 540
Alder 39,42 Astrogastraceous Series 742
Alder False Truffle 757 Auriculariales 669
Allegro, John 894 Autodigestion 342,343*
Allergies to Mushrooms 892, 896
B
Almond Mushroom 336
Alpine Jelly Cone 674 Balsamiaceae 852
Amadou 581 Banana Slug 28*, 29*
Amanitaceae 262 Banded Agaricus 321
Amanitas 263 Barrows, Chuck 314, 331, 340, 529
egg stage 262* Bartelli, Ingrid 787
Amanita-Toxins 892 Basidia 4*, 6*
Amanitins 893 in jelly fungi 669*
Amatoxins 892 Basidioles 5,6*
Amethyst Laccaria 172 Basidiomycetes 4, 52-54*, 57
Ammirati, Joseph 430, 455 Basidiomycotina—see Basidiomycetes
Angel Wings 135 Basidium—see Basidia
Anise Agaricus 340 Basket Stinkhorn 770
Anise Mushroom 162 Bay Laurel 40
Annulus 16 Bearded Milk Cap 73
different types (Agaricus) 312 Beefsteak Fungus 553
Anonymous Amanita 275 Beefsteak Morels 799
Ant Fungus 881 Bell Morel 794
Ants 30,31 Bell-Shaped Panaeolus 356
Anywhere and Everywhere, Mush¬ Ben’s Bitter Bolete 524
rooms Found 47 Bigelow, Howard 154, 207
Aphyllophorales 57,548 Big-Foot 788
Apple Bolete 528 Big Laughing Mushroom 410
Apricot Jelly Mushroom 672 Bigwood, Jeremy 294
Artist’s Conk 576 Birch 41
Artist’s Fungus 576 Birch Bolete 541
Arvenses 313,731 Birch Conk 584
GENERAL INDEX 927

Birch Polypore 584 Bull Nose 800

Bird’s Nest Fungi 54*, 778 BUM’S 209,417,451,455,463
Bitorques 313 Burk, William 686, 769, 776
Bitter Bolete 523 Burned Areas, Mushrooms of 45
Bitter Hedgehog 620 Butter Bolete 525
Bitter Polypore 560 Buttery Collybia 216
Black Chanterelle 665 Button Mushroom 319
Black Cup Fungus 829 Butt Rot 550
Black Diamond 842*
C
Black Earth Tongues 866
Blackening Russula 89 Cabaniss, Michael 539
Blackfellow’s Bread 564 Caesar’s Amanita 284
Black-Foot 562 Caldwell, Katy 306
Black Jelly Drops 876 California Agaricus 327
Black Jelly Roll 672 Candlesnuff Fungus 885
Black-Leg 562 Candy Cap 80
Black Morel 786*, 790, 959* Canning Mushrooms 891
Black Truffle 854 Cannon Fungus 781
Black Witch’s Butter 672 Cantharellaceae 658
Blah Ramaria 651 Cantharellales 548
Bleeding Agaricus 325 Cap 15
Bleeding Coral Mushroom 653 cuticle 19*
Bleeding Milk Cap 68 shape of 17*
Bleeding Mycena 231 Carbon Balls 887
Blewit 9,10,11,153 Carbonizing Decay 550
Blue-Capped Polypore 558 Caterpillar Fungus 882
Blue Chanterelle 668 Cauliflower Mushroom(s) 657
Blue-Green Anise Mushroom 161 Cedar 41
Blue Leg 154 Cep 530
Blue Stain 878 Champignon 318
Blue-Staining Slippery Jack 504 Chanterelle 662
Blusher, The 276 Chanterelles 52*, 658
Blushing Amanita 276 Chaplin, Charlie 168
Bohemian Truffle 712 Chemical Characteristics of Mush¬
Bolbitiaceae 466 rooms 20
Boletaceae 488 Chestnut Bolete 510
Bolete Eater 883 Chicken Lips 874
Boletes 52*, 488 Chicken of the Woods 572
Boletivores 84,546 Chinquapin. 38
Booted Amanita 279 Chrome-Footed Bolete 533
Boring Brown Bolete 517 Classification 8
Boring Brown Cup Fungus 821 Clavariaceae 630
Boring Ubiquitous Mushrooms 209, Cleaning Mushrooms 889
417,451,455,463 Cloud Ears 669
Bracket Fungi 52*, 549 Club Corals 632
Brain Mushroom 801 Club Fungi 53*, 630
Bread Molds 782 Coccoli 284
Brown Crumbly Rot 578 Cocconi 285
Brown Field Mushroom 319 Coccora 265,284
Brown Rot 550 Cole, C. 185
Brown Witch’s Butter 673 Collecting Mushrooms 11, 26
Bufotenine 894 Colorful Earth Tongues 868
Bulbopodium 418,441 Comb Hericium 615
928 GENERAL INDEX

Commercial Collecting 26 Dead Man’s Hand 710

Commercial Mushroom—see Culti¬ Death Angel—see Destroying Angel
vated Mushroom Death Cap 269
Common Collybia 215 Deer Mushroom 255
Common Dung Cup 823 Deer Truffles 862
Common Earthball 708 Delicious Milk Cap 68
Common Puffball 693 Delignifying Decay 550
Common Puffballs 690 Dermocybe (subgenus) 418
Common Stinkhorns 766 Desert Shaggy Mane 725
Compactae 84 Deserts, Mushrooms of 46
Compost, Shalom 325 Desjardin, Dennis 207, 208, 210, 214,
Conch Rot 583 854
Cone Heads 470 Destroying Angel 271
Conical Waxy Cap 116 Devil’s Dipstick 771
Conifer Coral Hericium 614 Devil’s Eggs 765
Coniophoraceae 604 Devil’s Stinkhorn 770
Conks 574 Devil’s Urn 829
Constricted Grisette 289 Dickenson, Emily 3
Cookery, Mushroom 888 Dictionary of Scientific Names (Latin
Coprinaceae 341 and Greek Word Elements) 899,903
Coprine 896 Discomycetes 783
Coral Fungi 53*, 630,645 Disturbed Ground, Mushrooms of 45
Corn Silk Inocybe 457 Dog Stinkhorn 771
Corticiaceae 604 Domicile Cup Fungus 822
Cortinariaceae 396 Donkey Ears 832
Cortinarius (subgenus) 418,446 Douglas-Fir 35
Cosmopolitan Mushrooms 47 Dow, Anne 547
Cottonwood 39,42 Doyle, Sir Arthur Conan 2
Cousteau, Jacques 573 Dryad Saddle 561
Cracked Cap Bolete 519 Drying Mushrooms 20, 890
Crampballs 887 Dry Rot Rungus 610
Crater Cup 829 Dunce Caps 470
Crepidotaceae 396,405 Dung Mushrooms 42
Crested Coral 641 Dutra, Frank 484
Crocodile Agaricus 334 Dyeing 418,454,455,571,712
Crown Coral Mushroom 642 Dyer’s Polypore 570
Crown Fungus 825
E
Crust Fungi 53*, 604
Cryptic Globe Fungus 585 Early Agrocybe 469
Cultivated Mushroom 319 Early Morel 793
Cultivation of Mushrooms 30 Ear Pick Fungus 629
Cup Fungi 55*, 783, 817 Earthballs 707
Cypress 36 Earth Fan 608
Cystidia 5,6*, 19*, 20 Earth Nuts 854
Czarnecki Family 176 Earthstars 54*, 677, 699
Earth Tongues 55*, 865
D
Edibility of Mushrooms 23, 265
Dacrymycetales 669 Elephant Ears 799, 803
Dark Bolete 534 Elfin Cups 805
Darwin, Charles 766 Elfin Saddles 55*, 783, 796, 805
Deadly Cortinarius 444 Emetic Russula 97
Deadly Galerina 401 Enokitake 220
Deadly Parasol 308 Entolomataceae 238
Dead Man’s Fingers 886 Erving, Julius 8
Dead Man’s Foot 712 Eucalyptus 39
GENERAL INDEX 929

Evenson, Verna Stucky 466 Garlic Mushroom 207

Everson, Bill 526 Gasteromycetes 57,676
Eyelash Pixie Cup 839 Gastroid Agarics 53*, 724
Gastroid Agaricus 729
F
Gastroid Boletes 544
Fairy Bonnet 352 Gastroid Coprinus 727
Fairy Clubs 634 Gastroid Cortinarius 734
Fairy Fan 871 Gastroid Lepiota 731
Fairy Fingers 637 Gastroid Liberty Cap 734
Fairy Hair 636 Gastroid Milk Cap 738
Fairy Ring Mushroom 7*, 208, 510 Gastroid Pholiota 735
Fairy Rings 6, 209, 687 Gastroid Pine Spike 732
Fairy Stool 568 Gastroid Russula 737
Fairy Tub 821 Gastrointestinal Irritants 895
False Chanterelle 479 Gelatinous Coral Mushroom 655
False Morel 801 Gemmed Amanita 281
False Morels 796, 799 Gemmed Puffball 693
False Sparassis 824 Geneaceae 849
False Tinder Polypore 581 Geode Truffles 849
False Truffles 54*, 739, 844 Geoglossaceae 866, 868, 872
False Turkey Tail 605 Gerry, Eric 733
Fan Pax 476 Giant Gel Cup 828
Fat-Headed Black Morel 791 Giant Gymnopilus 410
Felted Heart Rot 580 Giant Horse Mushroom 333
Felt-Ringed Agaricus 326 Giant Puffball 678*, 682
Fetid Russula 92 Giant Puffballs 680
Field Mushroom—see Meadow Mush¬ Gilbertson, Robert 543
room Gilkey, Helen 843, 858
Field Puffball 695 Gilled Bolete 480
Fir 41 Gilled Mushrooms 52*, 57, 58
Fires 45 Gilled Polypore 589
Flask Fungi 55*, 878 Gills 16
Flesh 16 arrangement of hyphae in 19*
Flower Pot Parasol 302 attachment to stalk 17*
Fluted Black Elfin Saddle 815 cross-section of 6*
Fluted Brown Elfin Saddle 814 Girolle 663
Fluted White Elfin Saddle 816 Glistening Inky Cap 348
Fly Agaric 265, 282 Glutinous Gomphidius 482
Fly Amanita 282 Goat’s Foot 560
Fogel, Robert 748 Golden Fairy Club 638
Foxfire 196 Golden Pholiota 390
Fragrant Russula 92 Golden Waxy Cap 115
Freezing Mushrooms 891 Gomphaceae 630
Fried Chicken Mushroom 174 Gomphidiaceae 481
Fries, Elias 58,418 Granulated Slippery Jack 502
Frost’s Bolete 528 Greek 899
Fungi 4 Green Earth Tongue 870
role in environment 6 Greening Goat’s Foot 559
Fungophobia 1 Green-Spored Parasol 295
Funnel Chanterelle 665 Grisette 288
Fuzzy Truffle(s) 846 Gristly Truffle 857
Guzman, Gaston 707
G
Gypsy Mushroom 412
Garden Mushrooms 43 Gyromitrin 893
930 GENERAL INDEX

H Iodine Polypore 560

Irregular Earth Tongue 871
Habitats 34
Isaacs, Bill 314,331
Half-Free Morel 791
Ivory Waxy Cap 119
Hall, Joseph 29
Hallucinogenic Mushrooms 29, 31, Jack-O-Lantern Mushroom(s) 147
265, 282, 354, 358, 359, 368, 370-374, Jacobs, Jim 752
409,410, 894, 895 JAR’s 84
Harding, Paul 69 Jelly Babies 874
Hard-Skinned Puffballs 707 Jelly Fungi 53*, 669
Haymaker’s Panaeolus 360 Jenkins, David 265
Hays, William Delisle 1 Jochelson, Vladimir 29
Heart Rot 549 Judas’ Ear 675
Hedgehog Mushroom 618 Juniper 41
Hedgehog Mushrooms 616 Just Another Russula 84
Helotiales 865 Kerrigan, Rick 314, 323, 324, 332,
Helvellaceae 796,848 379,729
Hemlock 40 Keys 21
Henis, M. 185 dichotomous (initial key) 52
pictorial key to gilled mushrooms 61
Hen of the Woods 564
pictorial key to major groups of
Hesler, L.R. 64,385
fleshy fungi 52
Hideous Gomphidius 482
King Alfred’s Cakes 887
Hole in the Ground 847
King Bolete 530
Honey Mushroom 34*, 196
Kurokawa 556
Hongo, Tsuguo 90
Hooded False Morel 802 L
Horn of Plenty 666
Horsehair Fungus 208 Laboulbeniomycetes 47
Horse Mushroom 332 Lackluster Laccaria 172
Hortenses 313 Landslide Mushroom 374
Huckleberry 40 Larch 40
Hydnaceae 611 Larch Boletes 505
Hygrophoraceae 103 Lantern Stinkhorn 776
Hygroscopic Earthstar 705 Largent, David 242, 244, 248, 249, 251
Hymenium 57,783 Latex 63,64
Hymenochaetaceae 604 Latin 9,899
Hymenogastrales 739 Latticed Stinkhorn 773
Hymenomycetes 57 Lawn Mushrooms 43
Hyphae 5, 19* Lawrence, D.H. 2
LBJ’s 33*
IJK LBM’s 32
Ibotenic Acid 894 Lenin, V.S. 3
Identifying Mushrooms 14 Leotiaceae 872, 875, 877
Incense Cedar 41 Lepidellas 265,275
Indecisive Bolete 535 Lepiotaceae 293
Indian Paint Fungus 613 Leprocybe 418
Indigo Milk Cap 69 Leucogastrales 759
Indoor Mushrooms 44 Liberty Cap 370
Inky Cap 9, 347 Lichen Agaric 223
Inky Caps 342 Lilac Inocybe 461
Inocybium 455 Lincoff, Gary 373,821
Inrolled Pax 477 Ling Chih 577
Insects 28,29 Linnaeus 785
Insidious Gomphidius 482 Lion’s Mane Hericium 615
Little Brown Jobs 33*
GENERAL INDEX 931

Little Brown Mushrooms 32 Morchellaceae 784

Little Helmet 352 Morel 787
Lizard’s Claw Stinkhorn 111 Morels 31*, 44*, 55*, 783, 784, 785
Lobster Mushroom 884 Moser, Meinhard 419
Long, W.H. 714 Mountain Gastroid Agaricus 730
Lorchels 799 Muk Ngo 675
LSD 895 Muscarine 455,894
Luminescence 146, 148, 196 Muscimol 894
Lumpers 10 Mushroom(s) 1-959
Lupine, Bush 40 allergies to 892, 896
Lycoperdales 677 carnivorous 29
chemical characteristics of 20
M
classification of 8
Macroscopic Characteristics 14 collecting 11,26
Madrone 38 color 14
Maggots 28, 29, 889 color changes 15
Magic Mushroom 373 cookery 888
Magpie Mushroom 346 definition of 4
Man On Horseback 179 distinctive 48
Mantis, Praying 28* dyeing with 418,454,455,571,712
Manure Mushrooms 42 edibility of 23, 265
Manzanita 38 evolution 57,724,739,841
Manzanita Bolete 539 fear of 1
Maple 39,41 growing 30
Marr, Currie 645 hallucinogenic 29, 31, 265, 282, 354,
Marsh, Ben 525 358, 359, 368, 370-374, 409, 410,
Marshall, Nina 769 894,895
Marshmallow Polypore 600 handling 27
Matsutake 49*, 189,191, 192 hunters 84, 224, 546-547
Mattoon, Judith Scott 576*, 876 hunting 11,25
Mcllvaine, Charles 113, 123, 141, 355, equipment 12
376, 419,445, 584, 690, 700, 765, 768, field notes 13
769,802, 866,871 identification of 14
Meadow Mushroom 318 macroscopic characteristics 14
Medicinal Value of Mushrooms 30 medicinal value 30
Melanogastraceae 756 microscopic characteristics 19
Melzer’s Reagent 20 names 8,899
Menser, Gary 854, 861 nutritional value 30
Methven, Andrew 79 odor 15
Mica Cap 348 parts of a 5*, 14
Microscopic Characteristics 19 picking 26
Milazzo, Ciro 145, 553 dangers of 27*
Milk Caps 64 poisoning 892
Miller, Luen 123 preservation of 20, 890
Miller, Orson K. 355, 689, 717, 793, role in the environment 6-8
826 seventy distinctive 48
Miniature Waxy Cap 113 sexuality 5-6
Minores 313 size 14
Miscellaneous Toxins 896 succession of 8
Mitchel, D.H. 466 taste 15,63
Mitchell, Craig 414, 666 texture 15
MMH —see Monomethylhydrazine toxins 892
Monomethylhydrazine 799,893 weeds 47
when and where they grow 25
932 GENERAL INDEX

Mycelium 5,18,43* Piercy, Marge 31

Mycorrhiza 7 Pigs 29,842
Mycorrhizal Fungi 7, 34 Pigs 29,842
Myxacium 418,429 Pig’s Ears 661,797,798
Pine 35
NO
Pine Conk 582
Narrow-Headed Black Morel 791 Pine Spike 485
Niche 7 Pine Spikes 484
Nidulariales 778 Pink-Bottom 318
Neruda, Pablo 888 Pinkish Coral Mushroom 654
Not So Tedious Tubaria 403 Pink-Tipped Coral Mushroom 656
Nutritional Value of Mushrooms 30 Pleated Marasmius 209
Oak 37 Pluteaceae 253
Oak Conk 582 Pocket Rot 550
Oak-Loving Collybia 215 Podaxales 724
Oak Root Fungus 196 Poisoning, Mushroom—see Toxins
Octopus Stinkhorn 774 Poison Oak & Ivy 27*
Oddballs 711 Poison Pax 477
Old Man of the Woods 543 Poison Pie 464
Old Man’s Beard 615 Polyporaceae 549
Orange Peel Fungus 837 Polypores 52*, 549
Orange Sponge Polypore 571 Poor Man’s Candle 123
Oregon Black Truffle 854 Poor Man’s Gumdrop 674
Oregon White Truffle 858 Poor Man’s Licorice 876
Ornate-Stalked Bolete 522 Poor Man’s Slippery Jack 504
Ornate Stinkhorns 772 Poor People’s Truffle 668
Ox Tongue 552*, 553 Poplar 42
Oyster Mushroom 29,134 Poplar Tricholoma 185
P Porcini 530
Pores 488,549
Paddy Straw Mushroom 258, 262 Potato Chip Conditions 35
Panther Amanita 280 Powdery Mildews 782
Paraphyses 5 Preserving Mushrooms
Parasitic Fungi 6 for study 20
Parasol Mushroom 298 for consumption 890
Parasol Mushrooms 293 Prince, The 337
Parchment Fungi 53*, 604 Psilocin 368,895
Parks, Harold 842 Psilocybin 368,895
Pasture Mushrooms 43 Psilocybin Mushrooms 31, 895
Paxillaceae 476 Psychedelic Mushrooms—see Hallu¬
Pear-Shaped Puffball 691 cinogenic Mushrooms
Peele, Stephen 294 Puffballs 54*, 677
Pellegrini, Angelo 889 parts of 677*
Peppery Bolete 517 Pungent Slippery Jack 503
Peppery White Milk Cap 71 Purple Fairy Club 637
Pezizaceae 817 Purple-Spored Puffball 687
Pezizales 783 Purple-Staining Milk Cap 75
Pfifferling 663 Pyrenomycetes 878
Phallaceae 766 Pyronemataceae 817,831, 834, 846
Phallales 764
Phallolysin 892 QR
Phallophobia 766 Queen Bolete 531
Phallotoxins 893 Questionable Stropharia 377
Phlegmacium 418 Quinine Conk 579
Pickling Mushrooms 892 Quinine Fungus 579
GENERAL INDEX 933

Rabbit Ears 833 Scaly Pholiota 389

Ramsbottom, John 611,773 Scarlet Cup Fungus 836
Ramsey, Robert 688 Scarlet Waxy Cap 114
Rattlesnakes 28* Scates, Catherine (Kit) 645, 651
Raverat, Gwen 766 Schizophyllaceae 590
Rea, Paul 777 Sclerodermatales 678, 699, 707
Red-Belted Conk 578 Sclerotiniaceae 877
Red-Brown Butt Rott 571 Scotch Bonnet 202
Red-Capped Butter Bolete 526 Sculptured Puffball 684
Red Chanterelle 664 Sellers, Bob 708
Red Coral Mushroom 655 Sericeocybe 418,448
Red Gravel 757 Sex 5,6
Red Morel 31*, 791 Shaggy Mane 9, 56*, 343*, 345
Red-Pored Bolete 528 Shaggy Parasol 297
Red-Stemmed Bitter Bolete 524 Shaggy-Stalked Parasol 309
Reduced Agarics 724 Sheep Polypore 557
Redwood 36 Sheep’s Head 564
Redwood Rooter 218 Shelley, P.B. 2,342
Reindeer 29 Shiitake 31*, 141
Rhodophyllaceae 238 Shingled Hedgehog 619
Ring(s)—see Annulus Shoehorn Oyster Mushroom 136
Riparian Woodland 39 Shoe String Root Rot 196
Ristich, Sam 672 Short-Stemmed Russula 87
Roody, Bill 771 Short-Stemmed Slippery Jack 501
Rosy Conk 580 Shrimp Russula 102
Rosy Gomphidius 483 Siberian Slippery Jack 498
Rosy Larch Bolete 506 Sickener, The 97
Rosy Russula 99 Sierran Puffball 684
Rots 549,550 Silver-Leaf Disease 606, 607
Rough-Stemmed Boletes 536 Singer, Rolf 207
Rufous Candy Cap 82 Slippery Jack 500
Russia 3 Slippery Jacks 491
Russulaceae 63 Slippery Jill 499
Russulales 63 Sloane, Hugo 822, 823
Russulas 83 Slugs 29
Rusts 57 Smith, Alexander 7, 64, 118, 138, 224,
Rusty Gilled Polypore 590 235, 261, 345, 360, 361, 384, 385, 419,
434, 435, 455, 466, 473, 489, 499, 585,
S
657, 691, 696, 740, 753, 754, 766, 769,
Saint Ciro 484 772, 804, 864, 874
Salting Mushrooms 892 Smith, Helen 740, 753
Saltshaker Earthstar 704 Smooth Parasol 299
Sand Amanita 273 Smooth-Spored Truffles 852
Sand-Loving Mushrooms 46 Smuts 57
Sandy, The 185 Snitow, Alan 1
Sanguinolenti 313 Snowbank False Morel 800
Saprophytic Fungi 6, 7 Snowbank Mushrooms 46
Sap Rot 550 Snow Mushroom 800
Sarcoscyphaceae 817,834 Snow-Puff Mushroom 220
Sarcosomataceae 817,826 Soapy Tricholoma 184
Satan’s Bolete 527 Soma 894
Saylor, Herb 652, 742, 744, 854, 856, Sordid Waxy Cap 122
864 Spawn—see Mycelium
Scaly Chanterelle 661 Specialized Habitats, Mushrooms
with 47
934 GENERAL INDEX

Sphaeriales 878 T
Sphaerocysts 63
Tamarack Jack 497
Sphere Thrower 781
Tanoak 37
Split-Gill 590
Tawny Grisette 287
Splitters 10
Taxonomy 8
Sponge Mushrooms 785
Teeth Fungi 53*, 611
Spore Color 18
Telamonia 418,450,451
Spore Prints 18
Terfeziaceae 854-855
Spores 4, 5, 18, 19
Terminology 14
developing 6*
Termites 30
numbers of 576-577, 680
Terrestrial Pholiota 389
truffle 842*
Thelephoraceae 604
Spreading Brown Cup Fungus 821
Thick-Footed Morel 788
Spring Agrocybe 469
Thick-Skinned Puffballs 707
Spring Mushrooms 46
Thick-Walled Maze Polypore 587
Springtime Amanita 286
Thiers, Harry 285, 489, 536, 742, 745
Spruce 40
Thimble Morel 794
Squirrels 29
Thimble Morels 793
Staghorn Jelly Fungus 674
Thin-Walled Maze Polypore 588
Stalk 16
Ticks 27*
ornamentation (boletes) 488-489
Tidepool Conditions 35
position of 17*
Tinder Polypore 581
shape of 17*
Toadstool, Definition of 25
Stalked Latticed Stinkhorn 776
Toadstool Tester 24*
Stalked Oddball 713
Toothed Jelly Fungus 671
Stalked Polypores 554
Top Rot 550
Stalked Puffballs 54*, 715
Totally Tedious Tubaria 402
Stamets, Paul 368
Toxins, Mushroom 892
Starving Man’s Licorice 828
Trappe, James 748, 758, 843, 860
Starving Man’s Slippery Jack 504
Tree Ears 675
Steinpilz 530
Tree Oyster—see Oyster Mushroom
Stem—see Stalk
Trees 34-42
Stereaceae 604
Tree Trunk Morel 788
Stickney, Larry 117, 672
Tremellales 669
Stinker, The 179
Tricholomataceae 129
Stinkhorn 768
True Truffles 854
Stinkhorns 54*, 764, 766, 772
Truffle-Hounds 842,855
Stinkopus 775
Truffle Hunting 842-843
Stinky Squid 776
Truffle-Like Peziza 822*, 824
Stone Fungus 563
Truffles 55*, 739, 841, 854
Strawberries and Cream 627
Truly Trivial and Totally Tedious
String and Ray Rot 565
Tubaria 403
Strophariaceae 367
Truly Trivial Tubaria 403
Stuntz, Daniel 373,419,434,455,
Trumpet of Death 668
458, 560, 645
Trunk Rot 550
Stuntz’s Blue Legs 372
Tuberaceae 854,855
Suburban Psathyrella 363
Tuberales 841
Suffixes, Latin and Greek 900, 901
Tubes 488,549
Sulfur Shelf 572
Tuckahoe 564,604
Sulfur Tuft 382
Tulostomatales 678,715
Sunny Side Up 474
Tumbling Puffball 697
Swamp Beacon 870
Tumbling Puffballs 696
Sweat-Producing Clitocybe 163
Tunbridge Ware 878
Sweetbread Mushroom 240
Turkey Tail 594
GENERAL INDEX 935

UV White Matsutake 49*, 191

Umbrella False Morel 804 White Morel 788*, 789
Van de Bogart, Fred 346 White Rot 550
Varnished Conk 577 White Truffle 854
Veil(s) 16 Willow 39,42
different types (Agaricus) 312 Wine-Colored Agaricus 326
different types (Amanita) 263-264 Wine-Red Stropharia 378
Veiled Polypore 585 Winter, Bob 143
Veiled Stinkhorn 771 Winter Chanterelle 665
Veined Brown Cup Fungus 796 Winter Mushroom 220
Velvet Foot 220 Witch’s Butter 673
Velvet Pax 478 Witch’s Hat 116
Velvet Stem 220 Woman on Motorcycle 299
Velvety Black Earth Tongue 867 Wood Blewit 154
Violet Cortinarius 446 Wood Ear 675
Violet Cup Fungus 824 Woodland Agaricus 334
Violet Hedgehog 622 Wood Truffles 848
Viscid Black Earth Tongue 866 Woolly Chanterelle 661
Vole, California Red-Backed 841 Worms—see Maggots
Volva 18 Wright, Greg 69, 161, 241, 346, 350,
different types (Amanita) 263-264 404, 475, 528
Volvariaceae 253 Wrinkled Thimble Morel 793
Vulcan Pixie Cup 840 XYZ
W Xanthodermati 313
Walnut 800 YAM’s 224
Wasson, R. Gordon 29, 894 Yeasts 782
Wasson, Valentina 320 Yellow Coral Mushroom 652
Waxy Caps 103 Yellow Morel 787
Weber, Nancy S. 740, 753, 816 Yellow-Staining Agaricus 329
Western Giant Puffball 684 Yellow-Staining Milk Cap 74
Western Grisette 290 Yellow-Veiled Stinkhorn 771
Wet Rot 611 Yet Another Mycena 224-225, 234
Wheat Ergot 895 YungNgo 675
White Chanterelle 662 Zeller’s Bolete 518
White King Bolete 529 Zygomycetes 844

Geopora arenico/a (see description on p. 847) is half cup fungus, half truffle. When young it looks
like a hole in the ground because all but the mouth is immersed in the soil. As it matures, however, the
hole broadens, and in old age the margin of the cup often splits into lobes as shown on left.
936

GENUS AND SPECIES INDEX

Note: This index includes only genera and species so that it can also serve as a check¬
list. See “General Index” for common names, topics, persons, etc.
Bold face numbers indicate detailed treatment of the subject; if treatment covers
several consecutive pages, only the first is given.
*—Indicates a photograph or illustration without accompanying text other than
the caption.
To avoid the confusion that results from using two sets of numbers, color plates are
not included in the index; they are listed on the pages where subjects are described.

A
Agaricus (cont.) Agrocybe (cont.)
Abortiporus lilaceps 323 arvalis 469
biennis 566 “luteovelatus” 324 cylindracea 470
Abstoma 680 macrosporus 334 dura 470
reticulatum 681,690 maritimus 322 erebia 467,470
townei 681,690 meleagris 329 firma 468
Agaricus 58*, 310, 896 micromegathus 340 paludosa 467
abruptibulbus 335 nivescens 334 pediades 43*, 468
albolutescens 335 osecanus 333 praecox 469
alb o sanguineus 325 pattersonae 316,324 retigera 468
altipes 317 perobscurus 339 semiorbicularis 469
amethystina 341 perrarus 338 sororia 468,469
amieosus 316, 326 pinyonensis 331 tuberosa 469
argenteus 319 placomyces 314,329, Albatrellus 554
arorae 311*,325 895 avellaneus 557,558
arvensis 12*, 312*, 332 pocillator 314,329 caeruleoporus 555,559
augustus 311*, 337 porphyrocephalus 319 confluens 556,558
“barrowsii” 331 praeclaresquamosus cristatus 556,560
benesi 316,325 312*,329,895 dispansus 556
bernardii 322 purpurellus 341 ellisii 559
bisporus 319 rhoadsii 317 flettii 558
bitorquis 312*, 321 rodmani 322 hirtus 560
blandianus 320 rubronanus 325 ovinus 557
brunnescens 320 rutilescens 317,319 peckianus 558
californicus 312*, 327, semotus 315, 339*, 340 pescaprae 558*, 560
895 sequoiae 317,332 similis 558
campestris 312*, 318, silvaticus 325 sylvestris 560
889* silvicola 312*, 334 Alboleptonia 249
chionodermus 315, smithii 315,338 adnatifolia 253
317,335 solidipes 318 ochracea 253
chlamydopus 322 spissicaulis 326 sericella 252
comtulus 315,340 spp. (unidentified) 317,
Aleuria 833
cretacellus 315 318,325
aurantia 837
crocodilinus 315,317, subfloccosus 320
rhenana 836
334 subrufescens 336
rutilans 837
cupreobrunneus 319 subrutile scens 326
summensis 315,339
Alnicola 399
diminutivus 340 escharoides 400
dulcidulus 341 sylvicola 335
melinoides 400
edulis 322 urinescens 334
scolecina 400
fissuratus 333 vaporarius 317
villaticus 334 Alpova 756
fuse ofib rillosus 325
vinaceo-umbrinus 326 alexsmithii 757,758
fuscovelatus 324
vinaceovirens 322 cinnamomeus 758
haemorrhoidarius 325
xanthodermus 329,895 diplophloeus 757
halophilus 322
luteus 758
hondensis 326,895 Agrocybe 467
nauseosus 757,758
hortensis 320 acericola 470
olivaceoniger 757,758
lanipes 316 aegerita 467,470
olivaceotinctus 757*,
amara 469
758
GENUS AND SPECIES INDEX 937

Alpova (cont.) Amanita (cont.) Armillaria (cont.)

superdubius 758 rubescens 264*, 266*, pitkinensis 194
trappei 757,758 276,884* ponderosa 49*, 191
Amanita 262,263 russuloides 281 straminea 190,194
abrupta 275 salmonea 269 tabescens 197
alba 266,288 silvicola 273,274* viscidipes 190
aspera 278 smithiana 275 zelleri 189
atkinsoniana 275 solitaria 275 Armillariella 189
baccata 273 spissa 269,280 bulbosa 190,197
bisporigera 273,892 sp/?. (unidentified) 268, mellea 34*, 196,896
boudieri 274 275,277 tabescens 150,190,197
breckonii 281 spreta 266
Ascobolus 838
brunnescens 267,279 strangulata 289
carbonarius 835,838
caesarea 284 strobiliformis 275
calyptrata 265,284, thiersii 275
sarcoides 875,877
679*,890* umbonata 284
calyptratoides 269,286 umbrinolutea 290 Ascotremella
calyptroderma 285 vaginata 288,289* faginacea 875,877
ceciliae 289 velosa 266*,286 turbinata 877
chlorinosma 275 ve/Twz 273,892
cinereoconia 269,276 virosa 267,273,892 772,775
cinereopannosa 275 volvata 266 Asterophora 200
citrina 267,279 wellsii 267 lycoperdoides 201
coker i 268,275 Amanitopsis 263 parasitica 201
constricta 289 (also see Amanita) Astraeus 699
cothurnata 268,280 vaginata 288 hygrometricus 700,705
crocea 287,288 Amylocystis pteridis 706
daucipes 275 lapponica 600 Aurantiop or ellus 572
farinosa 267,276 Anellaria Aurantioporus
flavoconia 278 separata 355 croceus 597
flavorubens 278 Anthopeziza Auricularia 669
flavorubescens 269,278 Jloccosa 836 auricula 675,958*
frostiana 279 delicata 675
Anthracobia
fulva 287 mesenterica 675
macrocystis 838
gemmata 281, 894 polytricha 675
melaloma 834,838
hemibapha 284
Ant hums Auriscalpium 629
hesleri 269
archer i 775 vulgare 629
inaurata 268, 289
aseroeformis 775 Austroboletus 508
junqudlea 281
borealis 778 betula 508
longipes 275
Arachnion gracilis 534
magniverrucata 274
album 679 subflavidus 508
mappa 279
muse aria 15,29,264*, /I raneosa
B
265,282,894,895 columellata 679
mutabilis 273 Arcangeliella 736 Baeospora
ocreata 12*, 266*, 271, camphorata 737,739 myosura 202,212
286*,892,893 crassa 738 myriadophylla 202
onusta 269,276 lactarioides 739 Balsamia 852
pachycolea 290 739 magnata 853
pantherina 264*, 280, variegata 737,739 Banker a 616
894 Ar miliaria 189 carnosa 618
parcivolvata 267,284 albolanaripes 194 fuligineo-alba 618
peckiana 267 bulbosa 197 violescens 618
phalloides 264*, 269, caligata 192 Barssia 852
892,893 decorosa 189 oregonensis 853
polypyramis 275 dryina 136 sp. (unidentified) 854
porphyria 279 fusca 190,194 Battarrea 715
praegraveolens 275 luteovirens 194 digued 716,718
ravenelii 275 mellea 196 laciniata 718
rhopalopus 275 olida 193 phalloides 111
roseitincta 269
stevenii 718
938 GENUS AND SPECIES INDEX

Bisporella Boletus (cont.) Boletus (cont.)

citrina 877 fibrillosus 523,529* vermiculosus 516
Bjerkandera 592 flammans 515,529 viridiflavus 516, 522
adusta 596 flaviporus 522 zelleri 518
fumosa 596 fragrans 514 Bondarzewia 554
Bolbitius 473 fraternus 521 berkeleyi 556,565
aleuriatus 475 frostii 528 montana 548*, 565
callisteus 475 griseus 516,522 Bovista 696
coprophilus 474,475 haematinus 513,528, californica 698
lacteus 474,475 895 color ata 696,698
reticulatus 475 hemichrysus 512 dakotense 697,698
sordidus 474 hypocarycinus 527 leucoderma 698
vitellinus 474 inedulis 515 long isp ora 697,698
Boletellus 508 longicurvipes 516 minor 696,698
ananas 508,509 luridus 515,528 nigrescens 697,698
betula 508 “marshii” 524 pi la 697
chrysenteroides 508 mendocinensis 513,520 plumbea 697
russellii 508 miniato-pallescens 521 pusilla 697,698
mirabilis 521 Bovistella 690
Boletinellus
mottii 530 radicata 691,694
merulioides 490
olivaceobrunneus 523 Brauniellula 732
Boletinus 491
ornatipes 522 albipes 733
(also see Suillus &
orovillus 513,527 nancyae 732
Fuse ob ole tinus)
pallidus 515
cavipes 495 Bulgaria 875
parasiticus 512
Boletopsis 554 inquinans 876
piedmontensis 515
griseus 557 melastoma 830
pinicola 530
leucomelas 557 mexicana 828
pinophilus 530
subsquamosa 556 rufa 827
piperatoides 517
Boletus 511,894 Byssonectria
piper atus 517
(also see Suillus, aggregata 828
porosporus 520
Leccinum, Tylopilus, projectellus 516, 522 C
etc.) pseudo separans 530
abieticola 514,525 pseudosulphur eus 515 Calbovista 680
aereus 50*,512*,531 pulcherrimus 528, 895 subsculpta 682,685
affinis 514,516 pulverulentus 514, 515 Callistosporium 202
albidus 525 radicans 525 graminicolor 211
amygdalinus 527 regius 512*, 526 luteo-olivaceum 211
appendiculatus 512*, retipes 522 Calocera
525
rubellus 515, 521 cornea 635,670,675
auriflammeus 516, 522
rubinellus 516 viscosa 674
auriporus 516, 522
rubripes 524 Calocybe 173
badius 516,519
rubroflammeus 515, 529 came a 176
barrowsii 512*, 529
rubropunctus 516 gambosa 183
betula 508 satanas 527,895,896 onychina 150,176
bicolor 521 sensibilis 521
Caloporus
caespitosus 522 separans 530 dichrous 552
calopus 523 smithii 513,520
campestris 521 Caloscypha
spadiceus 518
fulgens 837
chromapes 534 speciosus 515,526
chrysenteron 519 Calostoma 715
sphaerocephalus 512
citriniporus 515,523 cinnabarina 718
spp. (unidentified) 514,
coccyginus 513,521 lutescens 716,719
523,528
coniferarum 514,523 microsporum 716,719
subglabripes 516
curtisii 516 ravenelii 716,719
subtomentosus 488*,
dryophilus 520 517
Calvatia 677*, 680
eastwoodiae 528 booniana 684
subvelutipes 515,527,
edulis 489*, 512*, 530 , 895
bovista 686
546,547,890* caelata 686
truncatus 514,520
erythropus 526 895 , variipes 516,531
Candida 682,686
fagicola 516
GENUS AND SPECIES INDEX 939

Calvatia (cont.) Cantharellus (cont.) Chroogomphus (cont.)

craniiformis 682,688 minor 660,665 ochraceus 485,486
cretacea 681,688 odoratus 660,664 pseudovinicolor 486
cyathiformis 687 purpurascens 660 rutilus 485,486
diguetii 688 subalbidus 662 tomentosus 487
elata 682,686 tubaeformis 665 vinicolor 20*, 484*, 485
excipuliformis 682,686 xanthopus 660,665 Ciboria 877
fragilis 681,687 Carbomyces 844 amentacea 877
fumosa 688 Catathelasma 189 Clathrus 772
gigantea 678*, 680, 682 imperials 195 archer i 774
hesperia 681,689 macrospora 195 bicolumnatus 773
lepidophora 681,689 singer i 195 cancellatus 773
Hoy dii 682,686 ventricosa 190, 195 cibarius 774
lycoperdoides 687
Caulorhiza 202 columnatus 773
maxima 683
hygrophoroides 219 crisp at us 773
owyheensis 681
umbonata 218 crispus 773
pachyderma 681,689
Cerrena preussii 774
pallida 682,686
unicolor 587, 589, 592, ruber 773
paradoxa 681
597 Claudopus 238
rubroflava 682,688
rubrotincta 682,687 Chaetoporus byssisedus 238
saccata 686 euporus 602 depluens 238
sculpta 684 Chamaeota 253 graveolens 238
Chamonixia 748 nidulans 141
subcretacea 688,689*
ambigua 751 parasiticus 239
tatrensis 682,686
umbrina 688 brevicolumna 748,751 C lav aria 630, 634
utriformis 686 caespitosa 748,751 (also see Clavulina,
caudata 748,751 Clavulinopsis,
Camarophyllus 103
Cheilymenia Ramaria, etc.)
angelesianus 112
coprinaria 838 amethystina 641
angustifolius 109
crucipila 839 cineroides 640
borealis 109
stercorea 838 fumosa 638
cinereus 112
theleboides 838 juncea 637
colemannianus 108,112
Cheimonophyllum kunzei 643
cremicolor 109
candidissimus 132 mucida 636
graveolens 108
ornatipes 640
niveus 105,109 Chlamydopus 715
phycophila 636
pallidus 112 meyenianus 721
pulchra 638
pauper tinus 112 Chlorociboria 877
purpurea 637
pratensis 110 aeruginascens 878
rosea 635,639
rainier ensis 112 aeruginosa 878
recurvatus 112 rubicundula 638
Chlorophyllum 293 vermicularis 631*, 637
russocoriaceus 109 molybdites 295,895, zollingeri 635,638
subviolaceus 111 896
virgineus 105, 109 Clavariadelphus 632
Chlorosplenium 878 borealis 632,634
Cantharellula 149 Choiromyces 855 fistulosus 637
umbonata 151,165
alveolatus 858,859* ligula 633
Cantharellus 658 cookei 858 lovejoyae 632,634
appalachiensis 660
Chondrogaster 748 mucronatus 632,633
aurantiacus 480
Chondrostereum pistillaris 632
cibarius 659*, 662
purpureum 605 sachalinensis 633
cinereus 666
Choriactis subfastigiatus 633
cinnabarinus 664
geaster 827 truncatus 634
c lav at us 661
Christiansenia Claviceps 880
confluens 660,664
mycetophila 216 purpurea 880,895
floccosus 662
formosus 664 Chroogomphus 20*, Clavicorona 640
ignicolor 660,665 484 avellanea 642
infundibuliformis 665 Jlavipes 485 pyxidata 642
lateritius 660,664 leptocystis 485,487 taxophila 635,636
lutescens 665
940 GENUS AND SPECIES INDEX

Clavulina 640 Clitocybe (cont.) Collybia (cont.)

amethystina 640,641 graveolens 149, 152 distorta 216
cinerea 641 harperi 152, 160 dryophila 215
cristata 641 illudens 148 erythropus 215
ornatipes 640 infundibuliformis 157 extuberans 205,217
rugosa 641,642,643 * inversa 156 familia 215
zollingeri 638 irina 149, 155 fuscopurpurea 214
Clavulinopsis 634 leopardina 161 fusipes 204
appalachiensis 638 lignatilis 136 iocephala 202
aurantio-cinnabar ina martiorum 149 luxurians 205,209
635,639 maxima 157 maculata 217
corniculata 639 morbifera 163 marasmioides 205,215
dichotoma 640 morganii 480 oregonensis 218
fusiformis 635,638, multiceps 174 peronata 205,213
639* nebular is 159 platyphylla 146
gracillima 638 nuda 9, 10, 11,153 polyphylla 205,213
helvola 638 obsoleta 163 racemosa 212*, 213
holmskjoldii 640 odora 161 radicata 220
laeticolor 638 olesonii 154 spongiosa 205
luteoalba 638 oramophila 163 spp. (unidentified) 205,
miniata 638 polygonarum 152, 156 213
subaustralis 639 praemagna 155 strictipes 209
subtilis 640 rivulosa 163 subsulcatipes 203,218
umbrinella 635,640 robusta 150, 160 subsulphur ea 216
Climacodon saeva 149,154 trullisata 211
septentrionale 612, sclerotoidea 164 tuber osa 212
613* septentrionalis 151,159 umbonata 219
Clitocybe 148 sinopica 157 velutipes 220
aeruginosa 151,162 socialis 176 Coltricia 566
alba 160 squamulosa 150, 157 cinnamomea 568
albirhiza 161 suaveolens 162 montagnei 567,568
americana 150, 157 subalpina 156 perennis 567,568
atrialba 165 subconnexa 155 Coniophora
atroviridis 151,162 sudorifica 163 puteana 605,611
augeana 163 tarda 152 Conocybe 470,895
aurantiaca 480 variabilis 151,161 coprophila 471,472
avellaneialba 152, 161 Clitocybula 149 crispa 471,473
brunneocephala 154 abundans 150,204,215 cyanopus 471,472
candicans 161 atrialba 151,165 filar is 471,892
Candida 151,159 familia 150,204,215, intrusa 471
,
cerussata 159 161 898* lactea 472
clavipes 160,896 Clitopilus 239 smithii 471,472
coniferophila 152, 161 orcellus 241 sp. (unidentified) 471
crassa 151, 160 passeckerianus 240 stercoraria 471,472
cyathiformis 164 prunulus 240 tenera 472
dealbata 163,894 Collybia 201 Cookeina 836
deceptiva 162 abundans 215 Coolia
densifolia 155 acervata 215 odorata 198
dilatata 159 albipilata 212 Copelandia 358
ectypoides 157 alkalivirens 204
Coprinus 342
epichysium 151, 166 atrata 166 alnivorus 344,346
fasciculata 155 badiialba 216 alopecia 348
flaccida 157 bresadolae 215 americanus 344
fragrans 162 butyracea 216 arenatus 351
geotropa 158 cirrhata 212 asterophora 344
gibba 157 confluens 213 asterophoroides 344
gigantea 158
gilva 157
conigena 212 atramentarius 9 347 , ,
cookei 212 896
glaucocana 149, 152 distort a 216 bulbilosus 352
“chevicola” 351
GENUS AND SPECIES INDEX 941

Coprinus (cont.) Coriolellus (cont.) Cortinarius (cont.)

cine reus 351 heteromorphus 603 claricolor 442
colosseus 346 sepium 603 cliduchus 442
comatus 9, 56*, 343*, serialis 603 collinitus 431
345 suaveolens 598 corrugatus 420
disseminatus 352 Coriolus corrugis 423
domesticus 349 (also see Trametes) cotoneus 445
ephemeroides 352 versicolor 594 crassus 423,434
ephemerus 344,352 Corticium 604 croceofolius 427,454
fimetarius 344,351 crocolitus 419*, 423,432
Cortinarius 417
impatiens 344,353 crystallinus 428,429
acutus 427,452
ins ignis 348 cyanites 424
adustus 423,450
lag op ides 351 cyanopus 441
aggregatus 422,439
lag opus 350 cylindripes 430
albidus 423
macrocephalus 351 damascenus 428,451
alboviolaceus 447
macrorhizus 351 decipiens 427,453
amethystinus 447
micaceus 348 delibutus 420,431
amoenelens 441
miser 344,352 dibaphus 438
angulosus 426
narcoticus 351 dilutus 428,451
anomalus 425,448
niveus 343,351 distans 428,451
argent atus 425,448
palmeranus 346 duracinus 451
armeniacus 428,451
picaceus 346 elatior 421,432
arm Hiatus 448
plicatilis 352 elegantioides 441
atkinsonianus 440
pseudoradiatus 351 elegantior 441
aurantiobasis 454
quadrifidus 344 evernius 450
aureifolius 425,454
radians 344,349 fasciatus 452
aureofulvus 440
radiatus 351 flavifolius 426
balteato-cumatilis 434
semilanatus 351 flavovirens 441
balteatus 433
silvaticus 349 biformis 428,451 fragrans 425,447
spadiceosporus 346 bigelowii 443 fulgens 441
spp. (unidentified) 342, fulmineus 441
bivelus 451
350,351 bo laris 425,446 gentilis 444,896
sterquilinus 343,346 boulderensis 423,449 glaucopus 437
sulphureus 344 griseoluridus 420,431
brunneus 423,428,450,
tectisporus 351 griseoviolaceus 425,431
451
umbrinus 346 haematochelis 426,449
builliardi 450
variegatus 344 heliotropicus 430
bulbosus 451
xerophilus 344 hemitrichus 428
cacaocolor '428,451
Coprobia caerulescens 439
hercynicus 446
granulata 838 herpeticus 441
caesiifolius 447
Cordyceps 878,879 caesiocyaneus 422,439 humboldtensis 453
canadensis 881 caesiostramineus 438 immixtus 436
capitata 880 californicus 426,454 impennis 423,450
clavulata 880,882 callisteus 427,454 incisus 427,452
entomorrhiza 880,881 calochrous 422,441 infractus 435
gracilis 880,881 calyptratus 421,444 iodes 420,430
melolanthae 880,882 calyptrodermus 421, iodioides 430
militaris 882,883* 444 laniger 451
myrmecophila 881 camphor atus 425,447 largus 434
ophioglossoides 880, caninus 425,448 latus 442
881 castaneicolor 420,422, lucorum 424,450
ravenelii 880,882 431 lustratus 428
sphecocephala 880,881 cedretorum 439 luteoarmillatus 422,
unilateralis 880,882 cinnabarinus 426,455 436
washingtonensis 880, cinnamomeo-luteus 453 magnivelatus 442
882 cinnamomeus 453 malachius 425,448
Coriolellus 603 citrinifolius 421,436 marylandensis 455
alaskanus 603 metarius 441
citrinipedes 441
carbonarius 602, 603 michiganensis 422,439
clandestinus 445
942 GENUS AND SPECIES INDEX

Cortinarius (cont.) Cortinarius (cont.) Crepidotus 405

miniatopus 426,450 sterilis 420 applanatus 405,406
montanus 421,440 stillatitius 430 calolepis 406
mucifluus 431 subargentatus 425,448 cinnabarinus 405
mucosus 429 subcuspidatus 452 crocophyllus 405,406
multiformis 442 subflexipes 423,453 fulvotomentosus 406
mutabilis 437 subfoetidus 434 fusisporus 406
nigrocuspidatus 427, subpulchrifolius 425, herbarum 405
452 448 maculans 406
obtusus 452 sub purpurascens All mollis 406
occidentalis 437 subpurpureophyllus variabilis 405*, 406
odorifer 422,436 All versutus 406
olivaceopictus 453 subpurpureus 424,450 Crinipellis 202
olympianus 439 subtestaceus 426,449 campanella 210
orellanus 427,444, 896 superbus 422 piceae 210
orichalceus 421,441 thiersii 453 stipitaria 210
osmophorus 436 torvus 423,450 zonata 210
paleaceus 427,452 traganus 447 Crucibulum 778
pallidifolius 421,432 triformis 428,451 laeve 779
percomis 436 triumphans All lev is 779
phoeniceus 454 trivialis 431 parvulum 779
pholideus 424,428, turmalis 423,442 vulgar e 779
446 uliginosus 426 Cryptoporus 584
pinetorum 428,451 uraceus All,428 volvatus 585
plumiger 424,428,450 urbicus 424,450
Cudonia 872,893
ponderosus 432 vanduzer ensis 432 , cireinans 872*, 873
prasinus 421,441 433*
grisea 873
privignus 428,451 variicolor All, 442
lutea 873,874
psammocephalus 427, varius All, 442
monticola 873
452 velatus 420,443
velicopia All, 439
Cyathipodia 805
pseudoarquatus 438
pseudobolaris All verrucisporus 420,443 Cyathus 778
pseudosalor 420,430 vibratilis 429 helenae 780
pulchellus 423,453 violaceus 446 olla 779,780
puniceus 455 virentophyllus 421,441 pygmaeus 779,780
volvatus All, AAA stercoreus 780
purpurascens All
pyriodorus 447 striatus 779,780
washingtonensis All,
rainierensis 426,444 453 Cyptotrama
raphanoides 453 wiebeae 443 chrysopeplum 131
regalis 443 zakii 454 Cy st o derma 198
renidens 426 Coryne ambrosii 199
rubicundulus All sarcoides 877 amianthinum 200
rubripes 449 car char ias 199
Cotylidia 604
rufo-olivaceus 421,441 cinnabarinum 199,200
aurantiaca 608
salor 430 fallax 199
decolorans 608
sanguineus 454 granosum 199
diaphana 608
saturninus 450 granulosum 199,200
Craterellus 658
scandens 452 gruberianum 199,200
caeruleofuscus 666
scaurus 440
caly cuius 659,668 D
semisanguineus 426, cantharellus 664
454 Dacrymyces
cinereus 665
sodagnitus 438 cornucopioides 666, deliquescens 670,674
sp. (unidentified) 423, 898* palmatus 670,674
441
fallax 668 Daedalea 586
speciosissimus 427,444 foetidus 659,666 berkeleyi 587,588
sphaerosporus 431 odoratus 664 confragosa 589
splendidus 430 sinuosus 659,666 juniperina 587,588
squamulosus 445
Creolophus mollis 596
stemmatus 427,452
cirrhatus 612 quercina 587
unicolor 589
GENUS AND SPECIES INDEX 943

Daedaleopsis 586 Elaphomyces (cont) Fomes 574

ambigua 587,589 subviscidus 865 (also see Fomitopsis,
confragosa 586*, 588 variegatus 864 Ganoderma, Phellinus)
Daldinia 878,885 verrucosum 864 annosus 579
concentrica 887 Elasmomyces 736 fomentarius 575,581
grandis 887 camphorata 739 juniperinus 583
vernicosa 887 odoratus 737,738 officinalis 580
Dasyscyphus 877 pilosus 738 pinicola 579
Datronia 592 roseipes 738 subroseus 580
mollis 595 russuloides 737,738 Fomitopsis 574
Delastria 855 Elfvingia 577 cajanderi 580
Dentinum 616 Elvela 805 ellisianus 579
fraxinophilus 579
albidum 619 Endogone 844
albomagnum 619 officinalis 579
lactiflua 844
repandum 618 pinicola 578
Endoptychum 727
umbilicatum 617,619 rosea 580
agaricoides 731
Dermocybe 418 arizonicum 732
Fulvifomes
(also see Cortinarius) depressum 730 juniperinus 583
cinnamomea 453 Entoloma 238,242, Funalia
sanguined 455 895,896 hispida 568
trogii 598
Destuntzia 748 (also see
fusca 750 Leptonia, Nolanea) Fuscoboletinus 505
rubra 750 abortivum 242 aeruginascens 507
say lor ii 750 clypeatum 244 glandulosus -506
solstitialis 750 ferruginans 245 grisellus 506,507
subborealis 750 grayanum 244 ochraceoroseus 506
lividum 243,244 paluster 506,507
Dictyocephalos 711
madidum 243 serotinus 506,507
attenuatus 713
nidorosum 244 sinuspaulianus 506
Dictyophora 766 spectabilis 506,507
duplicata 768,771 nitidum 242,250
weaverae 506
indusiata 767*, 770 pernitrosum 245
multicolor 768,771 prunuloides 243,244 G
rubrovolvata 771 rhodopolium 243
Discina 797 sinuatum 244 Galera 399
ancilis 799 speculum 243,244 Galerina 399
apiculatula 799 strictius 246 autumnalis 401,892
leucoxantha 799 trachysporum 242,250 cedretorum 402
macrospora 799 vernum 248 corneipes 369
olympiana 799 violaceum 249 heterocystis 402
perlata 798 Exidia hypnorum 402
Disciotis 784,796 alba 671,673 marginal a 400,401,
venosa 796 glandulosa 672 892
nucleata 670,673 mutabilis 395
Disciseda 696
recisa 671,674 paludosa 400,401
a/er 698
semilanceata 402
brandegeei 698 F tibicystis 402
Candida 698
Favolus triscopa 402
luteola 698
alveolaris 563 venenata 400,401,892
pedicellata 698
subterranea 698 Fayodia 151 Galeropsis 733
anthracobia 166 angusticeps 735
Dissoderma
cucullata 735
paradoxum 198 Fischerula
polytrichoides 735
subcaulis 845
E Galiel/a
Fistulina 553
rufa 827
Eccilia 245,246,249 hepatica 552*, 553
pallida 554 Ganoderma 574
Echinodontium 612 adspersum 577
tinctorium 613 Flammula 407
annularis 577
Elaphomyces 862 Flammulina 202
applanatum 576
granulatus 864 velutipes 220
brownii 577
muricatus 863 Floccularia 189, 194
944 GENUS AND SPECIES INDEX

Ganoderma (cont.) Genea 849 Gomphus 658

curtisii 574,578 arenaria 851 bonari 661,662
europaeum 577 cerebriformis 852 clavatus 661
lucidum 577 compacta 849,850 foccosus 661,895
oregonense 575,578 gardneri 850 kauffmanii 660,662
tsugae 575,578 harknessii 850 pseudo clavatus 661
Gastroboletus 544 hispidula 851 Grifola 554
amyloideus 544,545 intermedia 851 frondosa 564
scabrosus 544 kraspedestoma 849, sulphurea 573
subalpinus 545 851 umbellata 556,564*,
suilloides 544,545 Geoglossum 866 565
turbinatus 544 affine 867 Guepirtiopsis
xerocomoides 545 alveolatum 867 alpinus 674
Gastrocybe 733 difforme 867 chrysocomus 674
lateritia 474,733,735 fallax 867 Gymnomyces 742
Gastrosporium glabrum 867 cinnamomeus 744
simplex 696 glutinosum 866 ferruginascens 743,744
Gautieria 746 intermedium 867 roseomaculatus 742,
Candida 747 nigritum 866,867 744
gautierioides 746,747 simile 867 socialis 743
graveolens 747 Geopora 841,846 Gymnopilus 407,895
monticola 747 arenicola 847,935* aeruginosus 409
morchelliformis 746, arenosa 846,847 bellulus 407,408
747 aurantia 846,847 favidellus 407,408
parksiana 746,747 clausa 846,847 fulvosquamulosus 407,
pterosperma 746,747 cooperi 846 410
Geaster 699 harknessii 847 harmoge 409
longii 847 junonius 411
Geastrum 699
magnata 847 liquiritiae 407,408
arenarium 705
pellita 846,847 luteocarneus 407,408
bryantii 702
campestre 701,705 Geopyxis luteofolius 409
coronatum 702 ("also see Tarzetta) parvis quamulosus 407,
delicatus 705 carbonaria 835,840 410
drummondii 700,705 vulcanalis 840 penetrans 407,408
fimbriatum 701,704 Gerronema 221 pulchrifolius 410
floriforme 700,705 Globifomes punctifolius 407,409
fornicatum 699*, 701 graveolens 567 sapineus 408
indicum 703 Gloeocantharellus sp. (unidentified) 408
limbatum 700,702, purpurascens 660 spectabilis 410
703* subspectabilis 411
Gloeophyllum 586
mammosum 700,705 saepiarium 588*, 590 terrestris 407,408
minimum 700,702 validipes 411
striatum 590
nanum 700,702 trabeum 586,590 ventricosus 411
pectinatum 702 Gloeoporus Gyrodon
pluriosteum 701,704 lividus 490
adustus 596
quadrifidum 700,702 merulioides 490
dichrous 552
recolligens 705 Glomus 844 Gyromitra 796,799,
rufescens 700,702
Gomphidius 481
893
saccatum 703 ambigua 803
glutinosus 482
schmidelii 702 brunnea 800,803
largus 482
sessile 704 californica 804
maculatus 482
smithii 702 caroliniana 800,802
nigricans 482
striatum 700,702 esculenta 801,893
oregonensis 482
triplex 703 fastigiata 801,803
rose us 483
umbilicatum 705 gz'gfls 800
smithii 482,483
xerophilum 702 infula 802,893
subroseus 483
Genabea 849 korfii 801
viscidus 486
cerebriformis 851 melaleucoides 799
Gomphogaster
fragilis 849,852 montana 801
leucosarx 733
spinospora 852 sphaerospora 800,804
underwoodii 803
GENUS AND SPECIES INDEX 945

Gyrophragmium 727 Helvetia (cont.) Hydnobolites 855

calif or nicum 727, 728 queletii 809 californicus 857
texensis 730 solitaria 809 cerebriformis 857
Gyroporus 510 stevensii 807,812 Hydnocystis
castaneus 510 subglabra 807,814 californica 847
cyanescens 510 sulcata 806, 815 Hydnopolyporus
purpurinus 510 villosa 805, 806, 810 palmatus 555,605
subalbellus 510 Hericium 611,613 Hydnotrya 848
abietis 614 cerebriformis 848,849
H
americanum 614 cubispora 848,849
Haematostereum coralloides 614 ellipsospora 825
sanguinolentum 606 erinaceus 615 michaelis 849
Hapalopilus lac inia turn 615 tulasnei 848,849
nidulans 568 ramosum 615 variiformis 848
Hebeloma 463,895, xveirii 614 yukonensis 849
896 Heterobasidion 574 Hydnotryopsis
albidulum 465 annosum 575,579 compactus 845
crustuliniforme 464, Heteroporus 554 setchellii 845
466* biennis 566 Hydnum 611,616
fastibile 466 Hirneola (also see Dentinum)
hiemale 465 auricula-judae 675 calvatum 621
insigne 465 Hirschioporus crassum 621
mesophaeum 465 abietinus 593 cristatum 617
sacchariolens 464,465 pargamenus 593 cyanellum 617,622
sarcophyllum 463,465 Hohenbuehelia 132 fennicum 620
sinapizans 465 angustatus 137 fuligineo-violaceum
strophosum 463,466 atrocaerulea 133, 137 617,622
syriense 463 geogenia 137 fumosum 618
Helvetia 796, 805, 893 mastrucatus 133, 137 fuscoindicum 621*, 622
(also see Gyromitra) petaloides 136 imbricatum 619
acetabulum 807 Humana laevigatum 618,621
albella 807, 811*, 812 hemispherica 839 martioflavum 621
albipes 813 repandum 618
Hydnangium 744
atra 813 rimosum 618,620
carneum 745
californica 804 scabrosum 620
roseum 745
compressa 811 stereosarcinon 618
soederstroemii 745
connivens 812 subincarnatum 617,
Hydnellum 622
corium 805,810 618,620
aurantiacum 626
costifera 806,808 caeruleum 625 Hygrocybe 103
crassitunicata 809 acuta 107,112
complectipes 626
crispa 816 acutoconica 115
concrescens 628
cupuliformis 806,810 albinella 105,109
conigenum 626
elastica 813 cruentum 624 atro-olivacea 112
ephippium 807,814 cumulatum 628 aurantiolutescens 116
fusca 814 cyanopodium 623,625 aurantiosplendens 115
griseoalba 806,808 diabolum 627 caerulescens 112
infula 803 caespitosa 105
earlianum 626
lac tea 816 calyptraeformis 117
ferrugipes 625,626
lacunosa 815 cantharellus 106,113
geogenium 623
leucomelaena 808 ceracea 115
mirabile 624
leucopus 812 chlorophana 106,115
nigellum 623,629
macropus 810 citrinopallida 115
peckii 627
maculata 814 coccinea 114
pineticola 624,627
mitra 815 conic a 116
piperatum 624
monachella 813 cuspidata 106,116
regium 623,625
pallidula 810 flavescens 115
scleropodium 623
palustris 815 flavifolia 115
scrobiculatum 627
pezizoides 807,814 spongiosipes 624 fornicata 109
philonotis 815 suaveolens 624 laeta 107,119
phlebophora 806, 816 subsuccosum 628 laetissima 106,115
zonatum 624,628 langei 116
marchii 114
946 GENUS AND SPECIES INDEX

Hygrocybe (cont.) Hygrophorus (cont.) Hymenogaster (cont.)

marginata 112 langei 116 subochraceus 749
miniata 113 laurae 107, 125 utriculatus 748,749
minutula 114 limacinus 127 Hypholoma 361,381
moseri 113 marginatus 113 (also see Naematoloma)
nigrescens 117 marianae 107 capnoides 383
nitida 106,115 marzuolus 129 dispersum 384
nit rat a 112 megasporus 127 fasciculare 383
olivaceoniger 117 miniatus 113 incertum 363
ovina 112 monticola 126 Hypomyces 878,882
parvula 106,115 morrisii 128 cervinigenus 883,884
persistens 116 nemo reus 108,111 chrysospermum 883
psittacina 118 occidentalis 127,128 hyalinus 883,884
punicea 114 odoratus 128 lactifluorum 884
pur a 104 olivaceoalbus 127 luteovirens 883,884
purpureofolia 112 pacificus 108, 126 transformans 883,884
reai 105,114 paludosus 107 Hypsizygus
ruber 105 penarius 123 tessulatus 133
singer i 117 perfumus 123
Hysterangium 762
splendidissima 115 persoonii 128
aureum 763
squamulosa 113 piceae 105, 120
clathroides 763
subaustralis 105, 109 ponderat us 104, 123
coriaceum 763
subminiata 106,113 pratensis 110
crassirhachis 763
subminutula 114 psittacinus 119
crassum 763
turunda 105,113 pudorinus 124
darkeri 763
unguinosa 106,119 puniceus 114
fischeri 764
virescens 118 purpurascens 124
fuscum 763,764
Hygrophoropsis 476 pusillus 110
occidental 763
aurantiaca 479 pustulatus 128
separabile 763
olida 476,480 pyrophilus 126
setchellii 763
Hygrophorus 103 recurvatus 112
sp. (unidentified) 763
(also see Camaro- roseibrunneus 125
stoloniferum 763
phyllus, Hygrocybe) russula 123
acutoconicus 116 saxatilis 108, 125 IJK
agathosmus 128 sordidus 122
albicastaneus 108, 125 speciosus 126 Ileodictyon
amarus 108, 124 subalpinus 109*, 121 cibarium 774
baker ensis 126 subpungens 125 Inocybe 455,894
borealis 109 subsalmonius 107 agardhii 462
brunneus 125 subviolaceus 112 albodisca 456,459
calophyllus 129 tennesseensis 125 bongardii 459
calyptraeformis 118 tephroleucus 128 caesariata 457,462
camarophyllus 107, 129 variicolor 107, 126 calamistrata 462
capreolarius 108, 124 vernalis 125 cookei 457,459
chrysaspis 121 vinicolor 126 corydalina 456,459
chrysodon 119 whiteii 121 fastigiata 456,457
coccineus 114 Hymenochaete flocculosa 457,462
conicus 117 rubiginosa 606 fuscodisca 456
cossus 121 tabacina 606 geophylla 460
discoideus 125 Hymenogaster 748 godeyi 459
eburneus 119 albellus 749 hirtella 459
erubescens 108, 124 albus 748,749 hystrix 457,462
flavescens 115 brunnescens 749 jurana 458
flavodiscus 121 diabolus 749 lac era 457,462
fuligineus 107, 127 gilkeyae 749 laetior 456,458
fuscoalbus 107, 127 luteus 749 lanatodisca 456,459
gliocyclus 120 mcmurphyi 749 lanuginosa 462
glutinosus 121 parksii 748, 749, 750* leucoblema 456
goetzii 108, 125 pyriformis 751 leucomelaena 457
hypothejus 126 ruber 750 lilacina 461
inocybiformis 107, 128 sublilacinus 749 maculata 458
kauffmanii 108 mixtilis 457,458
na pipes 457
GENUS AND SPECIES INDEX 947

Inocybe (cont.) Lactarius (cont.) Lactarius (cont.)

oblectabilis 456,458 alpinus 67,82 pseudo deceptivus 71
obscurioides 461 aquifluus 67 pseudo deliciosus 68
olympiana 457 argillaceifolius 63*, 76 pseudomucidus 77
picrosma 457 aspideoides 65,76 pubescens 41*, 73
pudica 460 atrobadius 80 representaneus 75
pyriodora 459 atroviridis 66,70 resimus 65,74
serotina 459 aurantiacus 79 riparius 80
sindonia 459 barrowsii 65, 69 rubrilacteus 68
sororia 457 caespitosus 76 rufulus 82
sp. (unidentified) 457, californiensis 75 rufus 79
462 camphoratus 81 salmoneus 65,68
suaveolens 456,459 cascadensis 75 sanguifluus 69
terrigena 457,462 chelidonium 65,69 scrobiculatus 73
Inonotus 566 chrysorheus 65,74 sordidus 70
andersonii 567 cinereus 77 speciosus 76
arizonicus 569 circellatus 67,77 subdulcis 82
circinatus 570 cocosiolens 67,79 subflammeus 79
cuticularis 569 controversus 70 subpalustris 75
dryadeus 567,569,570 corrugis 66,78 subplinthogalus 66
dryophilus 569 croceus 65,75 subpurpureus 65,69
hispidus 569 deceptivus 66,71 subserifluus 67,82
obliquus 567 deliciosus 68 substriatus 79
radiatus 569 fallax 77 subvellereus 66, 71
texanus 569 fragilis 80 subvernalis 66
tomentosus 569 fuliginellus 78 subvillosus 66,73
Irpex fumosus 78 subviscidus 80
lacteus 598 gerardii 78 thejogalus 67,82
mollis 601 glutigriseus 77 thiersii 82
glyciosmus 66 thyinos 68
Ischnoderma 573
griseus 67 tomentoso-marginatus
benzoinum 573
helvus 67 71
resinosum 573
hemicyaneus 69 torminosus 73
Ithyphallus 766
hepaticus 80, 82 trivialis 77
Jafnea
herpeticus 82 uvidus 75,897*
semitosta 834,840
hygrophoroides 67,78 vietus 76
Kuehneromyces hysginus 80 vinaceorufescens 74
mutabilis 395 indigo 69 volemus 78,79*
insulsus 72 xanthogalactus 75
L
kauffmanii 67,77 yazooensis 66,72
Laccaria 171 lignyotus 65,78 zonarius 72
altaica 172 luculentus 79 Laetiporus 572
amethystea 173 luteolus 66, IS persicinus 573
amethysteo-occidentalis maculatipes 75 sulphureus 572,895,
173 maculatus 75 896
amethystina 172 manzanitae 67,80 Lamprospora 839
bicolor 172, 173 mucidus 77 Lartgermannia
lac cat a 172 necator 70 gigantea 683
ochropurpurea 172, 173 neuhoffii 71
Laricifomes
ohiensis 172 occidentalis 67
officinalis 580
proximo 172 oculatus 67,82
Laternea
stria tula 172 olivaceoumbrinus 70
columnata 773
tortilis 172 olympianus 66,72
triscapa 773
trullisata 172, 173 pallescens 65,75
Laxitextum
Lacrymaria 366 pallidiolivaceus 66
bicolor 607
velutina 366 paradoxus 65,69
payettensis 66,72 Leccinum 536
Lactarius 63,64,895
peckii 67 aeneum 540
affinis 66
piper at us 71 alaskanum 541
allardii 67
psammicola 66, 72, 73 albellum 537, 542
alnicola 71
arbuticola 539
948 GENUS AND SPECIES INDEX

Leccinum (cont.) Lenzites 586 Lepiota (cont.)

arctostaphylos 540 (also see Gloeophyllum) scabrivelata 303
armeniacum 538,540 betulina 589 seminuda 306*, 307
atrostipitatum 538,541 saepiaria 590 sequoiarum 307
aurantiacum 537,539 Leotia 872 sistrata 307
541 albiceps 873 spp. (unidentified) 294,
“aurantioscaber” 538, atrovirens 873,875 305,307
539 chlorocephala 875 subincarnata 309
brunneum 538,540 lubrica 874 tinctoria 295,301
californicum 537,542 viscosa 874 “ tomentodisca” 307
chromapes 534 Lepiota 293 ventriosospora 310
cinnamomeum 538 acutesquamosa 294, Lepista 148
clavatum 538 303 (also see Clitocybe)
constans 538,540 americana 301 inversa 157
creiaceum 542 asperula 294,303 irina 155
crocipodium 539 atrodisca 304 nuda 153*,154
discolor 537,539,541 badhamii 301 saeva 154
fallax 539,541 barssii 303 sp. (unidentified) 154
fibrillosum 538,540 birnbaumii 302 tarda 152
griseonigrum 538,541 brebissonii 302 Leptoglossum 132
griseum 538,542 breviramus 302 Leptonia 248
holopus 537,542 brunnescens 305 asprella 252
idahoensis 540 bucknallii 295 carnea 250
incarnatum 538 cpstanea 307,892 convexa 250
insigne 540 castaneidisca 307 corvina 251
largentii 540 cepae stipes 301 cupressa 252
manzanitae 539 clypeolaria 309 cyanea 249,250
montanum 538,541 clypeolarioides 310 cyaneonita 250
ponder osum 538,540 cortinarius 309 decolor ans 251
potteri 538 cristata 306 diversa 251
roseofracta 542 decorata 307 exalbida 249,252
rotundifoliae 542 eriophora 303 fuligineo-marginata
rufescentoides 537 ex coriata 303 249,252
rugosiceps 539 felina 295,309 gracilipes 252
scab rum 541 flammeatincta 304 incana 249
snellii 538,541 flavescens 302 jubata 246
subalpinum 539,541 fragilissimus 302 nigra 249,252
testaceoscabrum 538, glatfelteri 306 nigroviolacea 250
541 helveola 309,892 occidentalis 249,250
vulpinum 538 hispida 303 parva 251
Lent aria 645 humei 294 porphyrophaea 249
byssiseda 644, 646, 649 josserandii 308,892 rectangula 251
pinicola 646,649 leucothites 300 rosea 249
Lentinellus 141 lilacinogranulosa 294 sericella 253
cochlea tus 142 longistriatus 302 serrulata 249,252
crinitis 133 lutea 302 undulatella 249,252
flabelliformis 144 luteophylla 295,302 vinaceobrunnea 252
montanus 144 molybdites 297 violaceonigra 250
omphalodes 142 morgani 297 zanthophylla 249, 251
ursinus 144 naucina 299,896 Leptopodia 805
vulpinus 144 naucinoides 300
Leucoagaricus 293
Lentinus 141 procera 298
(also see Lepiota)
detonsus 141 pulcherrima 291
naucinus 300
edodes 31*, 141 rachodes 297,896
procerus 298
kauffmanii 142 rhacodes 298
rachodes 298
lepideus 142 roseatincta 305
roseifolia 295,305
Leucocoprinus 293,
ponder osus 43*, 142*,
143 roseilivida 295,306,307 302
rubrotincta 305 (also see Lepiota)
sulcatus 142
sanguiflua 295 birnbaumii 302
tigrinus 141
GENUS AND SPECIES INDEX 949

Leucocoprinus (cont.) Lycoperdon (cont.) Marasmius (cont.)

brebissonii 302 pusillum 698 copelandi 207
breviramus 302 pyriforme 691 de lee tans 203,206
cepaestipes 302 rimulatum 694 epiphyllus 206
flavescens 302 subincarnatum 690 foetidus 203 .
fragilissimus 302 umbrinum 691,694 fulvoferrugineus 210
lilacinogranulosus 294 Lyophyllum 173 fuscopurpureus 214
longistriatus 302 atratum 166, 174 haematocephalus 210
luteus 302 carneum 176 iocephalus 202
Leucogaster 759 connatum 175 magnisporus 206
carolinianus 760 decastes 174,898* nigrodiscus 205,209
odoratus 760 infumatum 176 olidus 208
rubescens 760 lor ica turn 175 oreades 7*, 208
Leucopaxillus 166 montanum 46*, 175 pallidocephalus 208
albissimus 166*, 167 palustre 174 plicatulus 209
amarus 168,918* rancidum 174 prasiosmus 208
candidus 159 semitale 174, 176 quercophyllus 206
gentianeus 168 5/?. (unidentified) 175 rotula 203,206
giganteus 159 Lysurus 772 scorodonius 202,208
laterarius 168 borealis 778 siccus 204,210
paradoxus 168 cruciatus 111 sp. (unidentified) 206
septentrionalis 159 gardneri 778 strictipes 205,209
tricolor 167 mokusin 776 thujinus 208
periphragmoides 776 umbilicatus 203
Leucophleps 759
urens 213
magnata 760
spinospora 760
M Martellia 742
boozeri 743
Leucoscypha Macowanites 736
brunnescens 743
rutilans 837 alpinus 738
californica 743
Limacella 262,291 americanus 737
cremea 742,743
glio derma 291 chlorinosmus 737,738
ellipsospora 743
glischra 291,292 iodiolens 737,738
fallax 743
guttata 291 luteolus 737,738
foetens 743
illinita 292 magnus 737,738
parksii 743
kauffmanii 291,292 roseipes 738
lenticularis 291 subolivaceus 738 Melanogaster 756
roseicremea 291,29 2 subrosaceus 738 ambiguus 759
solidipes 291,292 euryspermus 758
Macrocystidia
intermedius 759
Longia cucumis 131
macrocarpus 759
texensis 730 Macrolepiota 293 parksii 758,759
Longula 727 (also see Lepiota)
variegatus 757*, 759
texensis 729 procera 298
Melanoleuca 169
Lycoperdon 677*,690 rachodes 298
alboflavida 169
americanum 691,694 Macropodia 805 brevipes 170
candidum 695 Macroscyphus 805 co gnat a 170
coloratum 698 Macrotyphula 634 evenosa 171,897*
curtisii 695,696 fistulosa 635,637 graminicola 169, 170
echinatum 694 juncea 636 lewisii 169
foetidum 692 Marasmiellus 202 melaleuca 169
gemmatum 694 albuscorticis 206 polioleuca 170
hie male 696 candidus 206 subalpina 171
marginatum 694 nigripes 203,206 vulgaris 170
mo//« 694
Marasmius 201 Melanophyllum
muscorum 694
albogriseus 204,209 echinatum 317
nettyana 691,695
alliaceus 208 Melanotus
nigrescens 693
androsaceus 208 textilis 405
peckii 691,694
bellipes 210
pedicellatum 692 Melastiza
borealis 210
perlatum 693 chateri 834,839
candidus 206
pulcherrimum 694 Meripilus
capillaris 208
giganteus 555,565
cohaerens 204
 950 GENUS AND SPECIES INDEX

Merulius Mycena 224 Mycena (cont.)

incarnatus 605,611 abramsii 235 sanguinolenta 232
tremellosus 605, 610*, 611 acicula 228 scabripes 233
Microcollybia 202,212 adonis 228 spp. (unidentified) 204,
albidula 227 209,226
Microglossum 868
atropurpureum 867, alcalina 234 stannea 235
869 amabilissima 226,228 strobilinoides 228
fumosum 867 amicta 226,231 stylobates 227
olivaceum 867,869,870 atroalboides 226,233, subcaerulea 226,231
rufum 871 235 subcana 233
viride 870 aurantiomarginata sub sanguinolenta 225,
Micromphale 202 225,228 232
arbuticola 202,208 capillaripes 229 /enax 237
foetidum 203 capillaris 227 tenerrima 227
penetrans 208 citrinomarginata 225, viscosa 237
sequoiae 204, 208 229 vitilis 236
clavicularis 237 vulgaris 225,237
Microporellus
clavularis 227,229* Mycenastrum 680
dealbatus 563
corticola 225,227 corium 689
obovatus 563
delicatella 227 Mycolevis
Microstoma
elegans 229 siccigleba 741
floccosa 835, 836
elegantula 226,230
protacta 835, 836 Mycorraphium
epipterygia 237
adustum 612
Mitrophora epipterygioides 237
semilibera 792 Myriosclerotinia 877
fibula 221
Mitrula 868 fHopes 235
Myriostoma 699
abietis 870 coliforme 704
galericulata 235
borealis 870 galopus 232 Myrmecocystis
elegans 869, 870 griseoviridis 225,237 cerebriformis 852
gracilis 870 haematopus 231 Myxomphalia 149
irregularis 871 ignobilis 227 maura 165
lunulatospora 870 inclinata 235
N
paludosa 870 iodiolens 234
Mollisia 877 juncicola 227 Naematoloma 381
Montagnea 727 laevigata 234 aurantiaca 382
arenarius 727 latifolia 235 capnoides 383
Morchella 784, 785 leaiana 225,236 dispersum 383,384
angusticeps 791 leptocephala 226,234
conica 791 lilacifolia 236,237* elongatum 384
luteopallens 226,228 ericaeum 384
crassipes 787, 788, 792*
maculata 235,236* fasciculare 382, 896
deliciosa 788*, 789
madronicola 225,227 myosotis 384
elata 31*, 786*.790
metata 234 olivaceotinctum 384
959*
monticola 228 polytrichi 384
esculenta 787
murina 234 radicosum 383
hybrida 792
occidentalis 226,234 squalidellum 384
semilibera 791
olivaceobrunnea 229 sublateritium 381,382
5/?. (unidentified) 787,
oregonensis 228 subviride 383
789
osmundicola 227 udum 384
Morganella 690 over holt sii 226,234
subincarnata 690 Nannfeldtiella
parabolica 234 aggregata 828
Mucilopilus paucilamellata 227 Naucoria 399
conicus 533 pelianthina 226,231 semiorbicularis 469
Mucronoporus pseudotenax 235 vinicolor 404
tomentosus 570 pur a 230 Neobulgaria
Multiclavula 634 purpureofusca 229 pura 875,877
mucida 636 rorida 237
Neocudoniella
vernalis 636 rosella 226, 229
albiceps 873
Mutinus 766 rubromarginata 230
Neogyromitra
bovinus 771 rugulosiceps 235
gigas 801
caninus 771 rutilantiformis 231
curtisii 771 sanguinolenta 232
elegans 768, 771 scab ripes 233
GENUS AND INDEX 951

Neolecta 868 Omphalotus 146,894, Panaeolus (cont.)

irregularis 871 896 solidipes 355
vile llina 871 illudens 148 sphinctrinus 357
Neosecolium 727 olearius 147, 148 subbalteatus 358,895
macrosporum 732 olivascens 147 tropicalis 358
Neottiella Onnia Panellus 132
rut Hans 837 tomentosa 570 longinquus 133
Neournula Osmoporus rams 132
pouchetii 827 odoratus 586,590 nidulans 141
Nidula 778 Osteina r ingens 132
Candida 780 obducta 555 serotinus 137
niveotomentosa 779, osrea 555 stipticus 138
781 O tide a 831 Panus 132
Nidularia 778 abietina 832 conchatus 138
far eta 779 alutacea 832,833* crinitis 133
pulvinata 779 auricula 832 dryinus 136
Nivatogastrium 733 bufonia 832 operculatus 132
nubigenum 735 cantharella 832,833 rwcfo 139
w right ii 736 concinna 832,833 strigosus 140
grandis 832 torulosus 138
Nolanea 245
cuneata 248 leporina 832,833 Paxillus 476
edulis 247 onotica 832 atrotomentosus 478
rainierensis 832 involutus 41*, 477, 896
fructufragrans 245,
smithii 832 panuoides 476
248
Oudemansiella 202 vernalis 478
hirtipes 246,247
holoconiota 248 longipes 220 Paxina 805
icterina 245,248 platyphylla 146 acetabulum 807
mammosa 248 radicata 218*, 219 leucomelas 809
murraii 245 Oxyporus recurvum 799
papillata 248 nobilissimus 549,575 Penicillium 782
salmonea 245 populinus 575 Peniophora 604
sericea 246 gigantea 607
staurospora 246,247, P
Perenniporia
248 subacida 603
Pachyella 818, 822
stricta 246 Peziza 818
babingtonii 819,822
verna 247 clypeata 819,822 (also see Pachyella,
Nyctalis 200 Pachyphloeus 855 Plicaria, Geopyxis,
citrinus 856 Tarzetta)
O ammophila 819,826
conglomeratus 857
Octavianina 744 melanoxanthus 855, W/a 819,821
asterosperma 745 857 badioconfusa 821,822
macrospora 746 virescens 857 brunneoatra 820
papyracea 746 Panaeolus 353,895 cerea 821
rogersii 746 acuminatus 354,357, domiciliana 822
echinospora 821
Omphalia 221 360
ellipsospora 822*, 825
Omphaliaster 151 cambodginensis 358
emileia 822
asterosporus 166 campanulatus 356
fimeti 823
borealis 166 castaneifolius 360
gautierioides 825
Omphalina 221 cyanescens 358,895
melaleucoides 799
chlorocyanea 162 fimicola 354,357
petersii 821
chrysophylla 221 foenisecii 360
praetervisa 824
epichysium 166 papilionaceus 357
proteana 824
ericetorum 223 phalaenarum 355
retirugis 354,357 pustulata 821
fibula 221
rickenii 357 repanda 821
grossula 162
semiovatus 355,357* sp. (unidentified) 822*,
hudsoniana 223
separatus 355,357* 824
luteicolor 221,223
sepulchralis 355 stuntzii 825
postii 223
pyxidata 223
strombodes 221
umbellifera 223
wynniae 162, 221
952 GENUS AND SPECIES INDEX

Peziza (cont.) Phellinus (cont.) Pholiota (cont.)

succosa 819,821 nigrolimitatus 583 lubrica 392
sylvestris 820*, 821 pini 582 malicola 388
varia 822 pomaceus 575,581 multifolia 386,388
venosa 796 rimosus 575,582 mutabilis 395
vesiculosa 823 robiniae 582 myosotis 384,387
violacea 824 robustus 575,582,583 polychroa 387
Phaeobulgaria taxodii 583 praecox 469
inquinans 876 texanus 583 prolixa 388
Phaeocollybia 413 torulosus 583 scamba 387,394
attenuata 414,415 tremulae 581 sp. (unidentified) 385
californica 415 Phellodon 622 spectabilis 411
christianae 416 atratus 629 spinulifera 388
deceptiva 414,415 confluens 624,628 spumosa 394
dis si lie ns 415 melaleucus 623,629 squarrosa 389
fallax 413,414 niger 623,629 squarroso-adiposa 391
festiva 413,414 tomentosus 628 squarrosoides 385*,
gregaria 414,415 Phellorina 715 386,390
jennyae 416 inquinans 723 subangularis 394
kauffmanii 416 strobilina 723 subcaerulea 380,385
laterarius 414,415 Phillipsia 836 sublubrica 393
lilacifolia 414,415 Phlebia subochracea 388
olivacea 414 merismoides 610 terrestris 389
oregonensis 416 radiata 610 velaglutinosa 387,393
piceae 415 vermiflua 470
Phlegmacium 418
pseudofestiva 413,414 vernalis 387,396
Phlogiotis
radicata 414,416 Pholiotina
helvelloides 672
rufipes 416 fHaris 472
scatesiae 414,415 Pholiota 384
Phylloporus 476
similis 414,415 abietis 391
arenicola 480
sipei 416 adiposa 391
rhodoxanthus 480
spadicea 414,416 albivelata 380,387
albocrenulata 392 Phyllotopsis 132
Phaeolepiota 411 nidulans 140
aurea 412 alnicola 388
astragalina 387 Physalacria
Phaeolus 566 inflata 548
aurea 413
alboluteus 571
aurivella 390 Pico a 852
fibrillosus 567,572
aurivelloides 391 carthusiana 854
schweinitzii 570
brunnescens 393 Piersonia
Phaeomarasmius caper at a 412 alveolata 858
confragosus 404 carbonaria 385,394 bispora 858
erinaceellus 386 confragosa 404 Piptoporus 584
Phallogaster 762 connata 391 betulinus 584
saccatus 762,764 decorata 387,393 Pisolithus 711
Phallus 766 destruens 395 arena rius 713
hadriani 769*, 770 elongatipes 384,387 arrhizus 713
imperialis 770 erinaceella 386 tinctorius 111
impudicus 768 ferruginea 393 Pithy a
iosmus 770 ferrugineo-lutescens cupressina 836
ravenelii 768,770 393 vulgaris 834,836
rubicundus 768,770 fibrillosipes 388
Plectania 826
Phellinus 574 filamentosa 391 coccinea 836
chrysoloma 583 flammans 391 hiemalis 836
everhartii 575,582, flavida 388
melaena 827,831
583* fulvosquamosa 381
melastoma 829
ferreus 575,582 fulvozonata 394 milleri 830
ferruginosus 575,582 graveolens 394
nannfeldtii 830
gilvus 582 hiemalis 391
nigrella 830
igniarius 581 highlandensis 385,394
laevigatus 575,581 lenta 387,393 Pleurocybella
limonella 391 porrigens 136
GENUS AND SPECIES INDEX 953

Pleurotellus Polyozellus 658 Psathyrella (cont.)

porrigens 136 multiplex 668 atrofolia 365
Pleurotus 132 Polypilus bipellis 362
(also see Hohen- frondosus 565 candolleana 363
buehelia) Polyporus 554 canoceps 366
candidissimus 132 (also see Albatrellus, carbonicola 366
columbinus 134 Meripilus, Osteina, circellatipes 364
cornucopiae 134 Ischnoderma, etc.) conissans 362,364
corticatus 136 arcularius 563 conopilea 365
dryinus 135*, 136 badius 42*, 562 elwhaensis 365
elongatipes 133 betulinus 585 epimyces 361
lignatilis 136 biennis 566 fuscofolia 364
ostreatus 133*, 134 brumalis 555,563 gracilis 365
petaloides 137 de cur re ns 561 hydrophila 364
porrigens 135 elegans 562 hymenocephala 363
sajor-cajou 134 fagicola 562 incerta 363
sapidus 134 frondosus 565 kauffmanii 362
ulmarius 133 hirtus 560,561* lacrymabunda 366
Plicaria 818 lentus 562 longipes 364
carbonaria 820 mcmurphyi 561 longistriata 362
endocarpoides 820 melanopus 563 maculata 366
leiocarpa 820 mori 555,563 multipedata 361,364
trachycarpa 820 mylittae 564 5/?. (unidentified) 362,
picipes 562 366
Plicaturopsis
radicatus 556,564 spadicea 362,364
crispa 591
squamosus 556,561 stercoraria 362
Pluteolus
sulphureus 573 sublateritia 362,364
aleuriatus 475
tuberaster 563 subnuda 365
callisteus 475
umbellatus 565 uliginicola 362,363*
Pluteus 253,254 velutina 366
varius 563
admirabilis 254,257
versicolor 594 Pseudobalsamia
atricapillus 256
Polysaccum alba 853
atromarginatus 254,
pisocarpium 713 magnata 853
256
nigrens 853
aurantiorugosus 254, Polystictus
tomentosus 570 Pseudocolus 772
257
versicolor 594 fusiformis 772,776,
californicus 258
111*
cervinus 255 Poria 602
javanicus 776
chrysophaeus 258 cocos 604
schellenbergiae 776
coccineus 257 corticola 603
cyanopus 254,258,895 incrassata 603 Pseudocoprinus
flavofuligineus 258 sequoiae 603 disseminatus 353
granularis 258 spissa 602 impatiens 353
leoninus 254,257 vaillantii 603 Pseudocraterellus
longistriatus 257 xantha 603 pseudoclavatus 661
lutescens 257 Porodisculus sinuosus 666
magnus 256 pendulus 552 Pseudofistulina
nanus 257 Poronidulus radicata 554
pellitus 254,256 conchifer 552 Pseudohydnum
petasatus 255 Porphyrellus gelatinosum 671
salicinus 254, 258, 895 pseudoscaber 534 Pseudopithyella
seticeps 258 Pouzarella 238,239 miniscula 836
Podaxis 715,725 Protogautieria Pseudoplectania
argent inus 726 lutea 741 melaena 831
longii 726 substriata 741 nigrella 830
microsporus 726 Psilocybe 367,368,895
Psalliota 310
pistillaris 724,725 angustispora 370
Psathyra 361
Podostroma 879 atrobrunnea 369
Psathyrella 361
alutaceum 879 baeocystis 372,895
ammophila 361
atrofolia 365
954 GENUS AND SPECIES INDEX

Psilocybe (cont.) Ramaria (cont.) Ramaria (cont.)

caerulescens 369,374, acrisiccescens 648, secunda 656
895 651 strasseri 648,656
caerulipes 369,373 amyloidea 646 stricta 631*, 648
californica 369 apiculata 646,649 stuntzii 647,655
castanella 369 araiospora 655 subbotrytis 647,655
coprophila 370 aurantiisiccescens suecica 649
corneipes 369 647,652 synaptopoda 647,653
cubensis 373,895 aurea 648,652 testaceoflava 647,654
cyanescens 371,895 botrytis 656 velocimutans 646
merdaria 369,370 botrytoides 648,656 vinosimaculans 648,
mexicana 373 brunnea 654 653
montana 369 cacao 646 Ramariopsis 640
pelliculosa 369,371 cartilaginea 648,655 californica 643
semilanceata 370,895 caulifloriformis 648 kunzei 643
silvatica 371 cedretorum 651 pulchella 641
squamosa 381 celerivirescens 646 Resupinatus
strictipes 372 claviramulata 647 applicatus 132
stuntzii 372 conjunctipes 641
Rhizina
tampanensis 374 cyaneigranosa 647,
inflata 797
umbonatescens 376 654,655
undulata 797
Pterula 631 cystidiophora 647,653
Rhizopogon 753
Ptychoverpa fennica 650
amyloideus 756
bohemica 794 flaccida 649
atroviolaceus 754
Pulcherricium flava 652,653
cokeri 754,755
caeruleum 605 flavigelatinosa 648,
colossus 756
655
Pulveroboletus 509 couchii 754
auriporus 509 flavobrunnescens
diplophloeus 758
647,652
hemichrysus 509 ellenae 755
ravenelii 509 formosa 654,896
evadens 754
fumigata 651
Pulvinula idahoensis 756
fumosiavellanea 651
archeri 838 maculatus 754,755
gelatiniaurantia 648,
carbonaria 838 occidentalis 754
655
Punctularia ochraceorubens 740*,
gelatinosa 655
strigoso-zonata 610 755
gracilis 649
Pustularia 840 parksii 754,756
grandis 647
Pycnoporellus 572 pinyonensis 754,755
invalii 649
Pycnoporus 592 roseolus 754
largentii 647,654
cinnabarinus 597 rube see ns 754
leptoformosa 654
sanguineus 597 smithii 754
longispora 654
subaustralis 754
Pyrofomes maculatipes 647,654
subcaerulescens 754,
juniper inus 583 magnipes 648,652
756
Pyronema murrillii 649
subsalmonius 754
omphalodes 835,838 myceliosa 649
succosus 754
obtusissima 647
QR truncatus 755
ochraceovirens 650
vinicolor 754
pusilla 649
Queletia 715 Rhodocollybia 202
rasilispora 652
mirabilis 722 butyracea 216
rubella 649
Radiigera 760 rubiginosa 653
maculata 217
atrogleba 761 Rhodocybe 240
rubribrunnescens
fuscogleba 761,762 aureicystidiata 241
647,654
taylorii 761,762 caelata 240,241
rubricarnata 654
Radulum rubrievanescens 648, mundula 240,241
orbiculare 610 656 nitellina 240,241
Ramaria 645,890* rubripermanens 656 nuciolens 241
abietina 650 sandaracina 647,648, roseiavellanea 241
acm 646,649 655 Rhodopaxillus
sanguinea 653 nudus 154
GENUS AND SPECIES INDEX 955

Rhodophyllus 238 Russula (cont.) Sarcosphaera 825

(also see Entoloma, in tegra 101 coronaria 826
Leptonia, Nolane a) krombholzii 85 crassa 825
sericellus 253 laurocerasi 93 eximia 826
strictior 246 lilacea 101 Schizophyllum 590
Rhodotus lutea 86,92 commune 590
palmatus 130 maculata 100 Schizostoma 715
Rhopalogaster maculosa 95 laceratum 716,722
transversarium 724 mairei 98 Scleroderma 707,896
Rickenella mariae 96 albidum 710
fibula 221 montana 86 arenicola 710
Ripartites 150 nigricans 85,90 areolatum 708
tricholoma 150 obscura 92 aurantium 708
Rozites 411 occidentalis 85,92 bovista 708,710
caperata 412 ochroleuca 85,92
cepa 709
olivacea 87, 102
Russula 63,83,896 citrinum 708
paludosa 92
abietina 101 flavidum 710
parazurea 87,95
adust a 85,91 floridanum 710
pectinata 94
aeruginea 95 furfurellum 711
pectinatoides 94
alachuana 96 fuscum 710
pelargonia 99
albella 96 geaster 710
placita 100
albida 96 hypogaeum 710
polychroma 101
albidula 96 /aeve 710
pue llaris 101
albonigra 89 lycoperdoides 708
pulverulenta 94
alutacea 102 macrorhizon 708
raoultii 97 michiganense 710
amoenolens 94
romagnesiana 88 polyrhizon 711
anomala 96
rosacea 99 polyrhizum 711
aquosa 99
sanguinea 100 reae 710
atrata 89
silvicola 98 texense 711
atropurpurea 85
sordida 89 verrucosum 708
bicolor 86
sororia 93 vulgar e 708
brevipes 87
5pp. (unidentified) 86,
brunneola 87,94 Sclerogaster 744
102,103
caerulea 101 columnatus 744
subalbidula 96
cascadensis 85,88 xerophilum 744
subfoetens 93
cerolens 94 Sclerotinia 877
subnigricans 90
chamaeleontina 101 Scutellinia
tenuiceps 102
claroflava 92 erinaceus 839
variata 86,94
compacta 85,91 scutellata 839
velenovskyi 100
crassotunicata 86,97 umbrarum 839
ventricosipes 93
cremoricolor 97
vesca 87,94 Scutiger
crenulata 97 ellisii 560
veternosa 102
crustosa 86,95 hirtus 560
virescens 95
cyanoxantha 94,897* pescaprae 560
xerampelina 83*, 102
decolorans 91
Sebacina
delica 88 S incrustans 609
densifolia 90
Sarcodon 616 Secotium 724
dissimulans 85,90
(also see Hydnum) agaricoides 732
emetica 97
flava 92 imbricatum 620 Sedecula 707
Sarcoscypha 834 pulvinata 707
foetens 93
foetentula 93 coccinea 836 Sepedonium
fragilis 98 floccosa 836 chrysospermum 884
fragrantissima 92 occidentalis 836 Sepultaria 846
gracilis 99 Sarcosoma 826 (also see Geopora)
gracillima 99 globosum 827,829 arenicola 847
granulata 94 latahensis 827,828 Serpula
grisea 87,95 mexicana 828 himantioides 611
lac ry mans 610
956 GENUS AND SPECIES INDEX

Setchelliogaster 733 Strobilomyces 543 Suillus (cont.)

tenuipes 733,735 confusus 542*,543 hirtellus 494
Simblum dryophilus 543 imitatus 497
sphaerocephalum 776 floccopus 543 kaibabensis 494,502
texense 776 strobilaceus 543 lakei 495
Simocybe 399 Strobilurus 202 luteus 500
centunculus 400 albipilatus 212 megaporinus 492,499
Skeletocutis conigenoides 202,211 monticolus 502
amorpha 552 kempt onae 211 occidentalis 494,502
Sparassis 657 lignitilis 212 pallidiceps 494,502
crispa 657 occidentalis 212 pictus 494,495
herbstii 657 trullisatus 211 pinorigidus 500
radicata 657 Stropharia 374 placidus 494,502
spa t hula t a 657 aeruginosa 380 ponderosus 492,496
Spathularia 868 albocyanea 380 proximus 497,506
clavata 872 albonitens 376 pseudobrevipes 500
flavida 871 ambigua 377 punctatipes 494,502
spathulata 872 aurantiaca 382 punctipes 493
velutipes 869,872 bilamellata 377 pungens 503
coronilla ill reticulatus 505
th ulariopsis
cubensis 374 riparius 492,499
velutipes 872
depilata 380 salmonicolor 500
Sphaerobolus 778 hardii 375,377 sibiricus 498
stellatus 781
hornemannii 379 sphaerosporus 492
Sphaerosoma 845 kauffmanii 380 subaureus 494
Sphaerozone 845 magnivelaris 378 sub luteus 493,500
Spongipellis melanosperma 375,377 subolivaceus 499
leucospongia 601 merdaria 370 tomentosus 504
pachydon 598,601 riparia 375,378 umbonatus 498
unicolor 601 rugoso-annulata 378 variegatus 505
Spongiporus semiglobata 376 volcanalis 493,501
leucospongia 601 siccipes 376 wasatchicus 502
Spragueola squamosa 375,381,382
irregularis 871 stercoraria 376 T
Squamanita 197 thrausta 382
Tarzetta
odorata 198 umbonatescens 375,
(also see Geopyxis)
paradoxum 198 376
bronca 835,840
umbonata 198 Suillus 491 catinus 835,840
Staheliomyces acerbus 494,504 cupularis 835,840
cinctus 768 acidus 493 rosea 835,840
Steccherinum albidipes 493,501
Tectella
adustum 612 albivelatus 492
pate liar is 132
americanus 494,499
septentrionale 612 Telamonia 418
borealis 493,501
Stephensia 845 Terfezia 855
brevipes 501
Stereum 604 arena ria 855
brunnescens 493,501
burtianum 608 Thaxterogaster 733
caerulescens 496
complicatum 606 pingue 734
castanellus 493
diaphanum 608 Thelephora
cavipes 494
fasciatum 606 laciniata 609
cothurnatus 500
gausapatum 606 multipartita 609
decipiens 494,495
hirsutum 605
elegans 497 palmata 609
lobatum 606
flavidus 498 spiculosa 609
ochraceoflavum 607
flavogranulatus 502 terrestris 608
ostrea 606
fuscotomentosus 504 vwr/w 609
rameale 606
glandulosipes 493,501 Toga ria
sanguinolentum 605,
granulatus 502 aurea 413
606
sericeum 607
grevillei 497 Trametes 592,603
helenae 494,498 (also see Coriolellus)
striatum 607
hirsuta 595
GENUS AND SPECIES INDEX 957

Trametes (cont.) Tricholoma (cont.) Tubaria (cont.)

mollis 596 matsutake 192 tenuis 403
occidentalis 550,595 myomyces 182 Tuber 841,854
pubescens 595 niveipes 181 aestivum 855
suave olens 598 nudum 154 besseyi 859
velutina 595 orirubens 182 californicum 860
versicolor 594 panaeolum 155 canaliculatum 856,859
Tremella pardinum 183,896 candidum 861*, 862
concrescens 670 personatum 154 citrinum 860
encephala 673 pessundatum 185,896 dryophilum 860
foliacea 673 platyphyllum 177, 179 gardneri 859
frondosa 673,674 ponder osum 191 gibbosum 858
fuciformis 669,671 populinum 185 harknessii 859,862
lutescens 674 portentosum 180 irradians 860
mesenterica 673 resplendens 183 levissimum 860
reticulata 644,670 robustum 189 linsdalei 859
Tremellodendron saponaceum 184 magnatum 854
candidum 644,670 scalpturatum 182 melanosporum 842*,
pallidum 640, 644, 670 sejunctum 180 854
schweinitzii 644 sordidum 152 monticola 860
Tremellodendropsis sp. (unidentified) 179, murinum 856,859
182 rufum 861
640
tuberosa 643 squarrulosum 178, 182 separans 859
subacutum 181 shearii 860
Tremellodon
sulphurescens 183 sphaerosporum 861
gelatinosum 671
sulphureum 179 texensis 856,859
Trichaptum 592 terreum 182
abietinus 593 Tulostoma 677* ,715
titans 178 berteroanum 720
biformis 592,593
ustale 179, 185 brumale 719
Trichoglossum 866 ustaloides 185
farlowii 868 campestre 720
vaccinum 186 cretaceum 716,720
hirsutum 867 venenata 183
velutipes 868 excentricum 720
virgatum 181 fibrillosum 720
Tricholoma 176, 189, zelleri 188 involucratum 720
896 Tricholomopsis 144 lysocephalum 721
acerbum 178,185 decora 145,146 macrocephalum 720
acre 178,182 edodes 141 meristostoma 720
aggregatum 174 fallax 146 opacum 720
albobrunneum 185 flammula 146 simulans 720
album 183 flavissima 146 striatum 720
argyraceum 182 platyphylla 146 Tylopilus 532
atroviolaceum 178, 181 rut Hans 145 alboater 532
aurantio-olivaceum 188 sulfur eoides 145, 146 ammiratii 533,535
aurantium 187 Trichophaea amylosporus 520,532
caligatum 192 abundans 838,840 atrofuscus 534
cheilolamnium 180 boudieri 834,840 badiceps 533
cingulatum 177
bullata 840 ballouii 533,534
columbetta 183
Trogia chromapes 533
equestre 180
crispa 132,591 conicus 533
flavobrunneum 185
flavovirens 179
Truncocolumella 752 eximius 532
citrina 752 felleus 533,535
focale 189
rubra 748,753 ferrugineus 533
fulvum 185
Truncospora fumosipes 534
gambosum 183
demidoffii 575, 583 gracilis 533,534
georgii 183
Tubaria 399 humilus 535
imbricatum 186, 188*
confragosa 403 indecisus 535
inamoenum 179
furfuracea 402 inter me dius 533
leucophyllum 177, 180
pellucida 403 minor 533
magnivelare 191
nebulosus 534
manzanilae 185
958 GENUS AND SPECIES INDEX

Tylopilus (cont.) UV WXYZ

olivaceobrunneus 534
pacificus 534
Underwoodia Weraroa 733
columnaris 797 cucullata 734
peralbidus 533
plumb eoviolaceus 533, Ungulina Whetzelinia 877
534 marginata 579 Wolfina
porphyrosporus 534 Urnula 826 aurantiopsis 834
pseudoscaber 534,535* craterium 829 lEy/i/iea
rhoadsiae 533 geaster 827 americana 831
rubrobrunneus 533,535 hiemalis 829 sparassoides 817
snellii 532 pouchetii 827
Wynnella
sordidus 534 Vaginata silvicola 832
subunicolor 535 plumbea 288 Xerocomus 511
tabacinus 533,535 Vascellum 690 (also see Boletus)
Typhula 634,636 curtisii 691,695, 696 chrysenteron 520
juncea 636 depressum 696 subtomentosus 518
Tyromyces 597 lloydianum 696 Xeromphalina 221
albellus 599 pratense 695 campanella 222
amarus 601 Veluticeps cauticinalis 222
balsameus 599 berkeleyi 607 fulvipes 221,222
basilaris 598,599 Eerpa 784,793,893 kauffmanii 223
caesius 599 bohemica 793 orickiana 223
chioneus 599 conica 794 p/cta 222
floriformis 598,599 Vibressea tenuipes 221
fragilis 600 truncorum 869,870, Xerulina
galactinus 599 873 chrysopepla 131
guttulatus 599
Volvaria 258 Xylaria 878,885
immitis 599
Volvariella 253,258 cornu-damae 886
lacteus 599
bombycina 261 hypoxylon 885
leucospongia 600
hypopithys 260 longipes 886
mollis 598,600
parvula 260 polymorpha 886
perdelicatus 599
pusilla 259,260 Xylobolus 604
spraguei 599
smithii 261 Xylosphaera
spumeus 599
speciosa 259 hypoxylon 886
stipticus 599
surrecta 259, 262 polymorpha 886
tephroleucus 599
taylori 259,261
transmutans 600 Zelleromyces 742
villosavolva 259,261 cinnabarinus 742
unicolor 598,601
volvacea 258, 259, 262 gardneri 742
sp. (unidentified) 742

A species of Auricularia (probably A. auricula) growing on a log. Brown color and rubbery-
gelatinous texture are distinctive (see p. 675 for details).
 959

The author with some giant black morels (Morchella elata group, p. 790). (Joel Leivick)
South San
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