Ascomycete Fungi of North America: A Mushroom Reference Guide 9780292754539

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Ascomycete Fungi of North America

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NUMBER SIX T Y-NINE

The Corrie Herring Hooks Series

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ASCOMYCETE FUNGI

of North America A MUSHROOM REFERENCE GUIDE

Michael W. Beug Alan E. Bessette UNIVERSIT Y OF TEX AS PRESS

Austin

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Arleen R. Bessette

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Copyright © 2014 by Michael W. Beug, Alan E. Bessette, and Arleen R. Bessette All rights reserved Printed in China First edition, 2014 Requests for permission to reproduce material from this work should be sent to: Permissions University of Texas Press P.O. Box 7819 Austin, TX 78713-7819 htt p://utpress.utexas.edu/index.php/rp-form

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

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

Library of Congress Cataloging-in-Publication Data Beug, Michael W. Ascomycete fungi of North America : a mushroom reference guide / by Michael W. Beug, Alan E. Bessette, and Arleen R. Bessette. pages cm. — (The Corrie Herring Hooks series ; number sixty-nine) Includes bibliographical references and indexes. ISBN 978-0-292-75452-2 (cloth : alk. paper) 1. Ascomycetes—North America—Identification. 2. Mushrooms—North America—Identification. I. Bessette, Alan. II. Bessette, Arleen Rainis, 1951– III. Title. IV. Title: Mushroom reference guide. QK623.A1B48 2014 579.5′6—dc23 2013019658 doi:10.7560/754522

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Kit Scates Barnhart, 1937–2003

Ascomycete Fungi of North America is dedicated to the late Catherine “Kit” Scates Barnhart for her devotion to sharing, teaching, and making accessible her knowledge and love of mushrooms. Kit founded the North Idaho Mycological Association and was a North American Mycological Association (NAMA) Vice President and Regional Trustee, as well as Chair of the NAMA Education Committee. In 1974 she worked with Dr. Daniel E. Stuntz to form the Pacific Northwest Key Council. Kit was the events organizer, principal recruiter, and fi rst President. Kit and Dan were responding to the dearth of information that prevented amateurs from identifying mushrooms found in the Pacific Northwest. At that time there were no readily accessible keys for amateurs. Kit wrote the fi rst Key Council keys to the genus Ramaria, plus keys to the Boletes and keys to several small genera, including Gomphidius and Chroogomphus. She designed a picture key to the common genera of gilled mushrooms, and a picture key to mushroom terms. Both are great tools for beginning mushroomers. It was her teaching that inspired the picture key that is so central to this book. She also taught mushroom photography, creating another enduring legacy. Kit’s mentoring, generosity, and gentle prodding guided numerous mushroom enthusiasts ever deeper into the world of fungi, and inspired several to pursue graduate work in mycology. We, and the mushroom community in general, owe her a great debt of gratitude.

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Contents

Pr eface ix Acknowledgments xi Chapter 1. Introduction: Ascomycota—The Ascomycetes xiv Chapter 2. Key to Included Ascomycetes 13 Using the Keys to Included Ascomycetes 14 Picture Key to the Major Types of Included Ascomycetes 16 Chapter 3. Hypogeous Ascomycetes: The Truffles 70 Key to the Hypogeous Ascomycete Genera 73 Chapter 4. Pezizomycetes 117 Chapter 5. Sordariomycetes 273 Chapter 6. Leotiomycetes 353 Chapter 7. Eurotiomycetes 415 Chapter 8. Geoglossaceae 418 Chapter 9. Neolectomycetes 430 Chapter 10. Orbiliomycetes 433 Chapter 11. Dothideomycetes 436 Chapter 12. Taphrinomycotina 440 Glossary 445 R efer ences 457 Photo Cr edits 469 Index to Common Na mes 475 Index to Scientific Na mes 477

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Gyromitra cf. ambigua p. 150

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Preface

um erous field guides to the fungi of North America have been published. Approximately seventy-five percent of all of these described fungi belong to the phylum Ascomycota and are commonly referred to as the Ascomycetes. Ascomycetes are commonly encountered and collected, and exhibit a remarkable range of biodiversity. Many, to the authors at least, are beautiful and visually complex—providing stunning photographic opportunities. Some are highly sought-after edible species, such as morels and truffles. Others play significant roles in plant ecology through their mycorrhizal associations. It seems remarkable, then, that given the above, no single book has been published that collectively describes and illustrates these fungi for all of North America. Numerous books describing Ascomycetes have been published in Switzerland, Great Britain, and continental Europe. One book, by Tylutki (1993), is dedicated to the Discomycetes of Idaho and the Pacific Northwest. Two works by Fred J. Seaver (1978a, 1978b) describe and illustrate a significant number of North American Ascomycetes known at that time. Many of Seaver’s described species were not illustrated or were illustrated using only blackand-white photographs or color paintings. Technical literature pertaining to Ascomycetes is generally widely scattered and often difficult to access, and popular field guides typically include a relatively small number of the more commonly encountered species.

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Our goal in writing Ascomycete Fungi of North America: A Mushroom Reference Guide is to fi ll this void by providing a scientifically accurate guide dedicated to this very significant group of fungi. Realizing that it would be impossible to describe and illustrate the tens of thousands of species that occur in North America, we have focused on those species found in the continental United States and Canada that are large enough to be readily noticeable to naturalists, photographers, and mushroom hunters, and on those that reproduce sexually (teleomorphs) versus those that reproduce asexually (anamorphs). To develop our list of more than 600 species, we scoured dozens of field guides, the Pacific Northwest Key Council Matchmaker program, the Mushroom Observer website, the Champignons du Québec website, numerous other websites, and the scientific literature. A number of species are described and illustrated for the fi rst time in color, with the total number of North American species included in this work being substantially greater than previously published in any other single book dedicated to this group of fungi. In organizing the species descriptions, we have grouped all of the hypogeous ascomycetes (the truffles) together, including the genus Elaphomyces (the only truffle genus that is not a member of the Pezizales), followed by the epigeous ascomycetes of the order Pezizales. Sordariomycetes are described next, followed by Leotiomycetes, the third class of Ascomycetes with fruitbodies likely to attract the attention of mushroom hunters. Short chapters cover the Eurotiomycetes class, the Geo-

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glossaceae family (a group usually placed in Leotiomycetes, but more likely being a clade in Lichinomycetes or a class-level lineage in itself), the Neolectomycetes class, the Orbiliomycetes class, the Dothideomycetes class, and the subphylum Taphrinomycotina. We have included microscopic and other advanced information of interest to students and professional mycologists, while emphasizing macroscopic identification field characters for easier use by a general audience. We have generally used MycoBank to determine the current accepted name. When Species fungorum lists a different accepted name, we have asked the advice of specialists in the field and/or have listed both names separated by an equals sign. If the equals

sign appears in parentheses, we have concluded that the second name is a synonym of the fi rst. Additionally, a color key to the species described in this book provides the reader with a visual guide to assist in the identification process. In most instances, we have been able to provide different images in the Picture Key than those that appear with the species descriptions. Creating this book has been a joy and an education. We fi nd ourselves more appreciative of, and entranced by, the Ascomycetes than when we fi rst began this work. Their variability, adaptability, and complexity are worthy of wonder and continued study. Hopefully, you will agree.

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Acknowledgments

t is our ple asur e to ack now ledge the mycologists who advised us in the writing of this work, as well as the many photographers whose keen vision and skill created the beautiful images that are so central to it. Rosalind Lowen and Harold Larsen took great care in reviewing the entire manuscript. Gary Samuels reviewed the early draft of the chapter on Sordariomycetes; Rosanne Healy reviewed the fi rstdraft chapter on hypogeous fungi; Sean Abbott reviewed the Discinaceae; Todd Elliott reviewed the Cordycipitaceae; and Michael Castellano reviewed the genus Elaphomyces. Steve Trudell, John Plischke III, Scott Redhead, Lorelei Norvell, Renee Lebeuf, Fred Rhoades, Matt hew Smith, Nancy Weber, Matt Trappe, James Trappe, Richard Korf, Donald Pfister, Brenda Callan, Jonathan Frank, Cathy Cripps, Steve Zelski, Kerry O’Donnell, and Andrus Voitk have provided valuable mycological advice and counsel. Colleagues Ian Gibson and Danny Miller each helped improve the grammar, clarity, and accuracy of the fi nal draft . Philippe Clowez, Franck Richard, Regis Courtecuisse, Kerry O’Donnell, Karen Hansen, and Pierre-Arthur Moreau as a team reviewed the section on morels and provided invaluable information about the results of their forthcoming morel DNA research. We are indebted to the following individuals not only for their photographic talent, but also

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for the labor and time spent identifying and documenting their mycological specimens, a process often involving considerable research. Our gratitude goes out to photographers Sean Abbott , Harley Barnhart, George Barron, Michael Castellano, Adolf and Oluna Ceska, Jules Cimon, Todd Elliott , Jonathan Frank, Rosanne Healy, Roger Heidt, Bruce Horn, Richard Kay, Sava Krstic, Jacques Landry, Harold Larsen, Renee Lebeuf, Lawrence Leonard, David Lewis, Raymond McNeil, Ronald Meyers, James Murray, Andrew Parker, John Plischke III, Fred Rhoades, Bill Roody, Gary Samuels, Christian Schwartz, Noah Siegel, Dianna Smith, Hugh and Sandi Smith, Matt hew Smith, Roger Smith, Walt Sturgeon, R. Greg Thorn, James Trappe, Matt Trappe, Steve Trudell, Debbie Viess, Andrus Voitk, and Michael Wood. Bruce Horn’s image of Entonaema liquescens was provided courtesy of the University Press of Kansas. We are also grateful to Gerald Sheine for providing us with scans of the photos taken by the late Harold “Hank” Marshburn and the late Emily Johnson; Jerry’s dedication has helped to preserve the collections of numerous mushroom photographers and mycologists who are no longer with us. Permission to use the Hank Marshburn images was generously granted by the Mycological Society of America, current owners of the Marshburn collection. Permission to use the images of Emily Johnson was provided by Steven Stephenson, Department of Biological Sciences, University of Ar-

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kansas. Harley Barnhart generously provided the photo collection of the late Catherine “Kit” Scates Barnhart. A special debt of gratitude is owed to Hope Miller, who welcomed us into her home, allowed us to acquire the ascomycete book collection of Orson Miller Jr., and gave us Orson’s extensive ascomycete reprint collection. Steve Trudell and John Plischke III provided invaluable service in locating hard-to-fi nd references. Sean Abbott sent his reprints on the Discinaceae. Philippe Clowez made a gift of his monograph, Les Morilles: une nouvelle approche mondiale du genre Morchella. Gary Samuels

gifted his book, Fungicolous Fungi of the Mid Atlantic States, and located important references on the Sordariomycetes. Renata Brunner Jass, Victoria Davis, Lynne Ferguson Chapman, Kaila Wyllys, Regina Fuentes, and the staff at University of Texas Press provided invaluable assistance in the preparation of this work. Thanks to you all! Finally, without the guidance and support of our editor, Casey Kitt rell, this book as you see it would not have been possible. Casey’s vision enabled us to create a work that far exceeded our original conception.

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Ascomycete Fungi of North America

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Chapter 1 Introduction Ascomycota—The Ascomycetes

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pprox im ately 75 percent of all described fungi are classified in the phylum Ascomycota, commonly referred to as the Ascomycetes. Many hundreds of millions of years ago, the ancestors of today’s Ascomycota divided into two lineages, the Taphrinomycotina and the Pezizomycotina. Millions of years later, the Pezizomycotina divided to produce the Saccharomycotina, the ascomycete yeasts. All three of these suborders are estimated to have been in existence for more than 400 million years, possibly more than one billion years. The Pezizomycotina, with more than 32,000 described genera, contains all of the fi lamentous fruitbody-producing species except for one, Neolecta, which is now classified in the suborder Taphrinomycotina. Most of the fungi that combine with algae to form lichens, and the majority of fungi for which we have no evidence of sexual reproduction, are found in the phylum Ascomycota. However, neither lichens nor asexual fungi are the subject of this book. Though we have included a few spectacular asexual species, our focus is on the Ascomycota that form sexual reproductive structures and are large enough to att ract the notice of mushroom hunters. These can be saprotrophs, which obtain energy from dead organic matter; endophytes, which form a cooperative partnership with the leaves and stems of plants; mycorrhizae, those forming a partnership with plant roots; or parasites, plant pathogens, and animal pathogens that obtain nutrients from their host organisms. The Ascomycota are defi ned by the presence of an ascus, a round to cylindrical sac-like structure within which nuclear fusion and meiosis take place. Meiosis results in the formation of four haploid nuclei. One round of mitotic division typically follows meiosis and results in eight nuclei and, eventually, eight haploid ascospores in each ascus. Many of the Ascomycota that form showy fruitbodies can also reproduce asexually via conidia

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Choiroactis geaster

(asexual spores), which contain mitotic nuclei each with a cell wall that is simply a modified hyphal wall. The hyphae walls themselves are composed of chitin and beta glucans. Ascospores are often shot from the ascus by turgor pressure and dispersed by the wind. In some of the larger Ascomycetes, including Chorioactis geaster, the mass spore discharge looks like a puff of smoke and is accompanied by an audible hiss. Ascospores are also spread by splashing or running water, and by animals. Ascospores, once released from the ascoma, germinate to form a new mycelium. In hyphal species, the species that form visible ascomata (fruitbodies), the ascomata (singular = ascoma) may be closed (cleistothecium), open with a narrow orifice (perithecium), broadly open like a cup (apothecium), or exist as an undifferentiated mass of tissue or stroma bearing the asci in locules (an ascostroma).

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Perithecium of Sordaria superba open, showing asci and black ascospores (±50×)

Cleistothecium of Erysiphe open, showing protunicate ascus and spores (±100×)

Apothecia of Ascobolus minutus showing exposed asci fi lled with dark ascospores (±100×)

The hyphae are regularly septate with a simple septum that possesses a single pore. The “cells” of most Pezizomycotina possess two haploid nuclei, although numerous exceptions to this condition exist. Indeed, it is the multinucleate nature of the “cells” of hyphae in the morels that contributes to making morels so difficult to identify to species. Species can reproduce both by the sexual phase (teleomorph), involving meiosis and the production of an ascus, and by an asexual (anamorph) mitotic phase that typically involves the production of conidia from specialized conidiogenous cells. The relationships between teleomorphs and anamorphs frequently are unknown because the two phases may occur in different seasons or on different hosts. Therefore, historically the anamorph and the teleomorph each has had its own separate Latin

binomial name. However, new rules for botanical nomenclature were adopted in 2011 requiring just one name per species with the other name placed in synonymy. Consequently, where the connection has been well established, we give the anamorph names as well as the teleomorph names, since anamorph names will take precedence over teleomorph names where they are better known or economically important; otherwise the teleomorph names will have precedence. The asci of macroscopic, sexually reproducing members of the Pezizomycotina can be operculate, inoperculate, or protunicate. Operculate asci release their spores through a defi ned operculum formed terminally or subterminally at the ascus apex. They are only found in the class Pezizomycetes. Inoperculate asci are typically thin-

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Arthrobotrys, an imperfect (asexual or anamorph) genus showing hyphae and conidia (±150×)

Penicillium (asexual), brushlike conidiophore-producing conidia, with phloxine stain (±250×)

Sordaria (sexual or teleomorph) showing mycelial network dotted with perithecia (±25×) I n t roduct ion

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Operculate asci of Ascobolus showing “lid” open with spore emerging (±500×)

Inoperculate ascus tip of Sordaria showing pore stained red with congo red (±500×)

walled and release their spores through a pore or canal, by rupturing at the ascus apex or by disintegration of the ascus wall. Apothecial Leotiomycetes, cleistothecial Leotiomycetes and Sordariomycetes, and perithecial Sordariomycetes all have inoperculate asci. Protunicate asci are thin-walled, typically globose to broadly clavate, and passively release spores by disintegration of the ascus wall. Protunicate asci are produced by cleistothecial (chaemothecial) Eurotiomycetes and perithecial Sordariomycetes. The following information applies to teleomorphs, the sexual reproductive stage that commonly att racts the attention of mushroom hunters. Usually, the ascospores of a typical teleomorph are smooth, single-walled, elliptical, and uniseriate (lined up end to end in the ascus), with eight spores per ascus. However, because there are nu-

merous exceptions to this, positive identification of most ascomycetes requires the use of a microscope. For example, some spore walls are thin, while others are thick. Ascospores can be round to long and slender, develop internal septa as they mature, or even divide into part-spores while they are still in the ascus. Some develop ornamentation and/or color as they mature. Therefore, it is important to study mature spores obtained from spore prints and not rely on the observed shape or color of immature spores that are still in the ascus. Also it is important to study fresh material, since microscopic features can change in dried herbarium material (Baral 1992). Additionally, since spore ornamentation may be soluble in KOH, it is important to examine mature spores in water mounts as well as in 3 percent KOH solution. Some spores develop internal gutt ules (oil drops). The presence

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Protunicate ascus of Microsphaera stained red with congo red (±250×)

or absence of gutt ules, or their size and number if present, can also aid in identification. Other suggestions for observation methods in the study of Ascomycetes can be found in Weber, Trappe, and Denison (1997). For some species, using a scanning electron microscope to examine the spores is vital to understanding spore morphology and, ultimately, the relationships between species and genera. Rosanne Healy (personal communication) did a developmental study of spore ornamentation in the Pachyphloeus carneus clade (a type of truffle). She found that a covering, which eventually erodes away, develops over the spores and connects the spines at their tips. You can see the spines (“tent poles”) between the spore and the outer covering in the picture (p. 10, lower) where the spore is broken. Species in other clades of Pachyphloeus do not develop this covering. The presence or absence of paraphyses (ster-

ile elements between the asci), as well as the shape and length of the paraphyses, can also be useful features to take note of. Melzer’s reagent is very useful to have. In it, some spores and/or some parts of asci or paraphyses turn blue (an amyloid reaction), some turn reddish brown (a dextrinoid reaction), and others remain unchanged (an inamyloid or nonamyloid reaction). Hairs on the outside of the apothecium of many species provide important clues for identification. It is important to ascertain whether the hairs are rooted or superficial, whether they are septate or nonseptate, and if any part is covered with crystalline material. A more in-depth examination of Ascomycetes involves studying the hyphal structure of the ascomata as well as the development of the ascomata in culture. While we make occasional reference to both in some descriptions, this level of detail is generally beyond the scope of this work.

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Ascobolus cf. immersus, immature and mature thick-walled elliptical spores (250×)

Octospora axillaris with fusiform thin-walled ascospores (note operculate empty ascus) (250×)

Gyromitra ancilis, with pointed apiculi and gutt ules (250×) A scom ycete Fu ngi of North A m er ica

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Peziza phyllogena showing verrucose spore ornamentation (250×)

Aleuria aurantia, coarsely reticulate ascospores (250×)

Tuber oregonense, honeycomb ridges on spores in ascus (100×) I n t roduct ion

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Cercophora, globose spores in thick-walled ascus (400×)

Coccomyces dentatus, hooked paraphyses with a fi liform spore emerging at center (100×)

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Geoglossum, ascus with emerging septate spore (100×)

Elaphocordyceps segmented spores in ascus, also showing thickened tip of ascus (100×)

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Pachyphloeus cf. carneus spore with unbroken outer membrane (±4,500×)

Pachyphloeus cf. carneus spore with broken outer membrane revealing the spinulose spores (±2,300×)

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Peziza domiciliana with amyloid asci tips, inamyloid paraphyses (100×)

Aleuria aurantia, dextrinoid paraphyses (100×)

Byssonectria terrestris, inamyloid paraphyses (100×)

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Lasiobolus intermedius, pointed aseptate hairs (50×)

Lachnum septate hairs with granular encrustations at tip (100×)

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Chapter 2 Key to Included Ascomycetes

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Using the Keys to Included Ascomycetes We have been able to obtain images of just over 500 of the more than 600 species fully or partially described in this book. These images are the basis for our Picture Key, a unique variation on the traditional dichotomous key. The key itself is mostly dichotomous, with occasional sets of multiple leads, which are indicated by down-arrows. When keying mushrooms, selecting the appropriate lead to follow is not always easy. Furthermore, in many cases gett ing to a correct answer requires the use of a compound microscope to observe features that are not discernible to the naked eye, even with the use of a hand lens. However, we wanted the keys to be useable by someone without access to a microscope. Our answer was to use photographs whenever we felt that we had narrowed the choices to the point where further refi nement would have required difficult key lead choices or microscopy. Nevertheless, you still may reach a choice that is difficult to make. A useful strategy is to take note of the place where you are uncertain so you can return and try the other lead if your original path does not yield a convincing result. To facilitate backtracking, whenever a lead does not sequentially follow the previous entry, we have included in parentheses the number of the lead that led to the current one. No key is perfect, so it is important to remember that even when the key leads to a seemingly clear choice, comparison with the full description, comments, and photographs must be made before accepting the resulting identification. The Picture Key includes both fully described species and species mentioned only in the Comments sections of the descriptions. Whenever an image was available, we have placed the image of a species mentioned in a Comments section in the Picture Key. We did not always use the image of the fully described species in the key, knowing that

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the look-alike species in the key will lead the reader to its full description and picture. For species that are somewhat variable in appearance, particularly noteworthy, and/or are choice edibles or poisonous species, we have included multiple images in the Picture Key and/or have included one image in the key and a different image in the description. In this way, you can gain a fuller appreciation of the range of appearances you might encounter in a given species. However, it is impossible to represent the full range of changes in color and surface features seen as Ascomycetes age and weather. It is also important to remember that the mushroom you have found may be very rare, a previously undescribed species, or one that appears so infrequently that good photographs were not available and so is not included in the choices before you. A feature of the Picture Key is the inclusion of information about the known geographic range of each included species. Th is may help you narrow your choices when using the key. Because there is much yet to learn about the range and distribution of Ascomycetes, please use the symbols as a guide but not a strict rule as to where a specific mushroom can be found. In addition to the Picture Key, we have included just one dichotomous key to genera: one for the truffles. We have not included dichotomous keys for the other sections of the book because the complexity that would be required in those keys defeats our central purpose of creating a readily accessible resource.

P I C T U R E K E Y S H O R TCU T S

1. The truffles . . . . . . . . . . . . . . . . . . . . . leads 2 to 8 2. Other Ascomycetes . . . . . . . . . . .leads 9 to end 3. Fruitbody greater than 2 cm broad with a distinct stalk . . . . . . . . . . . . . leads 11 to 21 + 22b 4. Fruitbody less than 2 cm broad, with a distinct stalk . . . . . . . . . . . . . . . . . . . . . leads 22 to 36

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5. Fruitbody with no stalk or with a short to indistinct stalk a. Growing on another mushroom . . . . . . . . . . . . . . . . . . . . . . . . . . leads 38a to 38e b. Growing on burned ground, burned wood leads 41 to 44b c. Growing on dung or well-manured ground . . . . . . . . . . . . . . . . . . . . leads 46 to 47 d. Not as above, fruitbody gelatinous . . . . . . . . . . . . . . . . . . . . . . . . . . . .leads 49 to 50 e. Growing on water-soaked wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lead 52a f. Not as above, fruitbody greater than 2 cm broad or tall

K e y to I nclu de d A scom ycete s

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i. Fruitbody cup- to ear-shaped . . . . . . . . . . . . . . . . . . . . . . . . leads 59 to 62 ii. Fruitbody not cup- to ear-shaped . . . . . . . . . . . . . . . . . . . . . . . . leads 54 to 58 g. Fruitbody not as above, less than 2 cm broad or tall i. Fruitbody fleshy, globular to cupshaped to disk-like . . . . . . leads 67 to 73 ii. Fruitbody pustules that break through the bark of small branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lead 63c iii. Fruitbody none of the above . . . . . . . . . . . . . . . . . . . . . . . leads 64 to 66

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Picture Key to the Major Types of Included Ascomycetes

2b. Interior hollow or of thick folds of mushroom tissue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3

1a. Epigeous (above ground) Ascomycetes (see also 3b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9 1b. Hypogeous (below ground) Ascomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2

3a. Interior hollow or folds of tissue, usually buried at maturity, spores forcibly discharged or not . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. Interior hollow, opening to the surface at the top by splitt ing into rays, spores forcibly discharged, sometimes with an audible hiss

2a. Interior near or at maturity ± completely fi lled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5

Sarcosphaera coronaria W, NE p. 250

Geopora pellita QC p. 89

Peziza ammophila A (sand dunes) p. 204

Geopora arenicola NE + CO p. 89

Chorioactis geaster Texas p. 139

Geopora sepulta E p. 90

DISTRIBUTION NOTES USED IN CAPTIONS: A = widespread; N, S, E, W = region in North America; M = mountains; C = coast; B = boreal; MW = Midwest; ? = uncertain; state and prov-

ince abbreviations are standard except that NE = northeast, not Nebraska; MX = Mexico; WC = west coast; EC = east coast.

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4a. (3a) Interior hollow or with just a few folds of tissue

Gilkeya compacta WA to MX p. 93

Genea harknessii W p. 86

Gilkeya compacta WA to MX p. 93

Genea hispidula E p. 88

Genea arenaria WA to MX + NE p. 85

4b. Interior densely folded

Geopora cooperi AK to MX + MI p. 91

Genabea cerebriformis WA to MX p. 84

Hydnotrya cubispora A p. 96

K e y to I nclu de d A scom ycete s

Beug-final.indb 17

Hydnotrya variiformis BC to CA to CO p. 100

Peziza ellipsospora W p. 98

Hydnotrya cerebriformis W p. 98

17

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5a. (2a) Peridium thin . . . . . . . . . . . . . . . . . . . . . . . .7 T5b. Peridium thick, interior not marbled— Elaphomyces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Pachyphloeus thysellii WA, WI, + ? p. 108

Pachyphloeus carneus clade A p. 105

5c. Peridium thick, interior marbled— Pachyphloeus

Pachyphloeus citrinus A p. 107

Pachyphloeus virescens CA + ? p. 108

Pachyphloeus carneus clade A p. 105

Pachyphloeus austrooregonensis OR, CA p. 106

6a. (5b) Cortex (outer peridium) thin, black; inner peridium thick, white when fresh, gray or brownish tinged in age

Elaphomyces cf. loebae NC p. 80

Elaphomyces cf. leveillei NE p. 80

Elaphomyces cf. maculatus NE p. 80

Elaphomyces cf. anthracinus OR, ID, QB p. 80

Elaphomyces cf. decipiens NE, OR, CA p. 80

Elaphomyces macrosporus MI, MA, NC p. 80

A scom ycete Fu ngi of North A m er ica

Beug-final.indb 18

18

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T6b. Peridium differentiated into three layers, outer layer felty, purplish; inner peridium layer (bulk of peridium) texture punk-like,

Elaphomyces cf. asperulus A p. 82

Elaphomyces americanum E p. 79

light pinkish cinnamon . . . . . . . . Elaphomyces appalachiensis p. 81 or 296 6c. Not as above

Elaphomyces cf. granulatus A p. 82

Elaphomyces verruculosus E + MX p. 82

Elaphomyces cf. muricatus A p. 79

Elaphomyces subviscidus OR, ID, CO p. 82

7a. (5a) Gleba with brown or gray to grayish green to grayish blue pockets of spores

Imaia gigantea SE p. 102

Leucangium carthusianum West Coast p. 104

7b. Gleba not with obvious globular pockets of spores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8

Balsamia nigrans OR, CA p. 77

8a. Marbled with translucent wide bands, some cavities, fruitbody depressed-spherical

Balsamia magnata OR to CA to CO p. 75

K e y to I nclu de d A scom ycete s

Beug-final.indb 19

Kalapuya brunnea West Coast p. 104

Barssia oregonensis NW p. 76

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T8b. Not as above, gleba with labyrinthine veins, fruitbody often folded-lobed, outer surface almost smooth

Choiromyces alveolatus W p. 78

Choiromyces meandriformis OR, CA, WV p. 78

Hydnobolites californicus A p. 94

8c. Not as above, gleba veined, globose to lumpy fruitbody, surface smooth or warted—Tuber

Tuber oregonense BC to CA p. 111

Tuber sphaerosporum TN, OR p. 109

Tuber lyonii E p. 113

Tuber californicum W, OH p. 109

Tuber shearii A p. 115

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Tuber gibbosum BC to CA p. 111

Tuber candidum A p. 113

A scom ycete Fu ngi of North A m er ica

Tuber canaliculatum E p. 110

Tuber quercicola WC p. 114

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Tuber lyonii E p. 113

Tuber cf. separans A p. 116

9a. (1a) Fruitbody growing on another mushroom, or sessile or with a very short stalk relative to cap diameter . . . . . . . . . . . . . . . . . . .37 9b. Fruitbody with a distinct stalk . . . . . . . . . . . .10

Verpa bohemica A p. 267

10a. Fruitbody less than 2 cm broad. . . . . . . . . . . 22 10b. Fruitbody greater than 2 cm broad . . . . . . . . 11 11a. Head not skirt-like, not attached only at the top of the stalk . . . . . . . . . . . . . . . . . . . . . . . . . . .12 11b. Head skirt-like, att ached only at the top of the stalk

Verpa bohemica yellow form WA, ID + ? p. 267

12a. Texture fleshy, fruitbody ridged and pitted, sponge-like, hollow. . . . . . . . . . . . . . . . . . . . . . . 13 T12b. Texture fleshy, fruitbody shape various but not as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16 12c. Texture hard or tough and leathery, fruitbody shape various . . . . . . . . . . . . . . . . . . . . . 22b

Tuber cf. rufum CA to AB p. 114

Verpa conica complex A p. 268

13a. (12a) Ridges and pits randomly arranged, edges typically yellow at maturity, base of cap attached to stalk with litt le or no sinus— the yellow morels . . . . . . . . . . . . . . . . . . . . . . . . 15 13b. Ridges and pits generally arranged in a vertical fashion and edges typically blackening at maturity, base attached to the stalk with a small to deep sinus—the black morels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 14a. Appearing in burned areas one year after a fi re

K e y to I nclu de d A scom ycete s

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Morchella tomentosa NW to AK + YT p. 193

Morchella exuberans CA p. 194

Morchella cf. anthracophila W p. 194

14b. Not associated with a fi re or appearing more than one year after the burn

Morchella snyderi W p. 191

Morchella snyderi W p. 191

Morchella angusticeps E p. 181

Morchella elata A p. 184

Morchella angusticeps E p. 36

Morchella elata A p. 184

A scom ycete Fu ngi of North A m er ica

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Morchella snyderi W p. 191

Morchella brunnea W p. 192

Morchella cf. elata A p. 184

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Morchella fr ustrata NW p. 185

Morchella populiphila W p. 189

Morchella punctipes E p. 189

15a. (13a) Small (1–3 cm broad) with sparse ridges and pits (8–16 primary ridges)

Morchella cf. diminutiva E p. 182

Morchella cf. sceptriformis SE, VA to MS p. 182

15b. Large (1.5–13 cm wide) with numerous ridges and pits (12–30 primary ridges)

Morchella cf. americana A p. 179

K e y to I nclu de d A scom ycete s

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Morchella americana A p. 179

Morchella americana var. lata NE, p. 180

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Morchella ulmaria E p. 180

Morchella cf. prava (sand dunes) E + MI p. 187

Morchella ulmaria E p. 180

Morchella rufobrunnea CA to MX p. 190

16a. (12b) Head absent, or head a distorted cluster of cups, or cup-shaped, saddle-shaped, or lobed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18 16b. Head brain-like, sometimes saddleshaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17

Gyromitra esculenta A p. 149

Beug-final.indb 24

Morchella rigida CA, A? p. 180

17a. Stalk diameter much less than cap diameter, round to flattened but not deeply fluted or internally folded, not thin-fleshed

Gyromitra esculenta A p. 149

A scom ycete Fu ngi of North A m er ica

Morchella prava N, MI to BC p. 187

Gyromitra esculenta var. alba QB + ? p. 149

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Gyromitra infula A p. 150

Gyromitra ambigua A p. 150

Sphaeronaemella helvellae A p. 150

T17b. Stalk diameter about half of cap diameter, stalk deeply fluted, thin-fleshed

Gyromitra californica W p. 147

Gyromitra californica W p. 147

Gyromitra sphaerospora E p. 154

17c. Stalk diameter nearly equal to cap diameter, stalk greatly folded, not thin fleshed

Gyromitra montana W, M p. 153

Gyromitra brunnea E p. 145

Gyromitra montana W, M p. 153

Gyromitra brunnea E p. 146

K e y to I nclu de d A scom ycete s

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Gyromitra korfii E p. 151

Gyromitra caroliniana S + MI to OH p. 148 25

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18a. (16a) Stalk round to flattened or slightly ribbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 18b. Stalk distinctly ribbed to lacunose (see also 17b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19

Helvella lactea WA, MB, ON, p. 160

19a. Head absent or saddle-shaped to lobed

Helvella crispa N p. 160

Underwoodia columnaris MW p. 263

Helvella dryophila W p. 170

Helvella sulcata E p. 171

Helvella vespertina A p. 170

Helvella maculata W p. 167

A scom ycete Fu ngi of North A m er ica

26

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19b. (also 59a) Head cup-shaped

Helvella leucomelaena A, p. 165

Helvella acetabulum W, p. 155

Helvella solitaria N, p. 169

Helvella costifera A p. 155

Jafnea fusicarpa NE p. 173

Helvella unicolor AB p. 169

20a. (18a) Head cup-shaped . . . . . . . . . . . . . . . . . . .21 T20b. Head a distorted cluster of cups . . . . .Wynnea sparassoides p. 271

Helvella latispora E + M, p. 164

Helvella ephippium AB, MB p. 156

Helvella pezizoides A p. 168

Helvella elastica A, p. 162

Helvella compressa W, p. 158

K e y to I nclu de d A scom ycete s

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20c. Head saddle-shaped to lobed

Helvella atra N, p. 157

Helvella albella A p. 156

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21a. (20a) Colors white to brown, gray, or black

Helvella corium NW to NE, p. 159

Helvella fibrosa EC, WC, M, B p. 163

Helvella cupuliformis E, OR, TX p. 161

Pseudoplectania vogesiaca A p. 237

Helvella macropus A p. 166

Tarzetta catinus W p. 258

T21b. Brightly colored, stalk obvious

Cookeina sulcipes subtropical p. 140

Cookeina tricholoma subtropical p. 140

Microstoma protractum W p. 177

21c. Brightly colored, stalk often buried and not initially obvious

Sarcoscypha coccinea W p. 246

Sowerbyella rhenana A p. 256

A scom ycete Fu ngi of North A m er ica

Beug-final.indb 28

Sowerbyella radiculata E p. 255

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22a. (10a) Fruitbody less than 2 cm broad; not hard or tough and leathery (see also 19a, b; 20a, c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23

Xylaria polymorpha A p. 351

Xylaria multiplex A p. 349

Xylaria cubensis (club) + Xylocoremium flabelliforme E p. 348

Xylaria hypoxylon A p. 349

Xylaria liquidambaris E p. 350

Xylaria longipes E p. 351

Xylaria filiformis A p. 350

Xylaria magnoliae E p. 350

K e y to I nclu de d A scom ycete s

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22b. (12c and 10a) Fruitbody size various, hard or tough and leathery, exterior white, gray, black, or dark brown

Xylaria atropictor NW p. 348

Xylaria cornu-damae A p. 349

Xylaria oxyacanthae E p. 349

Xylaria tentaculata E p. 352

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23a. (22a) Growing on the ground, on wood, or on vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . .25 23b. Growing on an arthropod or on a buried truffle: the Cordycipitaceae family . . . . . . . 24

Elaphocordyceps capitata A p. 296

24a. Growing on buried Elaphomyces (Truffle) species

Elaphocordyceps longisegmentis E p. 296

Elaphocordyceps ophioglossoides A p. 298

24b. Growing on an arthropod

Cordyceps washingtonensis WA p. 290

Cordyceps militaris A p. 290

Cordyceps hesleri SE p. 289

Elaphocordyceps subsessilis A p. 299

A scom ycete Fu ngi of North A m er ica

Beug-final.indb 30

Cordyceps olivascens SE p. 290

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Akanthomyces aculeatus A p. 279

Beauveria bassiana A p. 284

Cordyceps tuberculata A p. 279

Beauveria bassiana A p. 284

Isaria farinosa A p. 280

Ophiocordyceps myrmecophila W p. 340

Beauveria bassiana A p. 284

Gibellula leiopus E p. 347

Ophiocordyceps dipterigena S p. 337

K e y to I nclu de d A scom ycete s

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Hymenostilbe sphingum A p. 280

Ophiocordyceps variabilis E p. 343

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Ophiocordyceps cf. gracilis A p. 337

Ophiocordyceps michiganensis E p. 341

Ophiocordyceps sphecocephala E p. 340

Ophiocordyceps super ficialis E p. 341

25a. (23a) Small stalked fruitbody not growing in deep moss or in bogs or water . . . . . . . . . . . . 26

Ascocoryne turficola NE p. 358

Bryoglossum gracile N p. 363

A scom ycete Fu ngi of North A m er ica

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Ophiocordyceps ravenelii E p. 339

Ophiocordyceps melolonthae E p. 338

Ophiocordyceps stylophora E p. 344

25b. Small stalked fruitbody on moss or watersoaked vegetation, in Sphagnum bogs, or in water

Cudoniella clavus N + CA p. 375

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Mitrula paludosa NE Canada p. 397

Mitrula elegans A p. 397

Gelatinodiscus flavidus NW p. 380

Roseodiscus subcarneus N p. 407

Vibrissea truncorum N + WV p. 414

Roseodiscus rhodoleucus N p. 408

26a. (25a) On soil, wood, vegetation, leaves, or cones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27

Onygena equina A p. 417

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Mitrula lunulatospora E p. 398

Neocudoniella radicella N p. 402

26b. On keratinous animal remains including hair, horns, hooves, and owl pellets

Onygena equina A p. 417

Onygena corvina A p. 417

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27a. (26a) Growing from a sclerotium

Sclerotinia veratri A p. 410

Dumontinia tuberosa A p. 378

27b. Not growing from a sclerotium . . . . . . . . . . 28 28a. Growing on soil . . . . . . . . . . . . . . . . . . . . . . . . . .32 28b. Growing on wood, cupules, cones, and leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

Heyderia abietis W + NE p. 383

Sclerotinia sulcata A p. 410

29a. Fruitbody cup-shaped to disk-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 T29b. Fruitbody club-shaped

Hypocrea alutacea A p. 302

29c. Fruitbody globose to convex (see also 25b)

Cudoniella acicularis E p. 374

A scom ycete Fu ngi of North A m er ica

Beug-final.indb 34

Neocudoniella albiceps E p. 375

Pachycudonia monticola W p. 403

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30a. (29a) Underside distinctly clothed in hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

Bisporella sulfurina BC + ? p. 362

Bisporella subpallida BC + ? p. 362

Kriegeria alutipes CA, NV, OR p. 380

Sarcoscypha occidentalis E p. 248

Lanzia luteovirescens W p. 409

K e y to I nclu de d A scom ycete s

Beug-final.indb 35

Bisporella citrina A p. 362

Hymenoscyphus virgultorum N + CO, CA p. 386

Peziza waltersii E p. 224

Crocicreas coronatum NF, QC p. 372

30b. Not as above, underside glabrous or minutely tomentose (see also 72b)

Bryoscyphus marchantiae W p. 386

Phaeohelotium epiphyllum A p. 404

Tatraea macrospora A p. 413

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Chlorencoelia torta E p. 366

1 (top) Holwaya mucida E p. 384 2 (bottom) Crinula caliciiformis E p. 384

Hymenoscyphus fr uctigenus NE p. 385

Chlorencoelia versiformis A p. 366

Urnula craterium E, MW p. 264

Chlorociboria aeruginosa A p. 367

A scom ycete Fu ngi of North A m er ica

Beug-final.indb 36

Rutstroemia bulgarioides E p. 409

Plectania nannfeldtii W p. 229

Chlorociboria aeruginascens A p. 367

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revivable; Lachnum calyculiforme on hardwood, not revivable)

31a. (30a) Hairs on underside colored yellow to brown (Lachnellula species on conifers, and

Lachnellula arida W p. 390

Lachnellula cf. flavovirens W p. 390

Brunnipila cf. calyculiformis W + ? p. 390

Velutarina rufo-olivacea A p. 390

T31b. Hairs on underside of cup hyaline to white, on conifers, revivable

Lachnellula calyciformis CA + ? p. 389

Lachnellula gallica BC + ? p. 389

K e y to I nclu de d A scom ycete s

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Lachnellula suecica N p. 389

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31c. Hairs on underside hyaline to white, on hardwoods, cones and cupules, sometimes on conifers, not revivable

Lachnum bicolor A p. 392

Lachnum virgineum A p. 393

Microstoma floccosum E p. 176

Lachnum virgineum A p. 393

32a. (28a) Fruitbody head spatula-shaped, or cup-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 32b. Fruitbody head globular to clubshaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33

Leotia lubrica A p. 394

Beug-final.indb 38

Lachnum pudibundum PNW + ? p. 393

33a. Head and stalk gelatinous

Leotia viscosa A p. 396

A scom ycete Fu ngi of North A m er ica

Dasyscyphella nivea PNW + ? p. 376

Coryne atrovirens A p. 394

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T33b. Head dark brown to black, globose, distinctly separated from the brownish stalk, in disturbed areas or in moss . . . . . . .Sarcoleotia globosa p. 419

Cudonia lutea E p. 373

Neolecta irregularis E p. 432

Hypocrea leucopus A p. 302

Neolecta vitellina A p. 432

Cudonia circinans A p. 373

Pachycudonia monticola W, M p. 403

33c. Not as above

34a. (32a) Fruitbody urn- to cup- to disc-shaped

Sowerbyella rhenana A p. 256

Neournula pouchetii W p. 196

Geopyxis carbonaria A p. 143

Tarzetta cupularis A p. 258

K e y to I nclu de d A scom ycete s

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Geopyxis vulcanalis W p. 143

Helvella leucomelaena N p. 165

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34b. Fruitbody a flattened to irregular club or spatula-shaped, colors various . . . . . . . . . . . .35

Neolecta irregularis E p. 432

Spathularia cf. rufa E p. 412

Spathulariopsis velutipes E p. 412

35a. Fruitbody brown to black . . . . . . . . . . . . . . . . 36 35b. Fruitbody tan to pale orange, bright yellow or greenish

Microglossum viride A p. 425

Spathularia flavida yellow form A p. 412

Microglossum rufum A p. 427

Spathularia flavida A p. 412

Microglossum fumosum N p. 427

Microglossum olivaceum A p. 425

36a. (35a) Head and stalk black to brown, stalk ± smooth, spore print black

Geoglossum alveolatum E p. 423

Geoglossum glabrum A p. 423

A scom ycete Fu ngi of North A m er ica

Beug-final.indb 40

Geoglossum peckianum E p. 420

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Geoglossum glutinosum A p. 420

Geoglossum umbratile A p. 424

Geoglossum simile E p. 422

Geoglossum simile asci/spores + paraphyses

T36b. Head and stalk black to brown or purplebrown, stalk ± smooth, spore print white

Geoglossum fallax A p. 423

Beug-final.indb 41

(G. fallax p. 423, white initially, then spores smoky at full maturity)

Thuemenidium arenarium E p. 428

K e y to I nclu de d A scom ycete s

Geoglossum difforme E p. 420

Thuemenidium atropurpureum A p. 428

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36c. Head and stalk black, hairy (use a hand lens), spore print brown

Trichoglossum farlowii E + ? p. 429

Trichoglossum velutipes A p. 429

37a. (9a) Not growing on another mushroom, shape various, sessile or short stalk . . . . . . . .39 37b. Growing on another mushroom or on wood or vegetation after decomposing the mushroom. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38

Hypomyces banningiae E p. 322

Hypomyces lateritius A p. 323

38a. Growing on a somewhat identifiable gilled mushroom

Hypomyces macrosporus E p. 322

Hypomyces lateritius A p. 323

A scom ycete Fu ngi of North A m er ica

Beug-final.indb 42

Trichoglossum walteri A p. 429

Hypomyces lactifluorum A p. 321

Hypomyces lactifluorum A p. 321

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Hypomyces porphyreus MI p. 326

Nectriopsis tubariicola A p. 335

Hypomyces luteovirens A p. 324

Hypomyces luteovirens A p. 324

Hypocrea avellanea E p. 327

Hypomyces hyalinus A p. 320

T38b. Growing on a more or less unidentifiable gilled mushroom

Hypomyces ochraceus A p. 325 (Cladobotryum verticillatum)

Hypomyces tremellicola E, MW p. 327

T38c. Growing on a bolete

Hypomyces chrysospermus A p. 319

Hypomyces microspermus A p. 319

K e y to I nclu de d A scom ycete s

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Hypomyces chlorinigenus E p. 318

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Hypomyces melanocarpus E p. 319

Hypomyces completus E p. 319

Hypomyces completus E p. 319

Hypocrea lixii A p. 308

Hypocrea pulvinata N p. 307

T38d. Growing on a polypore

Hypocrea americana N p. 316

Hypomyces aurantius A p. 315

Protocrea farinosa A p. 316

Protocrea pallida E p. 316

Sporophagomyces chrysostomus E p. 326

A scom ycete Fu ngi of North A m er ica

Beug-final.indb 44

Hypomyces rosellus A p. 316

Hypomyces rosellus A p. 316

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38e. Growing on a slime mold or a non-gilled mushroom other than a bolete or polypore

Hypocrea latizonata A p. 303

Hypocrea sulphurea A p. 312

Nectriopsis violacea A p. 336

Helminthosphaeria clavariarum NW, NE p. 301

Hypomyces cervinigenus A p. 317

K e y to I nclu de d A scom ycete s

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Sphaeronaemella helvellae A p. 150

Hypomyces leotiicola A p. 394

Hypomyces cervinigenus A p. 317

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39a. (37a) Not growing on the ground in a burned area or not growing on burned wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 39b. Growing on burned wood, or on ground in a burned area, or among mosses a few years after a burn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 40a. Fruitbody typically greater than 2 cm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

Tricharina gilva NW p. 259

Trichophaea abundans W p. 262

Anthracobia macrocystis W p. 128

41a. Fruitbody underside glabrous to slightly scurfy to tomentose but lacking long hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 41b. Fruitbody with hairs on the underside of the disk or cup

Sphaerosporella brunnea A p. 257

Trichophaea hemisphaerioides NW p. 261

Anthracobia melaloma W p. 128

A scom ycete Fu ngi of North A m er ica

Beug-final.indb 46

40b. Fruitbody typically between 0.5 cm and 2 cm in diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .41

Trichophaea contradicta NY, BC p. 262

Trichophaeopsis bicuspis E + ? p. 262

Anthracobia maurilabra E p. 127

46

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42a. (41a) Fruitbody deeply cup-shaped to goblet-shaped

Geopora arenosa N p. 89

Geopyxis carbonaria A p. 143

42b. Fruitbody cup-shaped to disk-like. . . . . . . . 43

Peziza griseorosea E p. 219

Rhodotarzetta rosea A p. 244

43a. Distinctly cup-like when young, not brightly colored

Peziza ostracoderma N p. 212

Plicaria carbonaria N p. 230

43b. Disk-like or if cup-shaped, brightly colored

Pulvinula laeterubra W p. 239

Pyronema omphalodes A p. 240

Pulvinula convexella W p. 239

Pyronema omphalodes A p. 240

K e y to I nclu de d A scom ycete s

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Pulvinula carbonaria N p. 238

Pyronema domesticum A p. 240

47

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Byssonectria fusispora A p. 132

Byssonectria fusispora A p. 132

Ascobolus carbonarius A p. 129

44a. (40a) Asci not amyloid

Gyromitra ancilis A p. 144

Rhizina undulata A p. 242

Plicaria endocarpoides W p. 231

44b. Asci amyloid (all in genus Peziza)

Peziza echinospora A p. 209

Peziza proteana f. proteana A p. 217

Peziza petersii N p. 213

Peziza praetervisa A p. 216

A scom ycete Fu ngi of North A m er ica

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Peziza proteana f. sparassoides A p. 218

Peziza sublilacina A p. 219

48

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45a. (39a) Fruitbody not growing on dung or decaying vegetation . . . . . . . . . . . . . . . . . . . . . 48 45b. Fruitbody growing on dung or decaying vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46

46a. Fruitbody less than 0.5 cm broad . . . . . . . . . 47 46b. Fruitbody greater than 0.5 cm broad

Peziza vesiculosa (2–8 cm broad) A p. 223

Peziza vesiculosa (2–8 cm broad) A p. 223

Peziza fimeti (0.5–2 cm broad) A p. 210

47a. (46a) Underside of cup or disk with distinct hairs

Pseudombrophila porcina NE p. 234

Pseudombrophila merdaria NE p. 234

Pseudombrophila cervaria N p. 233

Iodophanus carneus A p. 234

Cheilymenia theleboloides A p. 137

K e y to I nclu de d A scom ycete s

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Cheilymenia stercorea A p. 135

Pseudombrophila theioleuca A p. 233

Lasiobolus cuniculi A p. 130

Cheilymenia fimicola A p. 135

49

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47b. Underside of the cup or disk glabrous to slightly tomentose

Ascobolus furfuraceus A p. 130

Ascobolus stercorarius A p. 130

Byssonectria cartilaginea WM p. 132

Byssonectria terrestris A p. 132

48a. (45a) Fruitbody woody to fleshy but not gelatinous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 48b. Fruitbody gelatinous or spongy . . . . . . . . . . 49

Ascocoryne cylichnium A p. 357

Ascocoryne sarcoides A p. 357

A scom ycete Fu ngi of North A m er ica

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49a. Fruitbody typically less than 2 cm broad and fruitbody gelatinous

Bulgariella pulla E p. 364

50

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49b. Fruitbody typically greater than 2 cm broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

Scorias spongiosa (young) E p. 439

50a. Fruitbody gelatinous when young, soon spongy and up to 15 cm or more broad and 6 cm or more tall

Scorias spongiosa (old) E p. 439

T50b. Gelatinous, upper surface dark purple to black

Pseudosarcosoma latahense PNW p. 266

Camarops petersii E, MW p. 287

Urnula padeniana NW p. 265

Plectania melastoma A p. 227

K e y to I nclu de d A scom ycete s

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Plectania milleri W, M p. 228

Bulgaria inquinans A p. 364

51

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50c. Gelatinous, upper surface not dark purple to black

Galiella rufa E p. 142

Sarcosoma globosum NE p. 249

Neobulgaria pura E p. 401

Camarops petersii, young E, MW p. 287

51a. (48a) Fruitbody typically less than 2 cm broad or tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 51b. Fruitbody typically greater than 2 cm broad or tall . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .52

Pachyella adnata A p. 203

A scom ycete Fu ngi of North A m er ica

Ascotremella faginea A p. 359

52a. Growing on water-soaked wood

Pachyella clypeata E p. 203

52b. Growing on wood or soil, but if on wood, not characteristically on very water-soaked wood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53a

Beug-final.indb 52

Entonaema liquescens SE + IL, KS p. 300

Miladina lecithina N p. 203

53a. Fruitbody cup-shaped to rabbit-ear-shaped, at least when young (see also 54b) . . . . . . . . .59 53b. Fruitbody not cup-shaped to rabbit-earshaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

52

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54a. Fruitbody shield-shaped to brain-like, light tan, attached to wood by a central stalk,

Hypocrea peltata SE p. 305

underside resembling a parasitized gilled mushroom, fleshy to leathery

Hypocrea peltata SE p. 305

T54b. Fruitbody a stalkless shallow cup with a tough texture that dries corky

Wolfina aurantiopsis E + CO p. 269

54c. Fruitbody woody to carbonaceous . . . . . . . .55

Biscogniauxia repanda N p. 286

55a. Inner material composed of both host material and fungus (hand lens)

Biscogniauxia marginata E p. 286

K e y to I nclu de d A scom ycete s

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Wolfina aurantiopsis E + CO p. 269

Biscogniauxia mediterranea A p. 286

53

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Biscogniauxia atropunctata SE p. 285

Apiosporina morbosa A p. 438

55b. Inner material composed of fungal material only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .56

Diatrype stigma E p. 295

Kretzschmaria deusta A p. 332

56b. Fruitbody more or less globular, though adjacent fruitbodies may fuse . . . . . . . . . . . . .57

Daldinia concentrica N p. 293

Kretzschmaria deusta A p. 332

57a. Cutt ing material in half reveals light and dark bands

Daldinia grandis W p. 294

A scom ycete Fu ngi of North A m er ica

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56a. Fruitbody eff used, margin generally adhering to substrate

Daldinia childiae A p. 291

54

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57b. Cutt ing material in half does not reveal alternating light and dark bands . . . . . . . . . . . . . . .58

Annulohypoxylon thouarsianum A p. 282

58a. Perithecia always surrounded by carbonaceous material, ostioles always higher than stromatal surface, ostioles may be encircled with an annulate disk

Annulohypoxylon multiforme A (not South) p. 281

58b. Perithecia never surrounded by carbonaceous material, ostioles usually lower than or at same level as surrounding stromatal

Hypoxylon fuscum A p. 329

K e y to I nclu de d A scom ycete s

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surface, ostioles never encircled with an annulate disk

Hypoxylon fragiforme E p. 330

Hypoxylon ferrugineum E p. 331

Annulohypoxylon cohaerens E p. 281

Hypoxylon rubiginosum A p. 331

Thuemenella cubispora E p. 346

Hypoxylon howeanum A p. 330

55

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59a. (53a) Cup thin-fleshed, ribbed on underside or with a buried ribbed stalk . . . . . . . . . . . . 19b 59b. Not both thin-fleshed and ribbed on the underside or with a buried ribbed stalk. . . . . . 60

Gyromitra olympiana W p. 145

Gyromitra leucoxantha A p. 145

Gyromitra ancilis A p. 144

Gyromitra melaleucoides A p. 152

60b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . .61

Sarcoscypha austriaca NE + N central p. 245

Gyromitra mwb060911 W p. 145

Disciotis venosa complex A p. 141

61a. Fruitbody upper surface scarlet (or white in albino forms), on wood

Sarcoscypha dudleyi E, MW p. 247

A scom ycete Fu ngi of North A m er ica

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60a. Upper surface sharply to broadly wrinkled to almost smooth, cap margin usually turned down, though may turn up into a broad cup shape

Sarcoscypha coccinea WC p. 246

56

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T61b. Fruitbody upper surface bright orange to reddish orange (or white in albino forms), on soil

Caloscypha fulgens A p. 133

Caloscypha fulgens A p. 133

Pseudaleuria quinaultiana WA OR p. 232

Aleuria aurantia A p. 126

61c. Upper surface not scarlet or orange (or bright white), on wood or soil . . . . . . . . . . . . 62

Otidea alutacea W p. 198

62a. Fruitbody rabbit-ear-shaped or a truncate ear, sclerotium usually absent, if present go to Wynnea americana p. 270

Otidea cf. concinna W p. 198

K e y to I nclu de d A scom ycete s

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Caloscypha fulgens A p. 133

Otidea leporina A p. 200

57

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Otidea unicisa E p. 200

Otidea smithii W p. 202

Otidea rainierensis WA p. 202

Otidea cf. tuomikoskii N or M p. 200

Wynnella silvicola A p. 272

Wynnea americana E p. 270

Otidea onotica A p. 200

Midotis irregularis E + OR p. 178

Wynnea americana E p. 270

62b. Fruitbody a fairly symmetrical cup, margin often inrolled when young, may expand to almost plane in age

Peziza arvernensis W p. 206

Peziza phyllogena A p. 215

A scom ycete Fu ngi of North A m er ica

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Peziza badia A p. 207

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Peziza domiciliana A p. 208

Peziza varia A p. 221

Peziza arenaria E p. 205

Peziza michelii OR, QC p. 220

Geopora sepulta E p. 90

Jafnea fusicarpa NE p. 173

Humaria hemisphaerica A p. 172

Jafnea semitosta NE p. 173

K e y to I nclu de d A scom ycete s

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Peziza succosa N p. 220

Peziza saccardoana E p. 219

Humaria hemisphaerica A p. 172

Tarzetta bronca E p. 258

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63a. (51a) Fleshy, globular to cup-shaped or disk-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 T63b. Neither fleshy nor composed of pustules that break through the bark of small branches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64

Diatrypella favacea A p. 295

Encoelia furfuracea A p. 379

64a. (63b) Not growing on grass or grain seedheads . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .65

Claviceps purpurea A p. 288

63c. Composed of pustules that break through the bark of small branches

64b. Evident as a purple-brown sclerotium on the seed-head of various grasses and grains, especially rye

Claviceps purpurea A p. 288

65a. (64a) Appears as minute pointed hyphae; at maturity with minute globose conidia (hand lens); found on the ground, leaves or

Chromelosporium fulvum A p. 212

wood, often on burned or sterilized soil of greenhouses

Chromelosporium coerulescens A p. 212

A scom ycete Fu ngi of North A m er ica

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Dermea cerasi A p. 377

Chromelosporium coerulescens hyphae 25X A p. 212

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T65b. Appears as deformed plant material

Mature Taphrina robinsoniana E p. 442

Mature Taphrina occidentalis W p. 442

65c. Fruitbody hard, composed of ± isolated perithecia or appearing as “tar spots” on leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66

Lasiosphaeria ovina E + MW p. 333

Young Taphrina occidentalis W p. 442

66a. Fruitbody hard, composed of isolated or small groups of perithecia

Rosellinia subiculata A p. 345

Rosellinia quercina A p. 345

Lasiosphaeria lanuginosa A p. 333

Lasiosphaeria spermoides NE + ? p. 333

K e y to I nclu de d A scom ycete s

61

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66b. Fruitbody resembles tar spots on leaves

Rhytisma cf. americanum E p. 406

Coccomyces dentatus A p. 371

Rhytisma punctatum A p. 406

Rhytisma salicinum A p. 406

67a. (63a) Fruitbody on the ground or herbaceous material . . . . . . . . . . . . . . . . . . . . . . . . . . .73 67b. Fruitbody on wood or on conifer needles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

68a. Fruitbody an apothecium, cup-shaped to disk-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69 68b. Fruitbody a perithecium often produced in a stroma

Hydropisphaera peziza A p. 334

Nectria cinnabarina A p. 334

A scom ycete Fu ngi of North A m er ica

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62

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Neonectria coccinea v. faginata E p. 334

Neonectria fuckeliana A p. 334

young Hypocrea cf. chlorospora A p. 311

Hypocrea cf. chromosperma A p. 311

young Hypocrea cf. strictipilosa A p. 310

Hypocrea cf. minutispora A p. 314

Hypocrea schweinitzii A p. 308

Hypocrea viridescens A p. 313

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Hypocrea cf. sinuosa A p. 311

Hypocrea pachybasioides A p. 304

Hypocrea cf. delicatula A p. 316

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69a. (68a) Fruitbody underside smooth or with fi ne hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .71 69b. Fruitbody underside with “eyelash” hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

70a. “Eyelash” hairs long and distinctive

Scutellinia umbrorum A p. 253

Scutellinia blumenaviensis NE p. 251

Scutellinia scutellata A p. 253

Scutellinia scutellata A p. 253

Scutellinia setosa A p. 252

Scutellinia kerguelensis E p. 254

Scutellinia pilatii E p. 254

Scutellinia scutellata var. leucothecia E p. 253

Scutellinia erinaceus A p. 252

70b. “Eyelash” hairs shorter or at least not immediately obvious

Scutellinia crucipila NE, NW p. 251

Scutellinia pennsylvanica E p. 253

A scom ycete Fu ngi of North A m er ica

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Cheilymenia vitellina E p. 138

64

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71a. (69a) Underside more or less glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

Arachnopeziza aurelia E, MW p. 356

71b. Underside distinctly clothed in hairs

Arachnopeziza aurelia E, MW p. 356

Hyaloscypha britannica WC + EC p. 360

Arachnopeziza aurata NE p. 356

Belonidium sulphureum N p. 360

72a. (71a) Fruitbody thin with a translucent-waxy appearance

Orbilia delicatula N p. 435

Orbilia xanthostigma N p. 435

K e y to I nclu de d A scom ycete s

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Orbilia inflatula N p. 435

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Orbilia luteorubella N p. 435

Orbilia piloboloides A p. 435

T72b. Fruitbody brightly colored, not thin and translucent-waxy (see also 30b)

Chlorosplenium chlora E p. 368

Phaeohelotium epiphyllum A p. 404

Pithya vulgaris W, M p. 225

Pithya cupressina A p. 225

A scom ycete Fu ngi of North A m er ica

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Pseudopithyella minuscula CA p. 226

Pithya cupressina A p. 225

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72c. Fruitbody dull-colored, on wood or conifer cone scales

Mollisia cinerea A p. 399

Tapesia fusca A p. 400

Mollisia discolor BC + ? p. 400

Hyaloscypha albohyalina BC + ? p. 360

Chlorencoelia versiformis E, MW + BC p. 366

Ionomidotis fulvotingens E p. 388

Catinella olivacea A p. 365

K e y to I nclu de d A scom ycete s

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Mollisia ventosa E p. 400

Patellariopsis clavispora A p. 400

Godronia urceolus N p. 381

Ciboria rufofusca NW p. 369

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73a. (67a) Underside clothed with hairs or feltlike from matted hairs

Acervus epispartius NE p. 125

Neottiella rutilans A p. 195

Wilcoxina rehmii N p. 134

Chaetothiersia vernalis CA p. 134

Podophacidium xanthomelum N p. 405

Byssonectria terrestris A p. 132

Belonopsis graminea A p. 400

Byssonectria tetraspora W p. 197

A scom ycete Fu ngi of North A m er ica

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Neottiella vivida N p. 195

Sphaerosporella hinnulea E p. 257

Rhodoscypha ovilla W p. 243

Melastiza chateri N + M p. 175

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73b. Underside glabrous or at most minutely tomentose

Peziza gerardii E p. 211

Peziza azureoides E p. 211

Peziza griseorosea E p. 219

Pseudoplectania nigrella A p. 236

Discinella boudieri BC + ? p. 215

Smardaea planchonis WC + QC p. 236

Smardaea planchonis WC + QC p. 211

Octospora humosa NE p. 197

Lamprospora crechqueraultii A p. 174

Pulvinula archeri NW p. 239

K e y to I nclu de d A scom ycete s

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Pulvinula convexella W p. 239

Pulvinula laeterubra N p. 239

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Chapter 3 Hypogeous Ascomycetes The Truffles

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y pogeous Ascom ycetes h ave evolved from many different lineages. A great variety of hypogeous Ascomycetes in North America are known to us because of the work done by Helen Gilkey (Gilkey 1916, 1939, 1947, 1954, 1961) and later by James Trappe, as well as many others (see Trappe et al. 2009; Fogel and States 2001, 2002a, 2002b, 2003; Frank, Southworth, and Trappe 2006a, 2006b) These fungi are rarely seen unless one is specifically searching for them, raking beneath the soil surface using a truffle rake or other similar implement. The most widely known and prized hypogeous Ascomycetes from a culinary standpoint are in the Pezizales order, predominately in the Tuberaceae family. The Tuberaceae in North America are represented by the genera Tuber, which has 60 described species in north temperate forests and numerous undescribed species, and Choiromyces, which has three species found in north temperate forests. The Morchellaceae family contributes four North American genera:

H

Fischerula1 (represented solely by a Pacific Northwest species, Fischerula subcaulis, distinguished by its large spores with warty, ridged agglutinated-spiny spore ornamentation; Imaia (one Appalachian Mountain species, Imaia gigantea; Gábor et al. 2008; Kovács et al. 2008); Kalapuya (one Pacific Northwest species, Kalapuya brunnea); and Leucangium2 (one Pacific Northwest species, Leucangium carthusianum).

The genera Balsamia (five species reported and named from north temperate forests, but having many additional unnamed species) and Barssia (having two described species in north temperate forests and one undescribed species) represent the Helvellaceae family. The Discinaceae family has one hypogeous genus, Hydnotrya (with twelve species described from north temperate forests plus additional undescribed species). The Pezizaceae family includes the following six genera: Cazia (Cazia flexiascus from the Pacific Northwest and Cazia quercicola from southwest Utah); Hydnobolites (Hydnobolites californicus Fischer from California as well as twelve to eighteen unnamed species, per personal communication with Rosanne Healy); Hydnotryopsis (two hardwood associated species from Pacific coastal states, Hydnotryopsis setchellii Gilkey and Hydnotryopsis compactus (Gilkey) Trappe, with others yet to be described); Pachyphloeus (six named species from north temperate forests, plus eight unnamed welldocumented species and probably an additional ten or more species that are less well documented); Peziza (with at least three hypogeous species); and Terfezia (no known North American species). The Pyrenomycetaceae family includes seven known genera as follows: Genea (with 25 species named from north temperate forests, plus some yet to be described); Genabea (with five species named from north temperate forests);

1. The placement of Fischerula in the Morchellaceae is not accepted by all authors, as some consider its proper placement to be unclear. 2. Some authors place Leucangium in the Helvellaceae.

Overview

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Geopora (twelve species from north temperate forests); Gilkeya (one species found from northern Oregon to central Mexico; Smith, Trappe, and Rizzo 2006); Hydnocystis (one new species from Minnesota, per personal communication with Rosanne Healy); Otidea (one species, Otidea subterranea); and Stephensia (four described species from north temperate forests and one undescribed spe-

H y pogeous A scom ycetes: Th e Tru ffl es

Beug-final.indb 72

cies known from Oregon; Trappe, Bushnell, and Castellano 1997). The order Eurotiales has one hypogeous family, Elaphomycetaceae, with one hypogeous genus, Elaphomyces (having thirty described species to date, with many more waiting to be described [Zhang and Minter 1989]).

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8a. Smooth to verrucose clean surface with a belly-button-like cavity; mild taste; exterior delicate pinkish to pinkish brown or orange; gleba solid white except for narrow veins; always with relatively young Douglas fi r . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Barssia

Key to the Hypogeous Ascomycete Genera 1a. Interior more or less hollow or with chambers that are often maze-like . . . . . . . . . . . . . .17 1b. Interior fi lled with mushroom tissue (possibly hollow when very young) . . . . . . . . . . . . . .2

8b. Surface minutely papillate to distinctly warted, excavated in places . . . . . . . Balsamia

2a. Peridium thin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2b. Peridium thick . . . . . . . . . . . . . . . . . . . . . . . . . . . .3

9a. (7a) Peridium white to brown; gleba gray to brown, marbled with white veins lined with peridium-like tissue . . . . . . . . . . .Hydnobolites 9b. Peridium pink to pinkish brown or orange; gleba white to pale gray with empty canals opening into a central depression . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Barssia

3a. Gleba marbled, spores not powdery at maturity . . . . . . . . . . . . . . . . . . . . . Pachyphloeus 3b. Gleba not marbled, spores powdery at maturity . . . . . . . . . . . . . . . . . . . . . . Elaphomyces 4a. (2a) Gleba marbled and not developing gray to dark gray to greenish blue pockets of asci .......................................... 6 4b. Gleba white to buff with meandering whitish sterile veins, developing gray to dark gray to greenish blue pockets of spore-bearing asci . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5 5a. Exterior light yellowish brown to orangebrown to brown, known from the Pacific Northwest . . . . . . . . . . . . . . . . . . . . . . . Kalapuya T5b. Exterior light yellowish brown to orangebrown to brown, known from the Appalachian Mountains . . . . . . . . . . . . . . . . . . . . .Imaia 5c. Exterior dark brown to blackish (greenish to grayish at maturity) . . . . . . . . . . . Leucangium 6a. (4a) Gleba veined with meandering, often dendritic, veins . . . . . . . . . . . . . . . . . . . . . . . . . .10 6b. Gleba with broad, sparingly forked, relatively straight, translucent veins. . . . . . . . . . . .7 7a. Peridium smooth to scabrous . . . . . . . . . . . . . .9 T7b. Peridium smooth to verrucose . . . . . . . . . . . . .8 7c. Peridium verrucose, with warts obscured by smooth tomentum . . . . . . . . . . . . . . . . . . . . . . . . .Otidea subterranea

Truffle K ey

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10a. (6a) Spores hyaline to pale yellow-brown or pale brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 10b. Spores pale brown to dark yellow-brown or dark brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11a. Spores red-brown with Melzer’s reagent, grayish to reddish brown columella often present, often with a short stalk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fischerula 11b. No reaction to Melzer’s reagent or otherwise not as above. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12 12a. Spores hyaline to pale brown, pitted and resembling miniature golf balls or ornamented with flexuous rods that have a rimmed depression at the tip (resembling golf tees) and whose length varies greatly from spore to spore . . . . . . . . . . . . . . . . . . . . . . . . . Choiromyces 12b. Spores pale brown to dark brown and not ornamented as above . . . . . . . . . . . . . . . . . Tuber 13a. (10a) Gleba purplish gray with white veins, sections in 3% KOH instantly greenishyellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cazia 13b. Not as above (see also 12a) . . . . . . . . . . . . . . .14

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14a. Asci not reactive to Melzer’s reagent . . . . . .16 14b. Asci turn blue with Melzer’s reagent. . . . . . . 15 15a. Fruitbody clay color or silver-white, aging yellowish, minutely scabrous, lobed; interior composed of winding canals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnotriopsis 15b. Fruitbody pink, roughened, irregular, deeply furrowed; interior flesh pink marbled with brown veins . . . . . . . . . . . . . . . . . Peziza stuntzii 16a. (14a) Fruitbody pale yellowish brown to dark brown, scattered brown tomentum, much lobed and convoluted; interior white to cream with short pale pinkish brown to pale brownish orange veins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stephensia 16b. Fruitbody sordid white to peach colored or brown, glabrous to scabrous, lacking tomentum, looks and feels like gristle; interior with broad translucent bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Hydnobolites 17a. (1a) Outer and inner surfaces verrucose, may or may not be pubescent . . . . . . . . . . . . . . . . . 20 17b. Outer and inner surfaces smooth to pubescent to minutely hairy but not verrucose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18 18a. Asci turn blue with Melzer’s reagent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Peziza 18b. Asci not bluing in Melzer’s reagent . . . . . . . .19

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19a. Fruitbody pale to dark brown, fuzzy, ± verrucose . . . . . . . . . . . . . . . . . . . . . . . Geopora (see also Otidea subterranea) 19b. Fruitbody white to yellow, pinkish, or redbrown to brown, smooth or pubescent but not fuzzy, not verrucose . . . . . . . . . Hydnotrya 20a. (17a) Outer surface dark brown, dark reddish brown, reddish gray, or black. . . . . . . . . Genea 20b. Outer surface creamy white to grayish yellow, tan, brown, or vinaceous . . . . . . . . . . . . .21 21a. Outer surface pink to vinaceous, nearly spherical, convoluted, with apical opening to interior; interior white to pinkish, hollow, interrupted by irregular projections from the outer wall . . . . . . . . . . . . . . . . . . . . . . . . . . Gilkeya 21b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22a. Outer surface distinctly tomentose (velvety) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geopora 22b. Outer surface at most minutely tomentose, glabrous or with long external hairs . . . . . . .23 23a. Covered with abundant to scattered long brown hairs, apical orifice often wide enough to give overall cup-like appearance, irregularly lobed or coarsely knobby . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genea 23b. Not covered with external hairs, roundish to knobby, brain-like with several openings to interior . . . . . . . . . . . . . . . . . . . . . . . . . . . Genabea

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Balsamia magnata Harkness fruitbody globular and hypogeous with no stalk; 0.5–2.0 cm broad; spherical to flattened, usually with an infolded top, covered with round to pointed warts, sometimes also with some small depressions; exterior whitish when very young but soon bright orange to reddish brown or brownish pink; interior with maze-like translucent canals lined by hymenium and usually fi lled with cottony hyphae; often with a persistent tuft of mycelium at base; odor and taste not distinctive. MICROSCOPIC FEATURES: spores 20–24 × 12–14 μm, elliptical, smooth, with irregular arrangement, hyaline, usually with 3 oil drops; asci 8-spored, MACROSCOPIC FEATURES:

irregular in shape, stipitate; paraphyses mainly a palisade. OCCURRENCE: solitary to numerous under many different trees and shrubs, including oaks, pines, fi rs, and Douglas fi r; year-round; common in the Pacific Northwest from Oregon to Southern California and east to Arizona and Colorado. EDIBILIT Y: reportedly fl avorless, but nice texture when added raw to various dishes. COMMENTS: Pseudobalsamia magnata (Harkness) Gilkey is a synonym. Balsamia alba Harkness (not illustrated) is sordid white with coarse warts, while Balsamia vulgaris Vittadini (not illustrated) has minute warts and an odor described as

Balsamia magnata

Balsamia

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similar to mouse feces or rancid buttered popcorn. Members of the Pachyphloeus carneus group (p. 105) can be bright orange like Balsamia magnata, but Balsamia magnata has a pale interior and smooth spores, while Pachyphloeus carneus group members have a yellow to green interior and spiny spores. Typically, Balsamia spores are difficult to distinguish from Barssia and Geopora spores. Barssia oregonensis Gilkey, a Pacific Northwest spe-

cies found under Douglas fi r, is similar but has larger spores (24–36 × 12–21 μm). Barssia oregonensis is lumpy, typically with a depression at the top, 1–2.5 cm broad, exterior fi nely warted or smooth and lumpy, and ochre-buff to orange-cinnamon to brick-red. The white to pale gray interior has canals lined by felt-like hymenium that mostly empty into the central depression.

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Balsamia nigrans Harkness fruitbody globular, hypogeous, lacking a stalk; 0.5–2 cm broad; spherical with a central depression; exterior coarsely and sharply warted, black; interior marbled, with irregular canals or sometimes chambers, whitish; with a more or less distinct mycelial tuft at the base; odor and taste not distinctive. MICROSCOPIC FEATURES: spores 20–37 × 12–15 μm, oblong, thin-walled with faint hyaline meandering lines around the circumference, hyaline, with one or more oil droplets; asci 40– 60 × 30–40 μm, more or less saccate, stipitate. OCCURRENCE: infrequent in dry areas under pines, oaks, and Douglas fi r at low to mid elevation; yearround; central California to northwest Oregon. MACROSCOPIC FEATURES:

reportedly has a nice texture but litt le flavor, used raw. COMMENTS: Pseudobalsamia nigrans (Harkness) Gilkey is a synonym. Th is species is listed in some sources as Balsamia nigrens Harkness. There are five or six species of Balsamia reported from north temperate forests. All have a basal or lateral cavity, a verrucose peridium, and white to pale yellow marbled flesh. There is usually a basal tuft of mycelium. See further comments about Balsamia under Balsamia magnata (p. 75). EDIBILIT Y:

Balsamia nigrans

Balsamia

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Choiromyces meandriformis Vittadini MACROSCOPIC FEATURES: fruitbody globular, irreg-

ular, hypogeous, lacking a stalk; up to 12 cm in diameter; exterior irregularly folded, lobed, smooth to slightly fibrillose in the folds, pale to pinkish buff to yellowish brown; peridium thin; interior solid, soft , drying hard, white to yellowish with wavy labyrinthine grayish to brown veins; odor faint when young, strong and aromatic when mature. MICROSCOPIC FEATURES: spores 22–30 μm including the ornamentation of spines and rods 3– 6 μm long, globose, biseriate or irregularly arranged, pale brown when mature; asci 100–200 × 50– 60 μm, saccate with a stalk up to 120 μm long, 8-spored. OCCURRENCE: occasional under Douglas fi r, pines, and hemlock at low elevation; fall; western Oregon, central California, and West Virginia, uncommon in North America, common in Europe.

Pig Tru ffle

considered a delicacy in northern Europe and poisonous in Spain, this species should be approached with caution. COMMENTS: Choiromyces venosus (Fries) Th . Fries, Tuber album Sowerby and Rhizopogon meandriformis Vittadini are synonyms. The spores have distinctive golf-tee-like rods. The fruitbodies can be quite large. Two of the three species described from northern temperate forests have been found in North America. Choiromyces alveolatus (Harkness) Trappe has distinctive pitted spores that resemble micro golf balls and a yellow-orange gleba with pale veins. Choiromyces alveolatus of the Pacific Northwest starts out white, ages yellow or brown, is slightly lobed with a scabrous, sometimes pubescent surface, and reaches 1.3 cm in diameter with globose yellow or brown spores 22–36 μm in diameter. EDIBILIT Y:

Choiromyces meandriformis

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Elaphomyces americanum Castellano (in preparation) fruitbody hypogeous, lacking a stalk or basal tuft of mycelium, up to 16 mm in diameter, more or less globose; outer surface covered in distinct tall yellow-brown warts, most acutely pointed, surface partly to completely embedded in a pale mycelial mat that obscures the warts; peridium two-layered, outer warty layer yellow-brown, inner layer marbled with white to off-white veins, matrix pale tan near the surface grading to tan to brown and then dark brown near the gleba; gleba spore mass powdery, dark brown with tan spiderweb-like hyphae; odor and taste not recorded. MICROSCOPIC FEATURES: spores when mature 28– 34 μm wide (mean 30.9 μm), globose, ornamented with nonreticulate tuft s of coalesced rods 1–2 μm tall, brown in KOH, with small (aborted?) globose spores roughly 16 μm in diameter; asci 42–46 μm in diameter, walls 3–4 μm thick, globose, hyaline. OCCURRENCE: in sand layer under fi r, pine, and hemlock; summer to early winter; eastern Canada south to Tennessee, west to Michigan; common. EDIBILIT Y: considered inedible. COMMENTS: What has been known as Elaphomyces muricatus in North America represents a species complex of several undescribed species and is distinct from Elaphomyces muricatus Fries of Europe (Castellano and Trappe, unpublished data). Elaphomyces americanum Castellano is an eastern member of this complex. Elaphomyces reticulatus

Vittadini, a European species found in deciduous forests but with a similar North American sister species, has an obscurely marbled peridium, flat warts on the outer cortex (in contrast to the more pointed pyramidal warts of the Elaphomyces muricatus complex), and on average smaller spores (19– 28 μm). Scleroderma species, which are Basidiomycetes, have a distinct base or att achment point.

El a phom yces

79

MACROSCOPIC FEATURES:

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Elaphomyces americanum

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Elaphomyces fallax Castellano and Trappe (in preparation) MACROSCOPIC FEATURES:

fruitbody hypogeous with a persistent circular basal tuft, up to 22 × 27 mm in diameter, subglobose, flattened to somewhat turbinate; outer layer mott led dark brown to black with distinct large, flattened, stellate warts, larger at apex, surface completely enveloped in pale brown to dark brown mycelium; peridium 1.5– 2 mm thick with a 200–250 μm britt le outer surface; subcutis 1–3.5 mm thick, uneven in thickness, zonate to somewhat mott led, off-white to pale tan exterior, brown interior, leathery; persistent basal tuft 19 mm in diameter × 6–7 mm long; gleba spore mass powdery, dark gray-brown with gray-brown spiderweb-like hyphae; odor slightly sweet, taste not recorded. MICROSCOPIC FEATURES: spores when mature 27– 31 μm (mean 29.3 μm), globose, walls about 1 μm thick, ornamentation a complete reticulum of alveoli up to 5 μm in diameter and 2–3 μm deep with coarse projections at the top; yellow-brown in mass; asci not seen. OCCURRENCE: hypogeous under oaks, hickories, other hardwoods, and pines; spring through fall; known from Florida, Maryland, North Carolina, Virginia, and West Virginia; common. EDIBILIT Y: not considered edible. COMMENTS: Elaphomyces fallax Castellano is similar to Elaphomyces cyanosporus Tulasne and C. Tulasne of Europe and has probably been misidentified as Elaphomyces cyanosporus in the past. There are a number of Elaphomyces species with a dark brown to black outer peridium. North American material similar to Elaphomyces anthracinus Vittadini of Europe has been found in Oregon, Idaho, and Quebec, though it is rare. The North American Elaphomyces anthracinus–like species has an outer surface that is mostly smooth with up to 0.1 mm broad warts and is coated with a thin, sparse layer of reddish-brown mycelium; the peridium is two-

layered with a thin black outer layer and a thick white inner peridium; the interior is initially hollow, gleba white and cottony at fi rst, turning powdery and black at maturity; spores are 12–21 μm, globose, densely covered with spines (contained in a thin epispore layer), and brownish black; and asci are not seen. Elaphomyces maculatus Vittadini and Elaphomyces leveillei Tulasne and C. Tulasne also have a black cortex that can be seen by cutt ing the fungus in half. These are both European species with sister species in North America. The American species have long been known by the European names. Elaphomyces cf. maculatus is found in deciduous forests, often associated with oaks, and has opaque spores 29–43 μm with densely clustered and coalescent spines 1–1.5 μm tall. Elaphomyces cf. leveillei is found in both deciduous and conifer forests, usually in black, peaty, acidic soil. Elaphomyces leveillei has 0.2–1 mm broad warts on the cortex resembling the surface of a dog’s nose, is usually enveloped in sulfur-yellow to pale greenish yellow mycelium, and has spores 18–26 μ in diameter. Elaphomyces macrosporus Castellano and Elliott (in preparation) is a rare species that also has a thin dark brown to black outer peridium with a thick white inner peridium and is distinguished by spores 45+ μm in size. Elaphomyces cf. loebae Castellano (in preparation), found in the southeast, has a unique peridium of small papillae that are contiguous and cover the entire surface (Michael Castellano, personal communication). The material illustrated is immature and still hollow. In mature specimens of Elaphomyces loebae, the spores measure 21–22(–23) μm including ornamentation that is 1–2 μm tall. Elaphomyces decipiens Vittadini is another European species with a similar American species found in the Northeast, Oregon, and northern California. Elaphomyces cf. decipiens, associated with conifers, has a pea-sized to

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Elaphomyces fallax

hazelnut-sized fruitbody that can be pale brown or orange, then black. The surface is nearly smooth to minutely warty, the inner peridium in cross section is white marbled with brown, and the globose spores are 18–28 μm in diameter. Elaphomyces appalachiensis Linder (not illustrated) is an uncommon but very distinctive litt le Elaphomyces species found in Great Smokies National Park (Linder

1939). It is typically 0.6 × 0.5 cm, exterior reddish purple, peridium differentiated into three layers: outer layer felty and purplish, inner peridium layer (bulk of peridium) texture punky and light pinkish cinnamon, and gleba white at the very center surrounded by distinctly greenish-blue-tinged gleba. The spores are (6.5–)7.5 × 9 μm, globose, subreticulate to reticulate-echinulate, hyaline.

El a phom yces

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Elaphomyces verruculosus Castellano (in preparation) MACROSCOPIC FEATURES:

fruitbody hypogeous, lacking a stalk or basal tuft of mycelium, irregularly subglobose to kidney-shaped, up to 22 mm thick × 27 mm long with an outer peridial surface of low, semirounded to angular or elongate warts; peridium pale yellow-brown when young, then mott led brown and yellow-brown when mature, covered with pale brown, brown, or yellow-brown hyphae and soil debris obscuring the warts and giving the surface a smooth appearance; peridium in cross section a roughly 140 μm exterior layer of red-brown to dark red-brown cells underlain with paler yellow-brown to off-white tissue 300–350 μm thick, immediately underlain with an off-white to pale gray-blue-tinged layer that is the basis for the warts, and fi nally a distinct roughly 100 μm thick brown layer at the interface with the gleba; gleba off-white and cottony when immature, spore mass powdery, dark brown to nearly black when mature with numerous gray to brown mycelial strands arising from the inner peridium wall and traversing the gleba; odor mild, taste not recorded. MICROSCOPIC FEATURES: spores when mature 35– 46 μm broad (mean 41.1 μm), globose, densely covered with rods and tuft s of spines 2–3 μm broad × 2–3 μm tall, brown to red-brown to dark red-brown in KOH; asci 42–50 μm broad, about 2 μm thick, globose, hyaline, 4–8-spored. OCCURRENCE: hypogeous under pines and Tsuga canadensis; late spring, summer, and fall; Quebec to Florida and northeastern Mexico; very common. EDIBILIT Y: considered inedible. COMMENTS: Elaphomyces verruculosus Castellano is aligned with Elaphomyces granulatus Fries of Europe, the deer truffle. Species similar to Elaphomyces granulatus are very abundant and widespread,

comprising a complex of species distinct from Elaphomyces granulatus found in Europe (Castellano and Trappe, unpublished data). They are often found by spott ing the Elaphocordyceps species that parasitize them. Elaphomyces asperulus (Vittadini) O. Kuntze is another European species with a sister species in North America. It has a pale brown outer peridium, a reddish gray to red-violaceous inner peridium, and mature spores are 25–38 μm in diameter with scarcely discernible coalescent spines 1 μm high, forming a cracking epispore. If the inner peridium is marbled, see Elaphomyces americanum (p. 79). If the outer surface is white to grayish and is smooth to somewhat verrucose with abundant pale yellow hyphae and hyphal tuft s emerging from the surface and encrusted with soil, consider Elaphomyces subviscidus (Zeller) Trappe and Guzmán. Elaphomyces subviscidus is found in the summer in Oregon, Idaho, and Colorado, is associated with lodgepole pine, ponderosa pine, Douglas fi r, and mountain hemlock, and can be up to 5 cm × 3 cm. The peridium is 0.25– 0.3 cm thick, with a thin, yellowish outer layer and a thick white to light gray inner layer. The interior spore mass is dark brown before maturity, nearly black when mature, and has an odor of onions. Elaphomyces subviscidus spores are globose, 12–29 μm in diameter including ornamentation of crowded spines 1–2 × 0.2 μm, and separated by 0.2– 0.5 μm or often joined in twos and threes by ridges but never forming a partial reticulum. Its spines are embedded in a gelatinous matrix and often separate from the spore surface when the cover slip is tapped. Its spores contain 1 large, round oil droplet and are of two sizes. The smaller spores have walls up to 0.5 μm thick and are very dark brown. The larger spores are thin-walled and tend to be lighter brown.

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Elaphomyces verruculosus

El a phom yces

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Genabea cerebriformis (Harkness) Trappe fruitbody globular, hypogeous, lacking a stalk; 0.5–2 cm broad, but usually less than 1 cm; exterior becoming irregularly lobed to brain-like with variably conic warts as it matures, with several openings to the interior, white to yellow-gray; interior a single cavity in small specimens, usually maze-like from infolding of the walls in larger specimens, less warted than the outer surface, color similar to exterior; odor mild to strong, pleasant, faintly of garlic. MICROSCOPIC FEATURES: spores 28–44 μm, globose, smooth when immature then covered with densely crowded spines 2–3 μm tall; spines often coalescing, giving the spore surface a rugose appearance, uniseriate to partially biseriate, hyaline when young, maturing grayish yellow; asci clavate to elliptic or irregular, 8-spored, though often not all spores mature; paraphyses clustered among densely crowded asci. MACROSCOPIC FEATURES:

solitary to gregarious in mineral soil or humus of coniferous or deciduous woods, often with Douglas fi r; western Washington to Mexico, common in the Pacific Northwest in the spring, found in the summer in Mexico. EDIBILIT Y: untested, too small to bother with. COMMENTS: Genea cerebriformis (Harkness) Gilkey, Myrmecocystis cerebriformis Harkness, and Myrmecocystis candida Harkness are synonyms. Gilkeya compacta (p. 93) also has round to nearly round spores but is distinguished by a reddish to vinaceous fruitbody, broad warts on the spores and cylindric asci. Genea species have brown to black fruitbodies and a basal tuft of mycelium, and while some spores in a collection may be nearly round, there will also be elliptic spores present. OCCURRENCE:

Genabea cerebriformis

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Genea arenaria Harkness fruitbody a convoluted hollow flattened sphere, 1–3 cm broad; exterior covered with scattered hairy warts, light brown to brown, with an apical orifice that opens into the interior; interior warted, paler than the exterior; tuft of hyphae at the base; odor mild to metallic-garlicky. MICROSCOPIC FEATURES: spores 20–28 × 15–23 μm, elliptic, exterior spore ornamentation of cones with obtuse, truncate, anvil-topped or forked tips, cones dissolve in KOH, strongly staining in cotton blue, 1 oil drop, hyaline. OCCURRENCE: associated with oaks in coastal and montane forests, often in sandy soils; western Oregon and California, also reported in Mexico and in northeastern North America; not uncommon regionally. EDIBILIT Y: reportedly palatable. MACROSCOPIC FEATURES:

Genea compacta Harkness (not illustrated) is also brown but is distinguished by larger spores (32–34 × 24–28 μm). The following eastern species are all very rare. Genea echinospora Gilkey (not illustrated) is blackish brown, 0.7 cm wide, hispid in the furrows, and scabrous on lobes; its spores are 36–40 × 24– 28 μm, elliptic and echinate with mostly truncate spines. Genea macrosiphon Gilkey (not illustrated) is buff y-citrine, 0.6 cm in diameter. Its inner and outer surfaces are minutely warted. Its spores measure 28–32 × 24 μm and are covered with scattered, mostly rounded-conical large and small warts. Genea thaxteri Gilkey (not illustrated) is verrucose, yellowish brown, and glabrous except for a basal tuft of mycelium, and its spores measure 27.5– 32.5(–35) × 22.5–27.5 μm including ornamentation of uncrowded, somewhat pyramidal warts.

COMMENTS:

Genea arenaria

Genea

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Genea harknessii Gilkey MACROSCOPIC FEATURES:

fruitbody globular to flattened, somewhat lobed, usually with an opening to the interior, hypogeous, lacking a stalk, 0.5– 2.5 cm broad; outside surface covered with stout pyramidal projections, dark reddish brown to dark gray or black; base usually with a thick tuft of brownish hyphae; interior convoluted and hollow with irregular projections of sterile tissue from the outer wall, warty, concolorous with outer surface; flesh britt le, white to gray; odor fragrant, garlicky. MICROSCOPIC FEATURES: spores 24–28 × 22– 27 μm, broadly elliptic to globose, smooth when young, developing 1–3 μm cones at maturity, uniseriate or partially biseriate, hyaline; asci average 225 × 32 μm, cylindrical to somewhat club-shaped, stipitate; paraphyses 2–8 μm, branched below secondary cortex. OCCURRENCE: usually solitary, sometimes gregarious in humus or soil under Douglas fi r and oaks plus other trees and shrubs; spring through fall; lowlands of western Washington and western Oregon and both lowlands and high country of California; common. EDIBILIT Y: edible but small and rarely abundant. COMMENTS: There are three somewhat similar black species known from western North America. Genea gardneri Gilkey (not illustrated) tends to be

darker colored, more convoluted, rarely has projections of sterile tissue into the inner cavity, and has larger spores (32-36 × 28-34 μm) with low rounded warts that dissolve in KOH. Genea bihymeniata M. E. Smith and Trappe (not illustrated) has slightly shorter spores and notably rounded spore ornaments compared to Genea harknessii, which has irregularly shaped spore ornaments with pointed, forked, or anvil-shaped tips. Genea anthracina Heblack and Stewart (not illustrated), known only from Minnesota, also is coal black and also has smaller spores (excluding ornamentation 22– 26 × 18–20 μm), and the warts are hemispherical, never angular or pointed—features very similar to those described for Genea bihymeniata. Genea balsleyi M. E. Smith (not illustrated), found in New Jersey, has an outer surface lacking obvious surface hairs, low rounded to subangular warts, a gray interior and spores 24–28(–30) × (20–)21–25 μm excluding ornamentation that has crowded conical to irregular knobs with truncate to angular tops (Smith 2007). Genea brachytheca Gilkey (not illustrated), an eastern species, is bone-brown to nearly black, minutely and sharply verrucose, somewhat densely hispid, with spores averaging 28 × 24 μm, covered with generally pointed or truncate spines of various sizes.

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Genea harknessii

Genea

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Genea hispidula Berkeley and Broome ex Tulasne and C. Tulasne fruitbody globular, hypogeous, attached by a tuft of mycelium, not a stalk, 0.3–1.2 cm broad; outer surface very densely covered with long brown hairs, the surface is reddish brown; distinct apical opening leads to the interior, which is hollow (single cavity) and more or less convoluted. MICROSCOPIC FEATURES: spores 30–42 × 25–35 μm including the isolated rounded warts, broadly elliptic, uniseriate, hyaline; asci cylindric, 8-spored, forming a distinct hymenium on the inside; paraphyses forming an epithecium above the asci. OCCURRENCE: solitary to several on the soil surface or shallowly buried in the soil of hardwood forests, rarely under conifers; widespread in eastern North America and Europe; rare. MACROSCOPIC FEATURES:

unknown. COMMENTS: Genea hispidula is a gorgeous and easily distinguished truffle. We know of nothing else with such a beautiful and thick coating of long brown hairs over the entire outer surface. Genea arenaria (p. 85) has a whitish outer layer and has long brown hairs from warts but the hairs are much more scattered. Genea brachytheca Gilkey (not illustrated), known from Quebec, has an appearance that suggests Genea hispidula but the surface hairs tend to be fewer, shorter, stiffer, and straighter. The outer surface color is bone-brown or nearly black, the spores are smaller (24–32 × 20–28 μm), and the asci are short compared to the proportionately long paraphyses. EDIBILIT Y:

Genea hispidula

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Geopora arenosa (Fuckel) S. Ahmad fruitbody spherical, slightly embedded in the soil, becoming cupshaped to saucer-shaped with the margin entire or splitt ing, lacking a stalk, less than 2 cm broad; interior of cup whitish to ochre, smooth; exterior brown with short sinuous to tufted hair-like hyphae and adhering sand grains; flesh fragile, relatively thick. MICROSCOPIC FEATURES: spores 20–25 × 11–18 μm, elliptic with slightly narrowed ends, usually with 2 oil droplets, mostly uniseriate, hyaline; asci 200– 240 × 15–20 μm, nearly cylindrical, inamyloid; paraphyses up to 200 μm long, gradually enlarged to 7– 9 μm with few septa; hairs 80–200 × 6–10 μm, thick-walled with thin septa, smooth, branched, brownish. MACROSCOPIC FEATURES:

solitary to gregarious on damp sandy soil and around burns; spring and fall; New York to Oregon; uncommon. EDIBILIT Y: not reported, too small to be of interest. COMMENTS: Sepultaria arenosa (Fuckel) Boudier is a synonym. Geopora arenicola (Léveillé) Kers, known from Colorado to eastern Canada, is macroscopically identical but has spores 23– 28 × 14–16 μm. Geopora pellita (Cooke and Peck) T. Schumacher, known from Norway and Quebec, differs from Geopora arenicola only in the orange interior surface of the cup. Geopora clausa (Tulasne and C. Tulasne) Burdsall (not illustrated) is found in the riparian zones of semiarid regions in the west, often in association with Salicaceae. Geopora clausa is also quite similar to Geopora OCCURRENCE:

Geopora arenosa

Geopor a

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arenosa. Geopora clausa is widespread in Europe and is also found in the Canary Islands, occurring in two forms, one with broadly elliptic spores and the other with narrowly elliptic spores. Geopora sepulta Korf and Burdsall, a widespread eastern species, is 2–7 cm wide, nearly round and enclosed at fi rst, and opening and becoming deeply cup-shaped as it matures. Its outer surface is smoky brown to dark brown, densely covered with dark brown flexuous hairs that are often encrusted with sand or dirt, and its inner surface is smooth, dull whitish, and becomes yellowish or brownish with

age. The margin is incurved and entire or splitting irregularly, sometimes reflexed with a somewhat star-shaped outline. Its spores measure 18– 22 × 10–12 μm, are broadly elliptic and smooth, and typically contain 1 large oil drop. Lachnea sepulta (Fries) Cooke, Peziza sepulta Fries, and Sepultaria sepulta Massee are synonyms. For species that are similar but are not partially or entirely buried in the ground, see Humaria hemisphaerica (p. 172) and Trichophaea hemisphaerioides (p. 261) if the spores have oil drops. See Tricharina gilva (p. 259) if the spores lack oil drops.

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Geopora cooperi Harkness

Fuzzy Tru ffle

fruitbody globular, hypogeous, lacking a stalk, 1–10 cm broad; exterior deeply convoluted, velvety, variably verrucose, pale to dark brown; interior very convoluted, the folds often touching each other but leaving some open spaces, white streaked with brown; odor mild but can be faintly aromatic, radish-like, or garlicky. MICROSCOPIC FEATURES: spores 18–30 × 10–21 μm, ovoid to elliptic, 1 oil drop, smooth, mostly uniseriate, hyaline; asci 140–200 × 16–28 μm, 8-spored, cylindric to clavate, sometimes pointed at the apex, forming a distinct palisade. OCCURRENCE: solitary to gregarious under hardwoods and especially conifers; spring through fall; western North America from northern Alaska to Mexico, also found in northern Europe; at times abundant. MACROSCOPIC FEATURES:

edible and good. Geopora harknessii E. Fischer ex Gilkey, Geopora magnifica Gilkey, and Geopora annulata Gilkey are synonyms. There are twelve Geopora species reported from north temperate forests, and all of the others more or less resemble tiny hollow balls that open up at the apex; see Geopora arenosa (p. 89). Geopora cooperi is the most common Geopora (Trappe et al. 2009) and looks very different from the other members of the genus. Geopora cooperi most closely resembles a Hydnotrya, but Hydnotrya species have a smooth to only minutely scurfy outer surface and have distinctive spiny to warted spores that are yellow to brown at maturity. Genabea cerebriformis (p. 84) is always small (rarely more than 1 cm broad) and has globose spiny spores. Otidea subterranea R. A. Healy EDIBILIT Y:

COMMENTS:

Geopora cooperi

Geopor a

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and M. E. Smith (not illustrated) is a fuzzy truffle known from mixed oak woodlands in Iowa (Smith and Healy 2009). It is 1–2 cm broad, globose to subglobose, with a basal mycelial tuft . The fruitbody is whitish when young, aging peach-cream with tanbrown areas. It has a slight, pleasant odor of fresh coconut. The gleba is solid to partly solid with dark brown fertile veins lined with a yellowish-brown

hypothecium and sterile veins of cottony whitish hyphae. The hyaline spores are elliptical, 14– 17 × 9.5–12 μm, and mostly unigutt ulate. The asci are 8-spored, inamyloid, with crozier at base, and noticeably shorter than the paraphyses. Paraphyses are 150–175 × 2–3 μm, with tips fused to form an epithecium of dark brown, thick-walled cells.

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Gilkeya compacta (Harkness) M. E. Smith and Trappe fruitbody globular, hypogeous, lacking a stalk, 0.5–2.5 cm across; exterior maturing lobed or knobby with a small hairless apical opening, pink to reddish brown; interior hollow with lobes and projections of sterile tissue from the outer wall, less verrucose than outer wall, pink to whitish; flesh fragile, britt le, white; odor and taste mild. MICROSCOPIC FEATURES: spores 28–43 × 25–40 μm excluding ornamentation of rounded granules 2– 3(–7) μm tall and 2– 6(–15) μm wide, nearly globose, uniseriate, hyaline; asci 250–280 × 31–45 μm, typically 8-spored, cylindric, walls up to 2 μm thick when young but thinning to about 0.5 μm by maturity, arranged in a palisade; paraphyses crowded, enlarging and coalescing beyond the asci to form a secondary cortex. OCCURRENCE: solitary to gregarious in leaf mold and mineral soil under conifers and oaks; winter through summer but mainly spring to early MACROSCOPIC FEATURES:

summer; Pacific Northwest to southern Mexico; uncommon. EDIBILIT Y: too small for culinary use, insipid. COMMENTS: Genea intermedia Gilkey and Hydnocystis compacta Harkness are synonyms. Genea compacta Harkness (not illustrated) is a different species that is light brown, has a superficial layer of short 1-celled hairs, and the spores are elliptic (24–28 × 32–34 μm). The distinctive features of Genea compacta are the pink to vinaceous purple coloration, absence of hairs on the peridium, lack of a tuft of mycelium at the base of the stalk, and spherical or very nearly spherical spores. Genea species are tan to brown to black, have a tuft of mycelium at the base, and do not have round spores. Genabea cerebriformis (p. 84) has more or less spherical spores, but the spores are densely spiny and the fruitbody is dull grayish yellow, very convoluted, and brain-like.

Gilkeya compacta

Gilk eya

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Hydnobolites californicus Eduard Fischer MACROSCOPIC FEATURES:

fruitbody 0.5–3.0 cm broad, globular, hypogeous, slightly to very irregularly folded, the canals opening at the furrows; surface with occasional short hairs, at fi rst yellowish tan, becoming brownish when mature; interior canals narrow, mostly long, winding, gray to yellowish brown marbled with white veins; consistency gristly. MICROSCOPIC FEATURES: spores 12–24(–30) μm, globose, very coarsely alveolate with 3–4 alveoli across the diameter of the spore, the angles projecting as coarse blunt spines 4 μm long, spores yellowish when mature; asci scattered irregularly through the hyphal structure, irregularly spherical-ellipsoid, faintly amyloid. OCCURRENCE: mycorrhizal with conifers and oaks; summer and fall; known from the Pacific Northwest, California, Indiana, Maine, Tennessee, and Europe; infrequently encountered. EDIBILIT Y: unknown. COMMENTS: Cazia fl exiascus Trappe (not illustrated), found in the Pacific Northwest under oaks and another member of the Pezizaceae, like Hydnobolites, has asci that are at most only faintly amy-

loid. It has hyaline to pale, globose to elliptical, reticulate spores borne in cylindric but crooked asci, and the spores measure 11–16 × 10–12 μm excluding the 1–1.5 μm thick warty-reticulate ornamentation. Cazia flexiascus has a white glabrous peridium, darkening when exposed, a solid pale purplish gray gleba marbled with white veins radiating from a basal pad, and a drop of KOH produces a strong yellow reaction. Cazia quercicola Fogel and States (not illustrated), known from southern Utah under oaks, is brain-like, white, bruising brownish orange, subtomentose, gleba fi rm, pale reddish purple when immature, and white when mature. Cazia quercicola is differentiated from Cazia flexiascus by smaller, globose to subglobose, warty, reticulate spores (11–14 × 10–12 μm, warts less than 0.5 μm high) and no reaction of the gleba to KOH. The other genera with globose, reticulate spores—Elaphomyces, Scleroderma (a basidiomycete genus), and Tuber—all have dark spores. A unique feature of Hydnobolites is the gleba marbled with veins that are lined with inflated cells similar to those of the peridium; the veins seem to be narrow insertions of the peridium into the gleba.

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Hydnobolites californicus

H y dnobolites

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Hydnotrya cubispora (E. A. Bessey and B. E. Thompson) Gilkey MACROSCOPIC FEATURES:

fruitbody spherical to irregularly lobed, typically flattened, lacking a stalk, 0.5–3.5 cm in diameter; outer surface minutely hairy to bald, pallid gray-brown to light brown or orange-brown, drying red-brown; interior highly convoluted, chambered with appressed folds (rarely a single large chamber), chambers white to pale gray even when mature; hymenium feltypubescent; tissue presumably britt le; odor and taste not distinctive. MICROSCOPIC FEATURES: spores 30– 60 × 25–36 μm including the thick epispore, broadly elliptic to subglobose at maturity but often appearing cubical while still in the asci, irregularly thickened wall giving a rugose to slightly nodulose appearance, hyaline when immature, then brown; mature asci cylindric, narrowed and truncate at apex, 8-spored; paraphyses fi liform, 7–12 μm at apex, clavate, granular contents, exceeding asci by 100+ μm, hyaline. OCCURRENCE: solitary to gregarious in woody debris in mixed woods; north temperate forests from coast to coast, reaching as far south as Tennessee in the east and California in the west, and north into Alaska; infrequent. EDIBILIT Y: crisp texture but lack of fl avor or aroma yields insipid dining; avoid alcohol as it can cause an adverse reaction. COMMENTS: The spores that sometimes maintain their elongated-cubical shape even into maturity are distinctive. The interior of the somewhat simi-

lar, rare, but widespread Hydnotrya tulasnei (p. 98) is red-brown to brown to purplish brown and is much more densely folded, and the 30–44 μm spores also appear angular when densely packed in the asci but are subglobose with a rounded-warty outer layer when mature. The pale brown to redbrown Hydnotrya michaelis E. Fischer and Trappe (not illustrated) is similar to Hydnotrya cubispora macroscopically, but the spore ornamentation of its broadly elliptic spores is close to Hydnotrya tulasnei. Hydnotrya michaelis has a pleasant to garlicky odor when young and is found from Oregon to Alaska and in the Rocky Mountain region. Hydnotrya inordinata Trappe and Castellano (not illustrated) is pubescent to smooth with adhering soil and organic matter, and is gray-pink becoming dark red-brown. The interior is complex, with folds forming canals and locules 0.05– 0.3 cm broad. The canals and locules are lined with pink-white, which becomes dark red-brown as spores mature. The flesh is colored like the surface of the fruiting body or paler, and the odor and taste are not distinctive. Th is uncommon species is usually associated with roots of pines, fi rs, and hemlock in the mountains, in spring and summer, in Oregon. The round to elliptic, thick-walled Hydnotrya inordinata spores have 1 large oil drop, and are 20–30 × 20–28 μm excluding ornamentation of the crowded, flexuous, tapered, mucilage-embedded, disorderly spines that are 2–4 × 0.2–1 μm.

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Hydnotrya cubispora

H y dnotrya

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Hydnotrya tulasnei (Berkeley) Berkeley and Broome MACROSCOPIC FEATURES:

fruitbody round to irregular, 1–3 cm broad, furrowed, brown to purplish brown; interior with canals and scattered small chambers, initially pink, then brown to purplish brown; when young, odor pungent to garlic-like, in age musty and unpleasant. MICROSCOPIC FEATURES: spores 30–44 μm, irregularly globose, initially angular in ascus, with an amorphous rounded-warty outer layer at maturity, initially hyaline, becoming brown in age; asci inamyloid. OCCURRENCE: in the upper layers of the soil under pines, true fi rs, Douglas fi r, and hemlock; worldwide distribution, in North America found in northeast and northwest coastal forests and occasionally in the mountains; common. EDIBILIT Y: palatable when young. COMMENTS: Synonyms include Hydnotrya carnea (Corda) Zobel and Hydnobolites tulasnei Berkeley. Hydnobolites tulasnei R. Hesse is a synonym of Hydnotrya cerebriformis (Tulasne and C. Tu-

lasne) Harkness. Hydnotrya cerebriformis Harkness is up to 2.5 cm in diameter, is salmon-colored, and has more or less globose brown spores that measure 25–32 μm in diameter; the minute spiny ornamentation is relatively orderly (Zhang 1991). Hydnotrya subnix Trappe & Castellano (not illustrated), known from a single site in Washington state, fruits near melting snow. It has a strong garlic odor and taste, and the exterior lacks openings from the interior. Peziza ellipsospora (Gilkey) Trappe (= Hydnotrya ellipsospora Gilkey) is a western species found under oaks. It is purplish brown with a minutely villose exterior and has elliptical, minutely papillose spores 14 × 10 μm and amyloid asci. Stephensia shanorii (Gilkey) Gilkey (not illustrated), found in the upper Midwest, has a disagreeable sewer-like odor, an outer surface that is light pinkish cinnamon to reddish brown, and a whitish interior that is densely folded (spores 16.5– 19.5 μm, only 1–4 maturing).

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Hydnotrya tulasnei

H y dnotrya

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Hydnotrya variiformis Gilkey MACROSCOPIC FEATURES:

fruitbody spherical to irregularly lobed, typically flattened, lacking a stalk, 0.7–4 cm in diameter; outer surface minutely velvety and lacking warts, off-white to yellowish, buff, or cinnamon-buff; interior varying from a simple cavity with a prominent opening to branching canals resulting from infolding of the outer wall, pallid when young but brownish to pinkish orange at maturity; tissue fragile and britt le; odor and taste not distinctive. MICROSCOPIC FEATURES: spores 28–37 × 18–29 μm, broadly elliptic, at fi rst lacking oil drops but at maturity with 1 large central globose oil drop, sometimes also with de Bary bubbles, smooth then minutely pitted and wrinkled at maturity, hyaline when immature, then yellow-brown; asci 240–280 μm, clavate then cylindrical at maturity, 8-spored, inamyloid; paraphyses extending 120+ μm beyond asci, 6–11 μm wide, enlarged gradually to apex, contents fi nely granular. OCCURRENCE: solitary to gregarious in woody debris or embedded in rotted wood, associated with conifers; montane and coastal forest regions of the west from California to Alaska, reported as far east as Wyoming; infrequent. EDIBILIT Y: insipid, the lack of an odor indicating a low desirability; avoid alcohol as it can cause an adverse reaction.

There are two varieties of Hydnotrya variiformis. Variety pallida starts white to cream and ages dull orange, while variety variiformis is orange-brown in all stages. Hydnotrya cerebriformis (p. 98) is distinguished by a strong garlicky odor when mature, and globose to broadly elliptic spores that are verrucose. Geopora cooperi (p. 91) has a minutely hairy, fuzzy brown exterior to which soil tends to cling, a white interior with some brown veins, ovoid to elliptic smooth spores, and a mild to garlicky odor. Geopora cooperi is densely folded. Genabea cerebriformis (p. 84) is warted on both the exterior and interior surfaces, and has spiny round spores and a mild to pleasant garlicky odor. Genabea cerebriformis is hollow to mildly infolded. About one dozen species of Hydnotrya have been described from northern temperate forests, and several more species remain to be described. The spores of each species tend to be distinctive. If a suspected Hydnotrya species is found on the west coast under oaks or other hardwoods, check to see if the asci turn blue in Melzer’s reagent. Hydnotrya species, like all members of the Discinaceae, do not blue. Hydnotryopsis species are similar, but turn blue in Melzer’s reagent, placing them in the Pezizaceae.

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COMMENTS:

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Hydnotrya variiformis

H y dnotrya

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Imaia gigantea (Imai) Trappe and Kovács MACROSCOPIC FEATURES:

fruitbody hypogeous to emergent, 3.5–6(–15) cm broad, globose to irregular with a sterile tuft or sterile projection at the base; peridium when young is obscurely verrucose with pale, prominent, rounded to subpolygonal warts at maturity, minutely scurfy to smooth between the warts, often cracked in age, 0.4– 0.5 mm thick with an outer colored layer and thicker white to pale brown subpellis; gleba solid, white, and moist, gelatinous when young, developing brown pockets of fertile tissue separated by sterile veins, not gelatinous at maturity; odor variably reported as potatolike, somewhat disagreeable, spicy-sweet, or rich, fragrant, and fruity; taste mild. MICROSCOPIC FEATURES: spores (30–)35–40 μm when globose and (35–)40–47 × 30–36 μm when subglobose, excluding the epispore, 35– 69 μm including the epispore, hyaline and smooth initially, brownish yellow to deep orange-yellow when mature, dextrinoid, thick-walled, randomly arranged in asci; epispore 2–5 μm thick, hyaline; asci 100– 200 × 80–155 μm with 2–3 μm thick walls, globose to irregular, in youth with a stalk up to 28 μm long, stalk disappearing at maturity, 8-spored, hyaline.

OCCURRENCE:

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in moss or the upper 10–15 cm of humus in broadleaf or mixed broadleaf and conifer forests, suspected mycorrhizal hosts unknown; fall (into December in Japan); Pennsylvania to North Carolina in the Appalachian Mountains, and in Japan; common. EDIBILIT Y: good (harvested commercially on a small scale). COMMENTS: Terfezia gigantea Imai is a synonym (Trappe and Sundberg 1977). Th is species is clearly in the Morchellaceae and is distinguished from Terfezia in the Pezizaceae by much larger spores enclosed in a unique amorphous epispore. The epispore appears under electron microscopy to be permeated with minute, meandering strands or canals and under oil immersion to be obscure minute spines embedded in mucilage. In addition, Terfezia species are found in xeric to arid habitats while Imaia gigantea is found in moist, temperate forests. Phylogenetic data place Imai gigantea close to Leucangium carthusianum (p. 104), sandwiched between Disciotis venosa (p. 141) and Morchella. Leucangium carthusianum has distinctly different thinwalled, smooth spores that are fusoid-apiculate.

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Imaia gigantea

Imaia

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Leucangium carthusianum (Tulasne and C. Tulasne) Paolett i Or egon Bl ack Tru ffle

fruitbody spherical to irregular, often resembling a piece of animal dung, hypogeous, stalk absent, 0.5–8 cm broad; exterior smooth to minutely warty, dark brown to slateviolet to black, greenish to grayish at maturity; interior more or less solid pockets of fertile tissue marbled with whitish sterile veins, buff when young, maturing grayish green to greenish blue; may exude a clear latex that can stain paper pale violet over several hours; odor pleasant and somewhat resembling pineapple when young, pungent and earthy when mature; taste pleasant and mild. MICROSCOPIC FEATURES: spores 55–100 × 19– 42 μm, spindle-shaped to lemon-shaped with a distinct apiculus at one or both ends, 1 large oil droplet when mature, smooth, greenish yellow to golden; asci globose to elliptic, 8-spored. MACROSCOPIC FEATURES:

solitary to several under relatively young Douglas fi r; fall through winter; western part of the Pacific Northwest, found under oaks in France; locally common. EDIBILIT Y: prized for its pleasant and interesting aroma. COMMENTS: Picoa carthusiana Tulasne and C. Tulasne is a synonym. Kalapuya brunnea M. Trappe, J. Trappe, and G. M. Bonito is a similar species found in western Oregon and northern California, distinguished by an orange-brown outer surface, a gray marbled interior, a garlicky odor, and 32–43 × 25–38 μm elliptic, smooth, inamyloid spores lacking apiculi (Trappe, Trappe, and Bonito 2010). Kalapuya brunnea is known as the Oregon brown truffle and is reportedly delicious. OCCURRENCE:

Leucangium carthusianum

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Pachyphloeus carneus Harkness fruitbody hypogeous, globular, lacking a stalk, about 1 cm broad; exterior orange with a smooth round to irregular, lighter orange, slight depression or sometimes a raised area at the top, rest of the exterior ornamented with polygonal warts with stellate markings at the apex of the warts and an irregularly outlined base; interior white when young, becoming grayish with maturity, veins white, obscure. MICROSCOPIC FEATURES: spores globose, average 14 μm diameter, uniseriate to occasionally biseriate, mature spores light brown, ornamented with spines whose tips are connected by a second layer that covers the whole spore and appears wrinkled in places, somewhat like tent poles holding up a tent that is not taut; asci elongate or ovoid, narrowed at terminus, with a short stalk, 8-spored. MACROSCOPIC FEATURES:

originally found under Sequoias in California; a look-alike is known from many parts of North America and Mexico. EDIBILIT Y: unknown. COMMENTS: Pachyphloeus conglomeratus Berkeley and Broome (not illustrated), known at least from Europe, is macroscopically similar but has much larger spores (21–23.5 μm including blunt warts covered by a gelatinous sheath) and a smooth, dark brown peridium with a few yellow stripes or spots. There are at least four unnamed North American Pachyphloeus species in the Pachyphloeus carneus clade, characterized by species with an orange coloration. Pachyphloeus marroninus Healy, Bonito, and Guevara (not illustrated) is in a separate clade and differs from other species of Pachyphloeus in the combination of a reddish brown peOCCURRENCE:

Pachyphloeus carneus clade

Pa c h y p h l o e u s

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ridium with polygonal warts, orange areolar area, and globose spores that are mostly 16–17 μm with coarse spines mostly 2–2.5 μm tall × 1–2 μm wide, not connected at the tips by a perisporal covering (Healy, Bonito, and Guevara 2009). Rosanne Healy, who is working on a revision of the Pachyphloeus genus, reports that mature spore color, spore ornamentation, ascocarp color, gleba color, and ascus shape all appear to be congruent with molecular data. Most Pachyphloeus species fall into the Pachyphloeus carneus clade (orange, with polygonal warts, and spores like Pachyphloeus carneus), the brown to tawny to green Pachyphloeus citrinus clade (with rounded warts and spores with golftee-shaped spines), or the Pachyphloeus melanoxanthus (Tulasne and C. Tulasne ex Burke) Tulasne and C. Tulasne clade (black with polygonal warts and spores that are spiny). The name Pachyphloeus citrinus (p. 107) is frequently misapplied to orangecolored Pachyphloeus species from North America, such as Pachyphloeus carneus. Pachyphloeus vi-

rescens Gilkey and Pachyphloeus thysellii Colgan and Trappe are in the Pachyphloeus citrinus clade. Pachyphloeus melanoxanthus sensu lato is in a more basal clade. The undescribed Pachyphloeus melanoxanthus (not illustrated) look-alike from Iowa is black, olive, or yellowish when young, aging dark brown to black, verrucose, about 1 cm in diameter with yellow-green to black gleba that has obscure yellow or greenish veins, and spores that are minutely spiny, globose, and 16–22 μm diameter. Pachyphloeus cf. melanoxanthus of North America does not cluster with Pachyphloeus melanoxanthus of Europe. Pachyphloeus austro-oregonensis J. L. Frank & Trappe, at the very base of the phylogenetic tree, is closely related to Scabropeziza cf. flavovirens (Healy, draft phylogenetic tree). Pachyphloeus austro-oregonensis is purple-brown with yellow veins, has globose spores 12–16 μm including ornamentation of rods 1–2.5 × 0.5 μm, and the asci are amyloid.

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Pachyphloeus citrinus Berkeley and Broome fruitbody 0.5–3 cm across, hypogeous, roughly spherical, sometimes slightly lobed or tapered in lower part, with a depression or opening at the top or a circular furrow or cluster of furrows, with a basal tuft of mycelium, no stalk; outer surface warted typically with polygonal warts; color variable, yellow, bright to dull orange or brown, blackening when old; interior more or less solid with sterile veins that often converge toward the top; sterile veins whitish, aging grayish to grayish olive or blackish; fertile tissue white to yellowish, ages pale olive; odor mild when young, becoming unpleasant or like rott ing weeds.

MACROSCOPIC FEATURES:

Ber ry Tru ffle

MICROSCOPIC FEATURES: spores (11–)13– 15(–21) μm, round, initially smooth but developing 2 μm spines (golf-tee-shaped pegs), yellowish when mature; asci average 200 × 50 μm, clavate to saccate, 8-spored. OCCURRENCE: solitary to gregarious, reported under both hardwoods and conifers; spring and autumn; throughout most of North America and Europe; not common. EDIBILIT Y: presumably unpleasant tasting. COMMENTS: Pachyphloeus austro-oregonensis J. L. Frank & Trappe, found in southern Oregon under Oregon white oak, has a warted, purple-brown

Pachyphloeus citrinus

Pa c h y p h l o e u s

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fruiting body with an interior of labyrinthine veins lined by golden yellow hymenia and separated by gray-brown sterile tissue. Pachyphloeus thysellii Colgan & Trappe, found under Douglas fi r in Thurston County, Washington, and also known from Wisconsin and California, is minutely warty and brown to tawny with yellow veins and patches between the peridial warts, and has an interior of alternating translucent white to yellow veins (Colgan and Trappe 2004). Its spores are predominantly globose and 12–17 μm broad excluding ornamentation of symmetrical rods up to 2 × 0.7–1 μm tall. The spore wall is about 1 μm thick and cyanophilic, and the asci are inamyloid. In Healy, Bonito, and Guevara (2009), the DNA analysis of Pachyphloeus thysellii falls out very close to Pachyphloeus virescens Gilkey of North America and fairly close to

Pachyphloeus citrinus. The type locality of Pachyphloeus virescens is Los Gatos, California. It is also found in Nebraska and Mexico. Pachyphloeus virescens is typically about 1 cm in diameter, coarsely verrucose, and dull green with livid yellow gleba, has a burnt-potato odor when mature, and has globose spores (18–24 μm) with golf-tee-like spines. Species in the Pachyphloeus cf. carneus (p. 105) clade in North America were long thought to be the same as Pachyphloeus citrinus due to an erroneous determination by a European mycologist in communication with Helen Gilkey. However, Pachyphloeus citrinus is easily distinguished from Pachyphloeus carneus by having a brown, rather than orange, exterior; rounded, rather than angular, warts; and broadly clavate, rather than nearly cylindrical, asci.

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Tuber californicum Harkness fruitbody somewhat spherical, usually irregularly lobed and pitted, hypogeous, lacking a stalk, 1–2.5(–5) cm broad; exterior covered with a minute, fi ne, white pubescence, white when young, mott led with olive to brown blotches when mature, sometimes cracking in age; interior solid, fi rm, marbled, whitish when young, dark brown with labyrinthine whitish veins when mature; odor mild when young, becoming slightly garlicky or cheesy when mature. MICROSCOPIC FEATURES: spores (30–)40–50 μm, globose, alveolate, usually with 6–7 honeycomb alveoli across the diameter, dark brown when mature; asci 70–100 × 70– 90 μm, nearly spherical, short-stalked, usually 1–4 spores, rarely 5 or 6 spores; peridium averages 200 μm thick. MACROSCOPIC FEATURES:

usually solitary, sometimes gregarious at the humus-soil interface under hardwoods and conifers at low to mid elevation; late fall until early summer; found from southwest Washington to southern California, reported from Ohio; abundant in the Pacific Northwest, but the small size and often solitary occurrence make it hard to fi nd in large numbers. EDIBILIT Y: rated edible by some, bitter or insipid by others. COMMENTS: The combination of round spores and pubescence is unique to Tuber californicum. Tuber sphaerosporum Gilkey of Colorado and eastern North America also has round spores but has fewer and larger pits on the spores and lacks a pubescent exterior. OCCURRENCE:

Tuber californicum

Tuber

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Tuber canaliculatum Gilkey fruitbody globular, irregularly furrowed, hypogeous, lacking a stalk, 2– 7 cm × 2–4 cm; outer surface covered with conspicuous low, polygonal warts, red-brown to orange-brown with yellow to orangish brown tomentum in patches or in the furrows; interior marbled, brownish gray to deep gray or dark brown in age with labyrinthine white veins; odor pungent, strong, pleasing; flavor mild. MICROSCOPIC FEATURES: spores 45– 70 × 40–50 μm excluding the 4– 6 cm tall honeycomb ridges (5– 9 honeycomb cells along the spore length), elliptic, dark brown. OCCURRENCE: solitary to gregarious in the top 2– 7 cm of mineral soil in mixed conifer-hardwood forests; summer to winter; from Quebec to OnMACROSCOPIC FEATURES:

tario, south to North Carolina and Kentucky, and wherever eastern white pine has been planted; common but rarely seen. EDIBILIT Y: exquisite according to Chef James Beard (Trappe et al. 2007), not edible according to Miller and Miller (2006). COMMENTS: Tuber bisporum Gilkey is a synonym. Tuber species have litt le taste and are valued for their aroma. They should be added raw to dishes just before serving or very late in the cooking process so that the aroma is not lost. Immature truffles, with litt le aroma, will add litt le interest to the dish. It is interesting that Orson and Hope Miller, who did not care for this species, reported that the odor when mature was sweet, but not pleasant (Miller and Miller 2006).

Tuber canaliculatum

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Tuber gibbosum Harkness

Spr ing Or egon W hite Tru ffle

fruitbody spherical to irregular, hypogeous, lacking a stalk, 1–5 cm or more across; outside smooth to scabrous with furrows that are minutely pubescent, white when young, maturing light buff to brownish yellow mott led with brown to purplish brown; interior white when young, maturing brown to purple-brown to brick-red with whitish meandering sterile veins; odor mild when immature, developing a complex “truffly” odor of garlic, cheese, and spices; taste mild, becoming garlicky when mature. MICROSCOPIC FEATURES: spores 25–50 × 17–40 μm, elliptic with bluntly rounded ends, smooth when young, developing honeycomb alveoli when mature with 7–10 alveoli across the length, dark yellow-brown to brown when mature; asci sacMACROSCOPIC FEATURES:

cate to pyriform, usually 1– 6 spores, rarely 7 or 8 spores. OCCURRENCE: solitary to gregarious mainly or exclusively with 10- to 60-year-old rapidly growing Douglas fi r; winter and spring; west of the Cascades from southern British Columbia to northern California, from sea level to about 2,000 feet; common in the Pacific Northwest. EDIBILIT Y: choice when mature, fl avorless when immature. COMMENTS: Tuber giganteum Gilkey is a synonym. Tuber oregonense Trappe, Bonito, and P. Rawlinson nom. prov. has only recently been recognized as a separate species with a fall rather than a spring maturity, a thin, almost translucent peridium, and sometimes with reddish tones. Tuber oregonense,

Tuber gibbosum

Tuber

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the fall Oregon white truffle, is white when young, soon yellow to olive mott led with cinnamon or brown to reddish brown blotches; peridium thin, almost translucent. The interior is fi rm, white when young, and ages brown with white marbling. The odor is mild when young, becoming a complex garlic, cheese, and spice odor when mature. The spores are 25–50 × 17–40 μm, elliptic or tapered to a point at both ends, and have honeycomb ornamen-

tation with 4– 6 alveoli spanning the length. Both Tuber oregonense and Tuber gibbosum are large and distinguished by growth under young, vigorously growing Douglas fi r, and both are avidly sought by commercial collectors. Unfortunately, widespread raking for these two species results in the dual shame of habitat destruction and harvest of mostly immature material that has litt le aroma and thus is of litt le interest as an edible.

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Tuber lyonii Butters

Peca n Tru ffle

fruitbody globular, lobed, hypogeous, lacking a stalk, 1–4 cm across; outside smooth, roughened in the furrows, orangish brown to reddish brown; interior white when young, maturing brown with white marbling; odor a complex mix of garlic, cheese, and spices. MICROSCOPIC FEATURES: spores 30–37 × 22–24 μm, elliptic, when mature developing tall spines connected by low lines. OCCURRENCE: solitary to gregarious under oaks and pecan; year-round; from northeastern Mexico to Ontario and the Great Plains to the East Coast; common. EDIBILIT Y: choice when mature. MACROSCOPIC FEATURES:

COMMENTS: Also spelled as Tuber lyoniae Butters. Tuber texense Heimsch is a synonym (Trappe, Jumpponen, and Cázares 1996). Tuber lyonii is frequently raked up with pecans during pecan harvest and can be abundant in pecan orchards where it is commercially harvested. It is the most popular native truffle of the eastern United States. Gilkey (1954) considered Tuber lyonii to be one of many synonyms of a widely variable species, Tuber candidum Harkness. However Trappe et al. (2007) implicitly recognize the two as separate species with different odors at maturity, different edibility, and different microscopic characteristics. Tuber candidum is smooth-skinned, light yellowish brown to

Tuber lyonii

Tuber

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reddish-brown, and associated with oaks, pines, and Douglas fi r, and has thick-walled spores that are 19–42 × 14–34 μm excluding ornamentation of spines that are 2–5 μm long at maturity. Tuber quercicola J. L. Frank, Southworth, and Trappe is closely related to Tuber candidum. Tuber quercicola has very similar thick-walled spores that are 20– 45 × 15–36 μm with 2–5 μm spines at maturity, but is distinguished from Tuber candidum by a rough,

scaly peridium that is light reddish brown to dark red. Tuber quercicola is found year-round under oaks and pines at warm, dry sites in western North America. The European species Tuber rufum Pico has similar long spines, and this name has been applied to Tuber quercicola in the past. Sean Abbott has found an unidentified species with long-spined spores in Alberta, which we have illustrated as Tuber cf. rufum in the picture key.

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Tuber shearii Harkness fruitbody globular, hypogeous, lacking a stalk, 0.5–2 cm wide; exterior smooth to roughened, light dull yellow to yellowish brown with white furrows; interior marbled light brown with white veins; odor and taste slightly pungent. MICROSCOPIC FEATURES: spores 32–56 × 28–42 μm excluding ornamentation of alveoli in a honeycomb pattern 3–4 μm tall with 3– 9 meshes along the spore length, broadly elliptical to subglobose, light to dark brown; asci 50–70 μm in diameter, nearly spherical. OCCURRENCE: solitary to several, most commonly under eastern white pine but also under Douglas fi r and mixed conifer-hardwood stands; summer MACROSCOPIC FEATURES:

through fall; transcontinental in southern Canada and the northern United States, sometimes as far south as North Carolina, Arizona, and California; infrequent. EDIBILIT Y: edible. COMMENTS: Terfezia oligosperma Shear is a synonym. Though widespread, T. shearii is not common. Because it is small and has only a slight odor, it is of litt le interest for cooking. Tuber anniae W. Colgan & Trappe is a highly similar small species found in the Pacific Northwest under Douglas fi r. Tuber anniae (not illustrated) has a crisp, bald exterior that is pale yellow when young and brown to dark olive-brown with grayish white furrows and patches when mature; the interior is dark

Tuber shearii

Tuber

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brown marbled with whitish veins alternating with brownish gray veins (Colgan and Trappe 1997). A mild to slightly paint-like odor may enable one to distinguish T. anniae from T. shearii without resorting to DNA analysis. Tuber monticola Harkness (not illustrated) is a small dingy white to gray species with mostly round spores (32–40 × 28–34 μm) that have 10–16 minute alveoli across the spore surface. Tuber monticola is common in the mountains of the Pacific Northwest under trees of the Pinaceae. Tuber separans Gilkey, infrequently found solitary to several under oaks and conifers in the Pacific Northwest and in eastern North America, is 0.5–1 cm across with a minutely scabrous exterior that is white when young and maturing earthy brown, with an interior similar in color to the exterior, and inconspicuous veins. Tuber separans spores are 48–58 × 40–56 μm, broadly elliptical, in a honeycomb pattern with 3–10 × 7–11 (commonly 6–7 × 8– 9) alveoli across the diameters. Tuber levissimum Gilkey (not illustrated) is another small, clay-brown species, with large internal veins that are inconspicuous in color. At maturity, the spores of T. levissimum are dark brown, 36–58(65) × 32– 52 μm, mostly broadly elliptical, and alveolate, with 3–10 × 4–13 alveoli across the diameter of the spore. T. levissimum has a smooth, thick peridium (400– 850 μm thick) and T. separans has a minutely scabrous thin peridium (200–260 μm thick). Seven additional small (< 1.5[–3] cm diameter), difficult-to-distinguish Tuber species (not illus-

trated) were described by Guevara et al. (2013). Of these, Tuber beyerlei Trappe, Bonito, and Guevara and Tuber lauryi Trappe, Bonito, and Guevara are western species. Tuber walkeri Healy, Bonito, and Guevara is found in the upper midwestern United States. Tuber miquihuanense Guevara, Bonito, and Cázares is known only from Tamaulipas, Mexico, in pine-oak forests. Tuber castilloi Guevara, Bonito, and Trappe, Tuber guevarai Bonito and Trappe, and Tuber mexiusanum Guevara, Bonito, and Cázares are found associated with Quercus species in eastern Mexico. Tuber mexiusanum is also found in the eastern United States under Populus deltoides, as well as in Minnesota and Iowa in mixed hardwood forests (Guevara et al. 2013). In these seven species, fruitbody color ranges from cream to reddish brown and all have globose to elliptical ascospores with alveolate reticulation. Two additional small Tuber species—Tuber gardneri Gilkey and Tuber whetsonense J. L. Frank, Southworth, and Trappe— are distinguished by thick-walled asci. Tuber gardneri (not illustrated) fruitbodies are yellow-brown to brown, verrucose, and have a brown to purplish brown interior with conspicuous white veins. Tuber whetstonense (not illustrated) fruitbodies are light brown to dark reddish-brown with a dark grayish brown interior marbled with narrow white veins (Frank, Southworth, and Trappe 2006a). Tuber gardneri is found under Douglas fi r while T. whetstonense is found under Oregon white oak.

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Chapter 4 Pezizomycetes

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he Pezizom ycetes conta in just one order, the Pezizales. Currently, there are sixteen recognized families within this order, with roughly 200 genera and between 1,600 and 2,000 species. The prized black truffles, white truffles, desert truffles, and morels belong to this class, along with all of the large fleshy ascomycetes. We have dealt with the truffles in a separate section. In this section, we deal with the remaining epigeous genera of the Pezizales. Twelve of the sixteen families contain species that are likely to att ract the attention of mushroom hunters. The fungi in this class are mycorrhizal, saprotrophic, or parasitic on plants. They can be found growing on soil, wood, leaves, decaying vegetation, or dung, or buried in the uppermost soil horizon (the truffles). Cold to temperate regions have a more diverse mycoflora than is found in tropical regions, with spring being the season of greatest abundance. The fruitbodies are open apothecia or various closed structures derived from apothecia. They range in size from less than 1 mm to greater than 15 cm. Some species have a stalk, although most are sessile. They are typically fleshy and often fragile. The asci of members of the Pezizales typically release their spores by rupturing at the apex to form an eccentric lid known as an operculum. The ascospores are single-celled, usually symmetrical, and can be spherical, ellipsoidal, or fusoid. Sometimes they are ornamented with warts, ridges, or spines. The anamorphs of some species are known, although the majority of species are only known from the teleomorphic state.

T

The Ascobolaceae

subimmersed or superficial, and 0.2–12 mm broad. The asci are weakly amyloid all over. The spores lack gutt ules and are hyaline, globose to elliptical, and ornamented at maturity with a purplish to brownish material that will dissolve in KOH. The Caloscyphaceae The Caloscyphaceae contains a single species, Caloscypha fulgens, which is a 0.5–4 cm broad conifer seed pathogen that grows on the soil in early spring, some years in tremendous abundance (Paden, Sutherland, and Woods 1978). Some authors include this species with the Pyronemataceae. The Chorioactidaceae The Chorioactidaceae is represented by four very different species: Chorioactis geaster, Neournula pouchetii, Wolfina aurantiopsis, and Pseudosarcosoma latahensis (Pfister, Slater, and Hansen 2008). Chorioactis geaster looks like a cigar stuck in the ground and opens to resemble a tulip flower that is brown outside and butterscotch inside. Neournula pouchetii is a stalked deep cup with a pinkish to gray interior and a whitish exterior that darkens on handling. Wolfina aurantiopsis is a stalkless shallow cup with a pale yellow, somewhat orange upper surface; a minutely roughened, brownish to black lower side, and tough, whitish flesh that becomes corky on drying. Pseudosarcosoma latahensis is a large, top-shaped, dark purple to black jelly-like fungus. As of 2013, MycoBank recognizes these four species as members of the Pezizales but does not place them in any family.

The Ascobolaceae is represented by the genus Ascobolus (Brummelen 1967). The species are mostly found on dung, although some are found on highpH soil, burnt ground, or on herbaceous stems. The apothecia are globose, turbinate, or disc-shaped,

The Discinaceae, Helvellaceae, and Rhizinaceae are found worldwide but are most diverse in north-

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ern temperate regions. We treat these three families together, while acknowledging that some authors recognize the close relationship between the Discinaceae and Rhizinaceae and consider them as being part of the Helvellaceae (e.g., Abbott and Currah 1997; Weber 1995). The Rhizinaceae is a small family centered on Rhizina undulata, a large, undulate species attached to the ground by numerous white rhizomorphs that cover the underside of the crust-like cup. Based on molecular data, the family now includes the genus Psilopezia and an anamorphic, pathogenic species, Phymatotrichopsis (Marek et al. 2009). The Discinaceae family includes just one epigeous genus, Gyromitra (although some authors separate Discina [McKnight 1969] and Pseudorhizina from Gyromitra), plus the hypogeous genus Hydnotrya (see chapter 3, Hypogeous Ascomycetes: The Truffles). Molecularly, Discina and Pseudorhizina are both nested with Gyromitra. Five subgenera are currently recognized within Gyromitra: Caroliniana, Discina, Pseudorhizina, Gyromitra, and Melaleucoides (Methven, Zelski, and Miller 2013). The largest fruitbodies of any of the ascomycetes are found in the genus Gyromitra. For example, some huge specimens of Gyromitra caroliniana, the red false morel, can weigh more than seven pounds. Gyromitra species vary from disclike subsessile species to tall stalked species with brain-like or saddle-shaped caps, with colors ranging from yellowish brown to reddish brown to purplish brown. The spores are elliptical to fusoidal, nearly smooth to distinctly rough or reticulate, and contain one, two, or three large gutt ules. The spores of some species have distinctive apiculi (Harmaja 1969b, 1973; Kempton and Wells 1973). Gyromitra esculenta is the type of the genus and forms a tight group with G. infula and G. ambigua. Gyromitra caroliniana, G. brunnea, and the European G. parma form a second tight group. A third

tight group includes Gyromitra ancilis, G. olympiana, G. leucoxantha, G. gigas (European), G. montana, and G. korfii. Gyromitra melaleucoides is very distantly related to the rest (Methven, Zelski, and Miller 2013). Members of the Pseudorhizina subgenus are dark brown to reddish brown, up to 14 cm wide × 8 cm tall, and thin-fleshed with a deeply fluted stalk that often has red to pinkish tints. The spores are elliptical to globose, and contain one or two large gutt ules. The Helvellaceae includes the epigeous genera Helvella, Midotis, Wynella, and Underwoodia, as well as the hypogeous genera Balsamia and Barssia (see chapter 3, Hypogeous Ascomycetes: The Truffles). The epigeous species found in these genera grow on the ground or sometimes on rott ing wood. They may or may not have a significant stalk. Their fertile surfaces can be disc-like, cup-like, an inverted cup, brain-like, or saddle-shaped and measure 0.5–8 cm broad. They are whitish, grayish, brownish, or black. The outer surfaces are glabrous to pubescent. Their spores are typically elliptical, smooth, and contain one large gutt ule (Dissing and Lange 1967; Kempton and Wells 1970; Weber 1972, 1975; Harmaja 1974b, 1979b; Häff ner 1987; Abbott and Currah 1997). Midotis species typically start out subglobose and closed, expanding and usually becoming vertically elongated on one side when mature. The fruitbodies can be yellow, brown, dark blue, or olivaceous, and the exterior is covered with short hair-like hyphae that typically turn momentarily violet with KOH. Midotis spores typically have 2–3 gutt ules. Wynella is monotypic, 3– 6 cm tall, rabbit-ear-shaped and blood-red on the interior surface. In North America, Under woodia is represented by one species, 4–12 cm tall, with a longitudinally grooved cream to tan stalk that is rounded at the top and lacking a distinct cap. Sean Abbott (personal communication) notes that “the few extant species of Underwoodia, each on separate continents, makes me think it may represent an evolutionarily primitive form in the Helvel-

Overview

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The epigeous genera of the Morchellaceae include Disciotis, Verpa, and Morchella. There are at least two unnamed species of Disciotis in America that are similar to Disciotis venosa of Europe. They are broadly cup-shaped to repand, subsessile, and light to dark brownish, and their surfaces are wrinkled or folded with a vein-like pattern. Their spores are elliptical and lack gutt ules. The genus Verpa is represented by V. bohemica and the V. conica complex. (Note: Some authorities recognize an additional genus, Ptychoverpa, in the Morchellaceae and consider Ptychoverpa bohemica the accepted name for what we are calling Verpa bohemica.) Verpa species have a distinct stalk and a cap. The underside of the top of the cap is attached to the top of the stalk, and the cap hangs like a skirt or a bell. They can be up to 10 cm tall. The genus Morchella is represented by the morels. Morels also have a distinct cap and stalk, but the cap is hollow, measures 2–10+ cm tall, and the bottom of the cap is att ached to the top of the stalk. Morels are probably the single most highly prized and collected culinary mushrooms in all of North America and are probably the most studied of the wild edible mushrooms. All morels known in the world come from a common ancestor in western North America that diverged from Verpa and Disciotis about 174 to 100 million years ago. Genetically, Morchella rufobrunnea lies at the base

of the genus Morchella, having evolved in western North America an estimated 125 million years ago (O’Donnell et al. 2011). The phylogenetic tree split roughly 100 million years ago to produce the Elata clade (black morels) in western North America and the Esculenta clade (yellow morels) in eastern North America. Representatives of the Elata clade started diversifying in western North America 75 million years ago, producing several more Elata clade species in the west. Elata clade representatives fi rst migrated from western North America to Asia and to Eastern North America 10 to 5 million years ago. Then 5 to 2 million years ago, new Elata clade species spread from Asia both to Europe and back to western North America. One of the new European Elata clade species made the migration to eastern North America, where a new species evolved. The Esculenta clade representative remained unchanged in eastern North America from about 100 million years ago until 60 to 55 million years ago, when a single new yellow morel species appeared in Europe that has since remained unchanged. Between 30 and 10 million years ago, several additional Esculenta clade species evolved in eastern North America. Then roughly 11 to 9 million years ago, the Esculenta clade had a second range expansion, this time to Asia. About 2 million years ago, two of the Asian species gave rise to two new Esculenta clade morels in Europe. All this time Morchella rufobrunnea remained relatively unchanged in western North America. Putt ing a name on a morel is complicated by the fact that morels are multinucleate, with as many as fi ft y or more haploid nuclei per cell. Only one pair of two compatible haploid nuclei migrates into each developing ascus, but each cell in morel hyphae contains dozens of different possible pairs. Each one of the unique pairs is capable of forming a genetically distinct ascus. Practically, this means that in a cluster of 4 to 20 morels, each morel in the cluster is likely to be genetically distinct. Th is is

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laceae, which is where it always comes out in the DNA trees. In this scenario, evolution of a cap would be an advanced trait.” We have included one unnamed Gyromitra (Gyromitra mwb060911) because it is encountered year after year at many sites in the Gifford Pinchot National Forest of Washington State. It is so distinctive that we have named it “mini peach pig’s ears” for its peach coloration and diminutive size relative to the common and widespread Gyromitra ancilis. The Morchellaceae

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Clowez monograph have precedence, although it is clear that Clowez named many but not all of the species studied by O’Donnell et al. (2011) and some species clearly not studied by O’Donnell et al. A European-American team is working on comparing the DNA sequences of morels included in the Clowez world monograph (Clowez 2010) and from the North American morels studied by Kuo et al. (2012), as well as other collections. The preliminary results of their work (with DNA molecular markers ITS, LSU, TEF1, and RPB2) as of March 1, 2013, were forwarded to us with the reservation that for some critical taxa a fi nal decision has not yet been made (F. Richard, R. Courteciusse, P. Clowez, K. O’Donnell, K. Hansen, and P. A. Moreau, personal communication).

a far different picture than is found with Basidiomycetes, which are dikaryotic (two nuclei per cell) and thus much simpler to study. You can read more about this and reference the extensive research on morels in Pilz et al. (2007). Two morphologically identical morels can be genetically very distinct, while two morphologically different morels can be closely related. Morels are able to exhibit a wide variety of forms and observable features and often intergrade among described or presumed species. A young morel can look like one species but, left alone to grow, will later resemble a second species, and still later the same morel will be presumed to be a third species. At all times, however, it will distinctively be a morel. The only question will be, which morel is it? It is difficult to differentiate morel species using either macroscopic or microscopic means. Ultimately, we must rely on a DNA analysis. Fortunately, all of the known morel species are choice edibles. O’Donnell et al. (2011) used DNA to demonstrate that the European species are different from North American species they had studied. They also showed that the three black morel species then known from eastern North America are different from the ten black morel species then known from western North America. While they identified five different yellow morel species in North America, they found only one to be present throughout North America. Th ree yellow morel species were found only in eastern North America, and one yellow morel species was found to have a narrow distribution from Michigan to British Columbia, reaching no further south than South Dakota. Kuo et al. (2012) published names for the North American morels whose genetics had been published in O’Donnell et al. (2011). These authors were unaware of a world monograph on morels completed in 2010 and published April 16, 2012, in French in the Bulletin de la Société Mycologique de France, a journal not covered by the major scientific abstract services (Clowez 2010). The names from the

Of the 29 genera in the Pezizaceae, five epigeous genera—Iodophanus, Pachyella, Peziza, Plicaria, and Sarcosphaera—contain species that are likely to att ract the attention of mushroom hunters in North America. Iodophanus is a small genus represented by one easily overlooked species, Iodophanus carneus. It is characterized by 1–3 mm broad apothecia, asci that are amyloid overall, and hyaline, wrinkled spores that lack gutt ules. Pachyella contains three North American species that are typically found broadly attached to waterlogged rotten wood. They are 0.5–1.2 cm broad, and have nonamyloid to weakly amyloid asci and bigutt ulate spores (Pfister and Candoussau 1981). Sarcosphaera contains a single species, Sarcosphaera coronaria, the 3–10 cm broad tulip cup (Pouzar 1972). Peziza contains the greatest number of large ascomycetes in the Pezizaceae family, with at least 50 species in North America (Hansen, Læssøe, and Pfister 2002; Hansen, LoBuglio, and Pfister 2005). Peziza apothecia are 0.1–10 cm broad, and sessile or, rarely, with a short stalk. The hymenium is whitish, yellowish, brownish, greenish, viola-

Overview

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The Pezizaceae

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Of the 80 genera in the Pyronemataceae, we have included representatives from the 30 genera with showy species. All are nonamyloid. The genus Otidea includes species having the largest apothecia. They typically are ear-shaped. The spores are narrowly elliptical, smooth, and bigutt ulate (Kanouse 1947; Liu and Zhuang 2006). The other included genera have very small to medium-sized apothecia and are sessile to substipitate, pulvinate, disc-shaped, or cup-shaped. The hymenium may be whitish, grayish, yellowish, orange, reddish, bluish, or purplish. They have glabrous to hairy exteriors. Spores are globose to fusoid, with or without gutt ules, and smooth or ornamented. The following genera are typically found on burnt ground. Anthracobia species are grayish brown, yellowish, or red, 0.2–1.2 cm broad, with appressed brownish hairs on the underside of the apothecium, and have elliptical spores that are smooth and bigutt ulate. Geopyxis species are ochraceous to greenish-brown, goblet-shaped with a yellowish white margin, and 0.5–1.2 cm broad. They have elliptical smooth spores that lack guttules. Neottiella species are orange or reddish cups measuring 0.2–1.5 cm broad. They have slender, hy-

aline, hyphae-like hairs on the underside. The elliptical spores are either ornamented or smooth and have gutt ules. Pyronema species are densely gregarious turbinate to pulvinate discs. They are pink or reddish, measure 1–2 mm wide, and grow on whitish mycelium. The underside of the apothecium is bald to furfuraceous. They have smooth elliptical spores that lack gutt ules (Moore and Korf 1963). Rhodotarzetta species have fruitbodies that are pink cups measuring 0.5–1.2 cm broad. The underside of the apothecium is bald. The smooth spores are elliptical and have two gutt ules. Sphaerosporella species are turbinate to disc-shaped, brownish, and measure 0.2– 0.6 cm wide. They have short, brownish, pointed hairs on the underside, and globose spores. Tricharina species have apothecia that are yellowish discs to shallow cups that measure 0.5–1 cm wide. They have projecting brownish hairs on the underside and smooth elliptical spores (Yang and Korf 1985). Trichophaea species are grayish white discs or shallow cups that measure 0.2– 1.2 cm broad, with hyaline or brownish hairs on the underside. The spores are subglobose to fusoid and smooth or ornamented, with gutt ules (Kanouse 1958). The species that are found on wood include the following. Chaetothiersia vernalis is deeply cupshaped, measures 0.5–3 cm wide, is colored white to gray, and has an exterior covered with erect brown hairs (Perry and Pfister 2008). The smooth spores are elliptical and egutt ulate. Miladina species grow on wet wood and have yellowish to orange pulvinate apothecia that measure 0.5–1 cm broad. The underside is bald, and the spores are broadly elliptical, verruculose, and multigutt ulate (Pfi ster and Korf 1974). Pseudaleuria quinaultiana is a reddish orange disc that measures up to 3.5 cm broad. Its underside is felty with long whitish flexuous hairs. The spores are elliptical, smooth, and egutt ulate. Scutellinia species are whitish to orange to scarlet discs or shallow cups that measure 0.5–1.5 cm wide. The exterior and margin

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ceous, or almost black with a furfuraceous exterior. The asci are cylindric and typically amyloid only at the operculum. The spores are elliptical to fusoid, smooth or ornamented, with or without guttules, and are hyaline or pale brownish. Peziza species are found on soil, on burnt ground, on dung, and on wood. We know of just three Plicaria species in North America. All are found only on burnt ground. The apothecia are sessile, cup-shaped, 2–8 cm broad, brownish to black, with a glabrous to furfuraceous exterior. The spores are globose, smooth or ornamented, hyaline, and pale brownish (Norman and Egger 1996). The Pyronemataceae

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of the apothecia are covered with rooting, dark, eyelash-like hairs. The spores are elliptical to globose and are ornamented (Denison 1959; Schumacher 1988, 1990; Yao and Spooner 1995). Wilcoxina species have disc-shaped apothecia that measure 1.5–2.5 mm wide and are pale mouse-gray, sessile, and centrally att ached. The outer surface is covered overall with hairs. The spores are egutt ulate and situated in the upper half of the ascus (Yang and Korf 1985). We have included the following taxa found on dung or decaying vegetation. Byssonectria species form gregarious, orange, 0.5–3 mm wide apothecia on a whitish subiculum. The spores are smooth, fusoid, and contain gutt ules (Pfister 1993, 1995). Cheilymenia species have disc- to cup-shaped apothecia that range from whitish to yellow to orange or red, have a distinct margin, and measure 0.5–20 mm broad. The underside of the disc has septate whitish to brownish hairs. The spores are elliptical, smooth or with a verrucose to echinulate episporium, and egutt ulate (Denison 1964; Moravec 1990). Species of Lasiobolus have white to yellowish or reddish apothecia that are globose to disc-shaped and measure 0.2–1 mm wide. They have thick-walled (usually nonseptate), stiff, pointed hairs on the underside. Spores are elliptical, smooth, with a gelatinous coating, and egutt ulate. We have included the following taxa that generally grow on soil. Acervus epispartius is a shallow cup that measures 0.5–1.5 cm wide and looks like the orange peel fungus Aleuria aurantia; it differs by having an apothecium that is covered with a membrane when young and elliptical, smooth spores that are unigutt ulate (Pfi ster 1975; Pfi ster and Bessette 1985). The fruitbodies of Aleuria aurantia are orange cups or discs that are 1– 12 cm broad. Their spores are regularly reticulate with prominent ridges and two gutt ules. Humaria species are deeply cup-shaped, measure 0.5–3 cm wide, and are white to grayish yellow. Their exte-

rior surface is covered with stiff, brownish, multiseptate hairs. The spores are elliptical and biguttulate. Jafnea species are typically 1–5 cm wide × 1–7 cm tall, have a short stalk, and are whitish to brownish. Their exterior is covered with soft brown hairs and their spores are elliptical to fusiform, bigutt ulate, and verrucose at maturity (Korf 1960). Lamprospora species are 1–3 mm broad and globose to disc-shaped, with an often dentate margin, orange to red hymenium, and a concolorous to more yellowish, glabrous underside. The hyaline, globose to subglobose spores have 1 large guttule and ornamentation of warts, spines, or reticulations that easily dissolve in 2–5 percent KOH. Melastiza species are often gregarious orange or red cups that are 0.3–2.5 cm wide and have an underside that is covered with appressed dark hairs. The spores are elliptical and ornamented (Yao and Spooner 1995). Octospora fruitbodies are yellow to orange discs or shallow cups, often found with mosses. They measure 0.1–1 cm broad, have a margin that is often dentate, and have a bald underside. Their spores are smooth or ornamented and unigutt ulate. Species of Pseudombrophila have turbinate, cupulate or disc-like apothecia that measure 0.2–2 cm wide. They range from whitish to ochraceous to brownish and have appressed hairs on the exterior surface. Their spores are elliptical, smooth to delicately ornamented, and lack guttules. Pulvinula species have white or yellowish to red disc-shaped to pulvinate apothecia. They are 0.5–12 mm wide with a bald exterior surface. Their spores are globose or elliptical, and unigutt ulate (Pfi ster 1976). The apothecia of Rhodoscypha species range from disc- to cup-shaped, sessile to substipitate with a prominent shaggy margin, and measure 0.5–1 cm broad. They are pink to whitish, with an exterior surface that is whitish due to being clothed in a dense mat of short, white, flexuous hairs. Their spores are fusoid and gutt ulate. The apothecia of Smardaea species are disc-shaped, measure 0.5–2 cm broad, and are purplish or vi-

Overview

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olet. Their spores are elliptical to fusoid and are coarsely ornamented with rounded warts. Sowerbyella apothecia are greenish yellow to orange and are cup-shaped with a rooting stalk. They measure 0.5–5 cm broad and have elliptical, ornamented spores (Moravec 1985; Yao and Spooner 2006). Tarzetta species are hemispheric to disc-shaped and measure 0.2–3 cm wide. The apothecia range from dirty yellowish to pale orange, with short whitish hairs covering the exterior surface. The spores are elliptical to fusoid and gutt ulate. Tricharina species have apothecia that are broadly sessile and shallowly cup-shaped to convex, and measure 0.2–1 cm broad. They are grayish, pinkish, or ochraceous, with marginal hairs and egutt ulate spores that almost fi ll the ascus.

cup-shaped, bright red to scarlet, measure 1– 6 cm broad, and grow on wood and sticks. Their exterior surface is tomentose and whitish to pinkish buff or reddish orange to pinkish red. The spores are elliptical to cylindric, blunt, smooth, and with granular contents at the poles (Harrington 1990, 1998; Harrington and Potter 1997). Wynnea is represented by two very different looking species. Wynnea americana resembles rabbit ears and forms a cluster of pinkish to brownish orange apothecia that arise from a tough brown sclerotium. The fruitbody is 2–14 cm wide × 6–13 cm high and has a blackish brown to reddish-brown exterior (Korf 1950; Pfister 1979). Wynnea sparassoides lacks a sclerotium. Its numerous apothecia are united into a beige to yellow-brown, cauliflower-like mass on a long unbranched stalk (Korf 1956).

The Sarcoscyphaceae Five of the eleven genera of the Sarcoscyphaceae are included in this work. Cookeina are tropical to subtropical species that have a stalk and form yellow to red, deeply cup-shaped apothecia. They grow on wood that is in the early stages of decay, on leaves, or sometimes on fruits. They are 1–2 cm wide × 1–3 cm high with an exterior that is clothed in short to long whitish to yellowish hairs. They have large, smooth to fi nely wrinkled, elliptical spores. Microstoma has just two species: M. protractum and M. floccosum. Microstoma species grow on wood, are red to scarlet, and are deeply cup-shaped with a short stalk. They are 1– 3 cm broad and up to 5 cm tall. The exterior surface is covered with short to long white hairs. Their spores are elliptical with small gutt ules. Pithya is a small genus represented in this book by two species. Pithya apothecia are disc-like, with or without a short stalk, colored dull to bright orange, and found on recently killed conifer foliage. They measure 0.1–1.5 cm wide and have globose spores with 1 large oil drop at maturity. Sarcoscypha species are

Pezizom ycetes

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The Sarcosomataceae Five of the nine genera of the Sarcosomataceae contain large, showy Ascomycetes (Korf 1957a, 1957b; Paden 1983). All are saprotrophic on rotten or buried wood. Galiella contains a single species, Galiella rufa. It has thick, cup-like apothecia with a pale orange to reddish brown inner surface, a brown to blackish brown exterior covered with matted wooly hairs, and rubbery-gelatinous flesh. Plectania species are at most slightly gelatinous cups ranging in size from 0.5 to 4 cm broad, that may or may not have a stalk. They are dark brown to black, with an outer surface covered with conspicuous to inconspicuous matted hairs. The spores are elliptical to fusoid, smooth, and usually eguttulate at maturity (Kanouse 1949). Pseudoplectania, represented here by two species, is distinguished from Plectania by having globose spores. Their exterior surface is glabrous to tomentose. Sarcosoma is represented by Sarcosoma globosum. The other included genus in Sarcosomataceae is Urnula.

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Acervus epispartius (Berkeley and Broome) Pfister fruitbody 5–16 mm wide, cup- to shallowly cup-shaped, stalkless or with a rudimentary stalk; inner surface orange to yelloworange; outer surface yellow, felty, completely covering the fruitbody of young specimens and tearing to form a flap that is discarded to one side, exposing the orange inner surface at maturity. MICROSCOPIC FEATURES: spores 6– 9 × 3–4 μm, elliptic with blunt ends, smooth, usually with 1 oil drop, hyaline; asci 100–140 × 7–10 μm, cylindric to clavate; paraphyses slender, clavate, curved, orange-colored. MACROSCOPIC FEATURES:

in groups or clusters on soil, among leaf litter or mixtures of wood chips, hay, and dung; summer and fall; eastern Canada and throughout New England; rare. COMMENTS: Peziza epispartia Berkeley and Broome is a synonym. Compare with Aleuria aurantia (p. 126), which forms much larger cups that do not have a flap of tissue that covers the inner cup surface. OCCURRENCE:

Acervus epispartius

Acerv us

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Aleuria aurantia (Fries) Fuckel

Or a nge Peel

fruitbody 1–10 cm wide, cup- to shallowly cup-shaped, stalkless or with a rudimentary stalk; inner surface bright orange to yellow-orange, smooth; outer surface pale yellowish orange to yellow, slightly scurfy; margin wavy and often torn at maturity. MICROSCOPIC FEATURES: spores 17–24 × 9–11 μm, elliptic, smooth at fi rst, soon covered by a coarse reticulum, usually with one or more projecting spines at each end, often with 2 large oil drops, uniseriate, hyaline; asci 175–250 × 12–15 μm, cylindric to subcylindric; paraphyses tips strongly clavate to subglobose, fi lled with orange granules. MACROSCOPIC FEATURES:

solitary, in groups, or in clusters in grassy areas on disturbed soil, in gardens, and along roadsides; summer and fall, also winter in California; widely distributed in North America; fairly common. COMMENTS: Sowerbyella rhenana (p. 256) is similar but has a distinct stalk. Acervus epispartius (p. 125) is also similar but the outer surface of its cup is felty and forms a flap of tissue that completely covers the inner cup surface of young specimens. OCCURRENCE:

Aleuria aurantia

Pezizom ycetes

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Anthracobia maurilabra (Cooke) Boudier fruitbody broadly cupshaped to disc-shaped, lacking a stalk, 0.2–1.2 cm broad; inner surface grayish brown; margin often somewhat inrolled and very wavy in outline; outer surface concolorous with the inner surface, covered with short brownish hairs. MICROSCOPIC FEATURES: spores 16–21 × 8–10 μm, elliptic, smooth, bigutt ulate. OCCURRENCE: gregarious on burnt ground yearround; reported from eastern North America and known from Europe. MACROSCOPIC FEATURES:

First named Peziza maurilabra Cooke, this species was reclassified as Humaria maurilabra (Cooke) Saccardo before being moved to Anthracobia. There are eleven named species in the genus and all are characterized by growth on burnt ground, an exterior covered with short brownish hairs, and a fertile inner surface that is grayish brown, yellowish, ochraceous, orange, or red.

COMMENTS:

Anthracobia maurilabra

A nthr acobi a

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Anthracobia melaloma (Albertini and Schweinitz) Arnould Bu r n Site Ochr e Cup

fruitbody cup-shaped to concave to flattened, lacking a stalk; 0.1– 0.5 cm broad; upper surface yellowish orange; underside slightly paler than upper surface, darkly punctate with tuft s of appressed brownish hairs. MICROSCOPIC FEATURES: spores 14–22 × 7–11 μm, oblong-elliptic, bigutt ulate; asci 8-spored, 160– 200 × 11–15 μm; paraphyses slender, septate, clavate (6 μm at tips), inamyloid; hairs up to 80 × 13 μm, multiseptate, thin-walled. OCCURRENCE: gregarious to massed on burned ground and burned wood; spring through early winter; western North America and Europe. COMMENTS: Initially this species was placed in the genus Peziza. Subsequently it was moved to the MACROSCOPIC FEATURES:

genera Humaria, Lachnea, Patella, and Pyronema before fi nally being classified in the genus Anthracobia. There are several tiny stalkless species associated with burnt ground and burnt wood that have a more or less orange upper surface. Anthracobia macrocystis (Cooke) Boudier is more reddish orange; the hairs are shorter, more rudimentary and have a single septum with paraphyses that stain green in Melzer’s reagent. Pyronema omphalodes (p. 240) is composed of confluent masses of minute orange cushion-shaped fruitbodies surrounded by cobwebby mycelium. It grows on burned or sterile ground, has a conspicuous white cobwebby subiculum, and hyaline, smooth, elliptic spores measuring 10–13 × 5–8 μm.

Anthracobia melaloma

Pezizom ycetes

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Ascobolus carbonarius P. Karsten fruitbody 3– 6 mm wide, rounded and cup-shaped when young, becoming shallowly cup-shaped when mature, stalkless; inner surface greenish at fi rst, soon becoming brown and fi nally purple-brown, conspicuously dotted by the ends of protruding asci; outer surface coarsely scurfy, yellowish green at fi rst, soon becoming dark brown. MICROSCOPIC FEATURES: spores 20–24 × 12–14 μm, elliptic, verrucose, purplish brown; asci 200–225 × 23–27 μm, clavate; paraphyses slender, slightly enlarged at the tips. MACROSCOPIC FEATURES:

Bur n Site Shield Cup

scattered, in groups, or in dense overlapping clusters on burned ground; spring, summer, and fall, year-round in California; widely distributed in North America; occasional to common. COMMENTS: Ascobolus atrofuscus (W. Phillips) Plowright is a synonym. Ascobolus geophilus Seaver (not illustrated), a burnt-ground species found from New York to Colorado, is very similar but the warts on its spores are very delicate. Other similar species of Ascobolus typically grow on dung. OCCURRENCE:

Ascobolus carbonarius

A scobolus

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Ascobolus stercorarius (Bulliard) J. Schröter MACROSCOPIC FEATURES:

fruitbody 0.5–5 mm wide, cup- to disc-shaped to more or less pulvinate, lacking a stalk; upper surface initially smooth, then rough from projecting asci, yellow to greenyellow, darkening to olive-brown to black-brown as the violet-brown spores mature; margin slightly projecting, fi nely scurfy, paler than upper surface; underside furfuraceous like the margin, often almost smooth in age, paler than upper surface, slightly whitish. MICROSCOPIC FEATURES: spores 17–28 × 9.5–14 μm, elliptic, surrounded by a gelatinous sheath, biseriate or irregularly disposed, initially hyaline then violet and fi nally violet-brown, with 7– 9 anastomosing light-colored, longitudinal, subparallel ribs that dissolve in KOH; asci 200–220 × 20–22 μm, 8-spored, amyloid overall; paraphyses slender, clavate, up to 8 μm long, septate, and occasionally some elliptic cells of the paraphyses are swollen up to 21 μm, sometimes branched. OCCURRENCE: solitary to crowded on animal dung of both predators and herbivores (often cow dung), also on rotten cabbage and other refuse; yearround; widespread. COMMENTS: Ascobolus furfuraceus Persoon: Fries is listed in Index fungorum and in MycoBank as a synonym of Ascobolus stercorarius. Some authors consider Ascobolus furfuraceus to be a distinct species with spores 22–28 × 10–14 μm, while Ascobolus stercorarius has smaller spores (17–22 × 9.5–12.5 μm). Ascobolus michaudii Boudier (not illustrated) has

a lemon-yellow upper surface versus a yellowish brown upper surface, and nearly indistinguishable spores (17–22 × 9.5–12 μm). Ascobolus crenulatus P. Karsten (not illustrated) is characterized by a greenish yellow upper surface, a distinctly scalloped margin, and smaller spores (14–16 × 7– 9 μm). Ascobolus lignatilis Albertini and Schweinitz (not illustrated) is distinguished by having a stalk and smaller spores (13–21 × 8–10 μm). A check of Index fungorum lists 257 entries under the genus Ascobolus. While many species have been moved to new genera over the years (in excess of 20 genera), a vast array of species remains in Ascobolus. Thecotheus species also have amyloid asci that protrude when mature, and the spores have a uniform gelatinous coating. Thecotheus species (about 30 are known globally) are characterized by having hyaline spores with or without apiculi, and 8 to 32 spores per ascus. Iodophanus species also have asci that are amyloid overall, but the hyaline wrinkled spores are not covered in a gelatinous sheath. Lasiobolus species have upright, stiff, nonseptate setae on the underside, inamyloid asci, and normally hyaline spores that are covered with a gelatinous sheath. Lasiobolus cuniculi Velenovský has elliptic spores measuring 16–26 × 10–15 μm. Cheilymenia species have hyaline to brown septate setae on the underside and inamyloid asci. The smooth spores may have a loose, irregularly verrucose to echinulate episporium.

Pezizom ycetes

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Ascobolus stercorarius

A scobolus

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Byssonectria terrestris (Albertini and Schweinitz: Fries) Pfister fruitbody up to 5 mm high and wide, at fi rst cylindric to cushion-shaped or nearly round to top-shaped, pale orange with a whitish scurfy coating overall, soon opening and forming a bright orange to orange-yellow stalkless cup with an uneven and scurfy margin that sits on a white subiculum measuring less than 2 mm thick. MICROSCOPIC FEATURES: spores 18–26 × 8–10 μm, uniseriate, fusiform, smooth, hyaline, bigutt ulate; asci cylindric to clavate, 200–225 × 10–12 μm, 8-spored, inamyloid; paraphyses slender, enlarged at the tips. OCCURRENCE: in dense clusters on an extensive white mat of mycelium covering leaf and needle litter, mosses, twigs, and soil where animals have urinated and where dung may be present; spring and early summer; widespread, mainly northern distribution; uncommon. COMMENTS: Humarina aggregata (Berkeley and Broome) Seaver, Peziza aggregata Berkeley and Broome, Peziza fusispora var. aggregata Berkeley and Broome, and Thelebolus terrestris Albertini and Schweinitz are synonyms. Byssonectria fusisMACROSCOPIC FEATURES:

pora (Berkeley) Rogerson and Korf = Inermisia fusispora (Berkeley) Rifai is similar and widely distributed in North America, but its fruitbody tends to be slightly larger, up to 6 mm wide, has a scanty mycelial mat, grows on burned areas usually containing charred wood, and has spores that measure 24–29 × 7–11 μm. Byssonectria seaveri Pfi ster (not illustrated) is similar and occurs on burned soil in eastern North America. Byssonectria seaveri has a scanty mycelial mat and much larger spores that measure 29–36 × 8–10 μm. Byssonectria cartilaginea (Kanouse and A. H. Smith) Pfister = Pseudocollema cartilagineum Kanouse and A. H. Smith, found under or near melting and retreating snow in the western mountains of Canada and the United States, is distinguished by a 2–5 mm thick cartilaginous subiculum binding rodent dung. Spores (20–24 × 8–10 μm) are similar to Byssonectria terrestris but Byssonectria cartilaginea has wider asci (230–250 × 15–17 μm), paraphyses slender, slightly clavate. In contrast, Byssonectria terrestris is only rarely associated with dung, and the subiculum is less than 2 mm thick.

Byssonectria terrestris

Pezizom ycetes

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Caloscypha fulgens (Persoon) Boudier Anamorph: Geniculodendron pyriforme G. A. Salt Blue-Sta ining Cup, Snow Ba nk Or a nge Peel Fu ngus

fruitbody 1–5 cm wide, irregularly cup-shaped and frequently split on one side, stalkless; inner surface smooth, bright yellow-orange to orange; outer surface yellow, often stained dark bluish green, especially toward the margin. MICROSCOPIC FEATURES: spores 5–8 μm, round, smooth, uniseriate, hyaline; asci up to 100 μm long and 10–12 μm wide, cylindric, tapered below into a long stem-like base; paraphyses slender, fi lled with orange granules. MACROSCOPIC FEATURES:

scattered or in groups on damp soil or on debris under conifers or in mixed woods; spring and summer; widely distributed in North America; common. COMMENTS: Peziza fulgens Persoon and Pseudoplectania fulgens (Persoon) Fuckel are synonyms. A rare form of this fungus does not have orange pigmentation and is eggshell white, revealing the true robin’s-egg-blue color of the stain that normally appears greenish due to the combination of blue and orange (Rogers and Bonman 1978). OCCURRENCE:

Caloscypha fulgens

Caloscypha

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Chaetothiersia vernalis Perry and Pfister fruitbody 0.5–3 cm wide, deeply cup-shaped when young, expanding to discshaped and becoming somewhat convoluted in age, stalkless with a narrowed base; upper surface smooth, pale gray to nearly white; margin distinct, covered with erect dark brown hairs, often in fascicles; underside densely to sparsely covered with pale to dark brown, appressed to projecting hairs. MICROSCOPIC FEATURES: spores 16.5–19 × 10– 12 μm, elliptic, smooth, egutt ulate, hyaline; asci 300–380 × 8–16 μm, 8-spored, inamyloid, arising from a pleurorhynchous base; paraphyses fi liform, straight, multiseptate, exceeding asci by approximately 10 μm. OCCURRENCE: gregarious to cespitose on decaying wood and bark of conifers, and on woody debris in soil; spring and summer, associated with snowmelt; known only from the high northern Sierra Nevada Mountains of California. COMMENTS: Species of Trichophaea, Trichophaeopsis, Tricharina, Geopora, Humaria, and Wilcoxina MACROSCOPIC FEATURES:

all superficially resemble Chaetothiersia vernalis. Trichophaea woolhopeia (Cooke and W. Phillips) Boudier (not illustrated) and Trichophaea abundans (P. Karsten) Boudier are distinguished by gutt ulate spores, as are species of the usually hypogeous genus Geopora. Wilcoxina species are more difficult to distinguish because they have eguttulate spores and hairs like Chaetothiersia vernalis, but their spores are generally smaller (up to 16 × 9.5 μm) and their apothecia are often orange to ochraceous. Exceptions are Wilcoxina rehmii Chin S. Yang and Korf and Wilcoxina sequoia (W. Phillips) T. Schumacher (not illustrated), which are colored much like Chaetothiersia vernalis. Wilcoxina rehmii, found in northern parts of North America and Europe on sandy soils, has spores that are 12–14 × 7– 9 μm, smooth, egutt ulate, and arranged in the upper part of the ascus. Wilcoxina sequoia is found in California on the bark and foliage of Sequoia gigantea. We were unable to fi nd information on spore size for this species.

Chaetothiersia vernalis

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Cheilymenia fimicola (De Notaris and Baglietto) Dennis Ey el ash Du ng Cup

fruitbody 3–10 mm wide, rounded and closed at fi rst, soon opening and becoming cup-shaped, fi nally shallowly cup-shaped, stalkless; inner surface bright yellow to orangeyellow or pale orange; outer surface and margin pale orange-yellow, with a sparse coating of long, pale brown, stiff hairs. MICROSCOPIC FEATURES: spores 17–22 × 9–12 μm, elliptic, smooth, uniseriate, hyaline; asci up to 200 μm long and 12–20 μm wide, cylindric or subcylindric; paraphyses fi liform, with clavate tips. OCCURRENCE: solitary or in groups on dung, manure, or compost; spring, summer, and fall, also winter and spring in California; widely distributed in North America; common. COMMENTS: Cheilymenia coprinaria (Cooke) Boudier, Patella coprinaria (Cooke) Seaver, and Peziza coprinaria Cooke are all synonyms. Cheilymenia stercorea (Persoon) Boudier is widely distributed MACROSCOPIC FEATURES:

in North America, has very small (0.5–3 mm wide) reddish orange shallow cups fading to yellowish or brownish, margin a membranous collar with long bristle-like hairs that are pale brown; underside with bristle-like rooting hairs intermingled with stellate brown hairs. Cheilymenia stercorea spores measure 12–19 × 6–11 μm. Cheilymenia fimicola has dark brown rather than light brown marginal hairs, and does not have stellate hairs on the underside. Cheilymenia raripila (W. Phillips) Dennis (not illustrated) is another dung-inhabiting species. Cheilymenia raripila differs from both Cheilymenia stercorea and Cheilymenia fimicola in having hairs that are not forked at the base, yellow apothecia that are often rather pale, and ascospores measuring 22– 26 × 12–14 μm. Cheilymenia theleboloides (p. 137) is very similar and grows on dung mixed with straw, or on the decaying remains of plants. Its fruitbody is pale orange-yellow and has hyaline to pale yel-

Cheilymenia fimicola

Ch e i ly m e n i a

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low hairs. Pseudombrophila species are sometimes confused with Cheilymenia species. Pseudombrophila species can be whitish, yellowish gray or pale ochraceous to darker brownish, with purplish tinges. They sometimes have a prominent membranous margin and an outer surface with blunt pale brownish hairs that is concolorous with or darker than the interior. The spores

are smooth, elliptic, more or less hyaline, egutt ulate, and sometimes contain de Bary bubbles. Two particularly att ractive and widespread dung-loving species, measuring 1–10 mm wide, are Pseudombrophila porcina (Svrček and Kubička) Brummelen and Pseudombrophila merdaria (p. 234). See also comments under Scutellinia crucipila (p. 251) and Ascobolus stercorarius (p. 130).

Pezizom ycetes

136

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Cheilymenia theleboloides (Albertini and Schweinitz) Boudier fruitbody 2–10 mm broad, saucer- to disc-shaped, broadly att ached; upper surface orange fading to golden yellow, then olive-yellow, becoming reddish brown in drying; margin with sparse and indistinct hyaline hairs, somewhat darker than the upper surface; underside paler than the upper surface, with comparatively few inconspicuous, scattered, hyaline to yellowish brown pointed hairs, the longest ones near the margin. MICROSCOPIC FEATURES: spores 12–20(–22) × (7–) 10(–11) μm, elliptic, smooth, egutt ulate, uniseriate, hyaline; epispore separates and forms an enMACROSCOPIC FEATURES:

velope when heated in lactic acid; asci 180–250 × 10–15 μm, 8-spored, cylindric, inamyloid; paraphyses slender, infrequently septate, subclavate at apices, 3–4 μm in lower part, 5–7 μm at apices, yellowish at tips; the longest hairs of the fruitbody margin are 170–800 μm long × 8–10 μm wide just above the base, the base often swollen to 15–25 μm wide, 1–3(6)-septate, straight, unbranched, narrowing slightly, usually with a blunt tip, originating from the superficial cells of the ectal excipulum, yellowbrown to nearly transparent. OCCURRENCE: gregarious to crowded on dung (typically cow or horse dung), rich soil, or decay-

Cheilymenia theleboloides

Ch e i ly m e n i a

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ing plant debris; nearly year-round, whenever temperatures are above freezing and there is adequate moisture; widely distributed; very common. COMMENTS: Synonyms include Scutellinia theleboloides (Albertini and Schweinitz) Kuntze, Lachnea ascoboloides (Bertero) Massee, and Neottiella ascoboloides (Bertero) Saccardo. Cheilymenia fimicola (p. 135) and Cheilymenia stercorea (Persoon) Boudier have numerous dark brown marginal hairs that arise from a branched “root” deeply embedded in the tissue under the upper fertile layer and are typical dung lovers found rarely on soil, while Cheilymenia theleboloides is often found on soil. Cheilymenia vitellina (Persoon) Dennis is characteristically found on soil and has much longer hairs,

reaching 1,000 × 40 μm. The disc is yellow and the hairs are yellowish, and the spherical ascospores are yellowish and measure 14–17 × 7– 9 μm. Scutellinia crucipila (p. 251) has numerous hairs that arise from a deep, unbranched “root,” and spores that are slightly warted (oil immersion) rather than smooth. Most importantly, it grows on soil, not dung. Scutellinia erinaceus (p. 252) has dense dark brown rooting hairs that are inconspicuously septate, has smooth spores, and grows on rotted wood. Tricharina gilva (p. 259) has light reddish brown hairs that are sparingly septate, thin-walled, and have a rounded point. Tricharina gilva grows on burnt ground, and, like all but Scutellinia crucipila, has smooth spores.

Pezizom ycetes

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Chorioactis geaster (Peck) Kuiper ex Eckblad fruitbody up to 10 cm long × 5 cm wide before opening to a fi nal width of 10–12 cm, club-shaped, nearly closed when young, the upper half splitt ing into 3– 6 rays at maturity, rays merging into a below-ground stalk; outer surface of the entire fruitbody covered with dense chocolate-brown tomentum; interior hollow, lined with a yellowish white to bright butterscotchyellow fertile layer that becomes leather-colored in age; stalk 1–5 cm long × 1–2 cm wide; flesh pure white, soft , and leathery when fresh, approximately 3.5 mm thick. MICROSCOPIC FEATURES: spores 50– 70 × 10–16 μm, oblong-fusiform, distinctly flattened on one side, uniseriate, slightly overlapping, with 3–5 prominent oil drops, hyaline, located in the upper twothirds of the ascus; asci 600–800 × 14–23 μm, cylindric to subcylindric, with an apical pore; paraphyses uniformly less than or equal to 2 μm in diameter, branched, septate. OCCURRENCE: solitary or in groups in moderately moist hardwood forests on roots and below-ground MACROSCOPIC FEATURES:

Dev il’s Ciga r

portions of old disintegrating stumps of the smallleaved elm; development begins in late spring, but is not noticeable aboveground until rainy periods occur late fall to early the next spring; known only from Texas and Kyushu, Japan; rare. COMMENTS: Urnula geaster (Peck) is a synonym (Heald and Wolf 1910; Rudy and Keller 1996; Pfister and Kurogi 2004). The name “devil’s cigar” comes from the fact that when the rays open, it gives off puffs of spores that appear similar to puffs of smoke. When giving off spores, an audible hiss can be heard from several feet away. The rays generally open flat to the ground and partially reclose after spore release in the manner of a Geastrum and thus the scientific name. Recent DNA analysis shows that even though the two populations of this species—one in Japan and one in Texas—diverged more than 19 million years ago, they remain quite closely related (Peterson et al. 2004).

Chorioactis geaster

Chor ioactis

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Cookeina tricholoma (Montagne) Kuntze fruitbody 1–1.5 cm wide × 1 cm high, cup-shaped, with a slightly incurved margin, nearly sessile or with a long stalk; inside of cup bright red, almost scarlet; outside of cup clothed in long hairs, most numerous near the margin, forming an incurved fringe; hairs 0.2– 0.3 cm long, 0.1– 0.2 mm at base and tapering upwards, whitish or pale brown; stalk virtually absent or up to 2–3 cm tall × 0.2 cm thick. MICROSCOPIC FEATURES: spores 27–33 × 12–14 μm, elliptic to fusoid, containing 1–2 large oil drops, usually marked with delicate, longitudinal striations, more or less hyaline; asci up to 350–375 × 20 μm, cylindrical, abruptly narrowed below to a short appendage-like base, 8-spored; paraphyses filiform, slightly enlarged at tips; hairs a dense fascicle of mycelial threads. OCCURRENCE: on old wood and bark; year-round; Central America, Mexico, and parts of very southern United States, plus South America, Australia, and the Philippines. MACROSCOPIC FEATURES:

All members of the genus are mainly tropical, brightly colored, usually clothed in external hairs, having 4–8 more or less hyaline, elliptic to subfusoid, striate spores (Pfister 1978a; Iturriaga and Pfi ster 2006). Cookeina sulcipes (Berkeley) Kuntze, found on old wood and bark year-round in tropical to subtropical regions of South America to the southern United States, is distinguished by very short yellowish hairs on the lower surface and marked with several concentric rings near the margin. Cookeina sulcipes is cup-shaped, 1–2 cm wide and up to 1 cm deep, upper surface deep orange to scarlet, fading when dry, stalk often absent but can be up to 3 cm long × 0.2 cm diameter and is microscopically virtually indistinguishable from Cookeina tricholoma. Cookeina tetraspora Seaver (not illustrated) is found on the leaves of some palms and is gregarious to the point of becoming confluent in age. It has a whitish, slightly tomentose fruitbody and, as the name implies, has four spores per ascus.

COMMENTS:

Cookeina tricholoma

Pezizom ycetes

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Disciotis venosa (Persoon: Fries) Boudier fruitbody 4–21 cm wide, cup-shaped when young, becoming shallowly cupshaped as it matures, with a short stalk; inner surface with conspicuous, often radially arranged vein-like folds and wrinkles, sometimes appearing reticulate, reddish brown to yellowish brown; outer surface slightly scurfy, grayish white to buff or brownish; margin wavy to irregular in age, often inrolled or turned up, sometimes splitt ing; flesh thick and britt le, sometimes reported as having a chlorine bleach odor; stalk up to 1.5 cm long and thick, typically ribbed. MICROSCOPIC FEATURES: spores 19–30 × 12–16 μm, broadly elliptic, smooth, lacking oil drops or sometimes with oil droplets on the exterior polar spore wall, pale yellow; asci 300–350 × 17–23 μm, cylindric, 8-spored; paraphyses fi liform, septate, with clavate tips and yellowish brown contents. MACROSCOPIC FEATURES:

V eined Cup

scattered or in groups on the ground under conifers and hardwoods; spring and early summer; widely distributed in North America; occasional. EDIBILIT Y: edible. COMMENTS: Peziza venosa Persoon is a synonym. Gyromitra ancilis (p. 144) is similar but the surface of its cup is folded, wrinkled or occasionally smooth and not radially veined. The underside usually has minute tuft s of hairs in small dot-like clumps. Its spores have prominent appendages and oil drops. Whether or not any North American material is the same as the European Disciotis venosa is as yet unclear. It is clear that we have at least two different species in North America and that the genus is in need of critical study. OCCURRENCE:

Disciotis venosa

Disciotis

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Galiella rufa (Schweinitz) Nannfeldt and Korf fruitbody 1.2–3 cm high × 1–3.5 cm wide, cup-shaped; inner surface pale orange to dull orange, reddish orange, reddish brown or tan, smooth; outer surface tough, brown near the margin, blackish brown below, covered with a dense layer of matted wooly hairs; margin fi nely toothed; flesh rubbery-gelatinous. MICROSCOPIC FEATURES: spores 18–20 × 8–10 μm, elliptic with narrow ends, fi nely verrucose, uniseriate, hyaline; asci 275–300 × 12–14 μm, cylindric; paraphyses fi liform. MACROSCOPIC FEATURES:

H a iry Rubber Cup

in groups or clusters on decaying hardwood; summer and fall; widely distributed in eastern North America; fairly common. COMMENTS: Bulgaria rufa Schweinitz is a synonym. Wolfina aurantiopsis (p. 269) is similar but the inner surface of its fruitbody is pale yellow to ochraceous or orange, the outer surface lacks the dense layer of matted wooly hairs, its flesh has a corky texture, and it has larger spores (27–33 × 16–18 μm). OCCURRENCE:

Galiella rufa

Pezizom ycetes

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Geopyxis carbonaria (Albertini and Schweinitz) Saccardo fruitbody 6–20 mm wide, deeply cup-shaped with a short, slender stalk; inner surface smooth, reddish brown to orange-brown; outer surface fi nely scurfy, colored like the inner surface; margin whitish, toothed or crenate, sometimes torn; stalk 0.5–2.5 cm long × 1.5– 6 mm thick, tapered downward, pale yellow. MICROSCOPIC FEATURES: spores 11–17 × 6– 9 μm, elliptic, smooth, egutt ulate, uniseriate, hyaline; asci 180–250 × 10–11 μm, clavate, 8-spored; paraphyses slender, septate, forked toward the base, tips typically not enlarged. OCCURRENCE: in groups or clusters on burned soil or charred wood; spring, summer, and fall, also late fall and winter in California; widely distributed in North America; occasional. COMMENTS: Peziza carbonaria Albertini and Schweinitz is a synonym. Geopyxis vulcanalis (Peck) Saccardo is similar but its fruitbody is MACROSCOPIC FEATURES:

Pi x ie Cup

rounded when young, soon becomes deeply cupshaped, then flattens out in age, and its stalk is generally short, up to 0.5 cm long and 0.1– 0.3 cm wide. It has a pale orange to yellowish upper surface, a fi nely scalloped margin, a paler to whitish underside, larger spores that measure 14–22 × 8–11 μm, elliptic, ends strongly narrowed, smooth, and eguttulate. Asci up to 275 × 15–17 μm and paraphyses slightly thickened in upper part, 4 μm at tips. Geopyxis vulcanalis (Peck) Saccardo grows on needle duff, among mosses, under conifers, or sometimes on burned ground, and usually has a sulfur odor when crushed. Geopyxis carbonaria occurs only on burned soil and charred wood, and its cup has less of a tendency to expand and is reddish tan inside and yellowish tan outside, contrasted with the more yellowish tan inside and whitish outside of Geopyxis vulcanalis. The spores of Tarzetta species are bigutt ulate.

Geopyxis carbonaria

Geopy x is

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Gyromitra ancilis (Persoon) Kreisel fruitbody 2–12 cm wide × 0.5–4 cm tall, discoid to shallowly cup-shaped to saucer-shaped, with an indistinct or rudimentary stalk; margin inrolled initially, soon reflexed; upper surface usually undulate-rugose, sometimes nearly smooth, red-brown to dark brown, drying dark brown to blackish brown; lower surface nearly bald to slightly pubescent, somewhat ribbed near the stalk, creamy white or pale brown, often somewhat translucent; stalk anywhere from a barely discernible thickened base to as large as 4.5 cm long × 3 cm thick. MICROSCOPIC FEATURES: spores 25–45 × (8–)11– 16 μm, spindle-shaped, some with a pointed thickened apiculus at each end, smooth to slightly MACROSCOPIC FEATURES:

Pig’s Ea rs

roughened, with 1 large central oil drop that is sometimes flanked by 2 smaller oil drops; asci 300– 375 × 17–25 μm, cylindrical, 8-spored, inamyloid; paraphyses gradually to abruptly clavate, 5–11 μm at the tip, septate, terminal cell 60–105 μm, with coarsely granular brownish contents. OCCURRENCE: single to clustered on the ground or on rotten wood; appearing soon after snowmelt under conifers and sometimes under hardwoods; widespread in North America, Europe, and Asia; common. COMMENTS: Discina ancilis (Persoon) Saccardo, Gyromitra macrospora (Bubák) Harmaja, Gyromitra fluctuans (Nylander) Harmaja, and Gyromitra warnei (Peck) Harmaja are all synonyms. Th is

Gyromitra ancilis

Pezizom ycetes

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species is also commonly known as Discina perlata Fries. Gyromitra ancilis, Gyromitra melaleucoides (p. 152), Gyromitra leucoxantha (Bresàdola) Harmaja, and Gyromitra olympiana (Kanouse) Harmaja can be most reliably differentiated by examination of their mature spores freely deposited in a spore print (not as seen in asci), since sizes, shapes, and colors of mature specimens overlap between these four species. Gyromitra melaleucoides has small spores (11.5– 14.5 × 8– 9.5 μm) that mature quickly and are elliptical in shape without appendages at the ends. The other three species all have much larger, slowly maturing spores that cannot be distinguished by size, but are distinguished by the shape of appendages at the ends of the spores when mature. Gyromitra ancilis develops a thick pointed apicu lus at both ends of some spores, Gyromitra olympiana develops a broad, blunt thickening at both ends of some mature spores, and Gyromitra leucoxantha has a depressed thickening at both ends of some mature spores, giving them a twohumped appearance. Gyromitra olympiana, named “Olympic pig’s ears,” is found both in the western mountains and eastern North America, typically has an upper surface that is yellow-brown to brown, becoming redbrown in age, sometimes with olivaceous tints, and

dries blackish brown. Discina olympiana Kanouse and Discina apiculata McKnight are synonyms. Gyromitra mwb060911, which we have named “mini peach pig’s ears,” microscopically is strikingly similar to Gyromitra olympiana but is distinguished by its consistently small size (1.5–4.5 cm wide; stalk, if present, up to 1 cm × 1 cm) and pinkish buff coloration of the upper surface, with the lower surface off-white, drying tan, and minutely pubescent. Gyromitra mwb060911 can be found from about 2,000′ elevation to about 4,500′ elevation in the Gifford Pinchot National Forest of Southwest Washington, where it is often the most abundant disk-like Gyromitra. It has also been found near McCall, Idaho. Gyromitra leucoxantha, known as “orange pig’s ears” or “yellow pig’s ears,” is uncommon throughout North America and Europe, and is typically yellow to orange or orange-brown, drying bright red-brown or brown. Discina leucoxantha Bresàdola is a common synonym. Other synonyms are Discina larryi McKnight and Gyromitra larryi (McKnight) Harmaja. Disciotis venosa (p. 141) has a veined upper surface, small more or less distinct dark tuft s of hair on the underside, and smooth spores that are eguttulate. It sometimes has a chlorine odor.

Gy rom itr a

145

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Gyromitra brunnea Underwood Ga bled Fa lse Mor el, Eleph a nt Ea r, Rooster’s Comb

fruitbody consisting of a cap and stalk; cap 4–12 cm wide × 4– 9 cm high, irregularly folded and deeply divided into uplifted wrinkled lobes comprising an upper and lower surface, sometimes saddle-shaped, moist and lubricous; margin wavy and folded; upper surface yellow-brown to reddish brown; lower surface pale yellow-brown to whitish, slightly granular, often fusing with any part of the stalk or cap; stalk 2.5– 10 cm long, 4– 9 cm thick, stout, enlarging downward or nearly equal, chalky white to pale grayish white, typically ribbed. MICROSCOPIC FEATURES: spores 25–30 × 13–15 μm, elliptic, fi nely reticulate, with several short projections on each end that appear only on the spores of mature specimens, containing 1–3 oil drops, uniseriate, hyaline; asci cylindric to subcylindric, 8-spored; paraphyses slender, with clavate tips. OCCURRENCE: solitary to scattered on the ground in hardwood forests; spring and early summer; eastern North America; uncommon. MACROSCOPIC FEATURES:

questionable and not recommended; although eaten by some individuals, this fungus is known to cause gastric distress and severe headache. COMMENTS: Gyromitra fastigiata (Krombholz) Rehm and Helvella underwoodii Seaver are synonyms. Gyromitra brunnea is distinguished by a cap that lacks ribs and cross-ribs, is frequently lobed, and forms seams where the creamy tan undersurface can be seen. In contrast, Gyromitra caroliniana (p. 148) has ribs and cross-ribs so that it looks ridged and pitted and somewhat resembles a true morel. The cap is never lobed, so the undersurface is never exposed except very close to the stalk. Gyromitra brunnea and Gyromitra caroliniana are in subgenus Caroliniana of Gyromitra (Methven, Zelski, and Miller 2013). EDIBILIT Y:

Gyromitra brunnea

Pezizom ycetes

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Gyromitra californica (W. Phillips) Raitviir fruitbody a cap and stalk; cap 2.5–14 cm wide × 1.5–8 cm high, very thin and fragile, puffed up like a delicate umbrella, margin usually free from stalk and incurved; upper surface uneven to convoluted but not brain-like, tan to brown, olive-brown, or grayish brown; lower surface strongly fluted with sharp-edged ribs, pubescent, white or creamy; stalk 3–12 cm long × 2– 6 cm thick, equal or narrowed at base, deeply fluted with widely spaced ribs that often reach nearly to the margin of the cap, usually not internally chambered in cross section, pubescent, whitish to yellowish, typically with rosy, wine-red, or purplish tints. MICROSCOPIC FEATURES: spores 13–20 × 7–10.5 μm, elliptic, bigutt ulate, lacking apiculi, smooth, hyaline; asci 160–200 × 10–12 μm, 8-spored; paraphyses 6–8 μm at apex, clavate, brown in mass. MACROSCOPIC FEATURES:

Umbr ell a Fa lse Mor el

solitary to several on soil or rott ing logs in coniferous woods; spring to summer and sometimes fall (winter in California); a common endemic to western North America. EDIBILIT Y: suspected of causing poisoning. COMMENTS: Since this mushroom looks like a hybrid of a Gyromitra cap on a Helvella stalk, it should be no surprise that synonyms include Helvella californica W. Phillips. Pseudorhizina californica (W. Phillips) Harmaja is the accepted name as of this writing, but recent molecular work places this species right in the middle of Gyromitra, in subgenus Pseudorhizina (Methven, Zelski, and Miller 2013). Gyromitra sphaerospora (p. 154) is the other member of the subgenus Pseudorhizina. OCCURRENCE:

Gyromitra californica

Gy rom itr a

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Gyromitra caroliniana (Bosc: Fries) Fries Ca rolina Fa lse Mor el, R ed Fa lse Mor el

fruitbody consisting of a cap and stalk; cap 5–18 cm wide and high, convoluted and brain-like, typically with several prominent seam-like vertical ridges, usually not saddleshaped, with ribs and cross-ribs, appearing ridged and pitted, moist and lubricous; margin appressed against the stalk, wavy and irregular; upper surface reddish brown; lower surface never exposed except very close to the stalk, white to whitish; stalk 4– 15.5 cm long × 2.5–8 cm thick, enlarged at the base, distinctly ribbed, white, interior multichambered. MICROSCOPIC FEATURES: spores 22–35 × 10–16 μm, elliptic, reticulate, with one or more short projections at each end, uniseriate, usually containing 1 large oil drop and two smaller ones, hyaline; asci clavate or subcylindric; paraphyses fi liform, with brownish, somewhat thickened tips. OCCURRENCE: solitary or in groups on the ground in hardwood forests or mixed woods; spring and early summer; widely distributed in eastern North America, west to Colorado and the Rocky Mountains; occasional. EDIBILIT Y: questionable and not recommended; although eaten by some individuals, this mushroom is suspected to contain toxins. COMMENTS: Helvella caroliniana (Bosc) Nees is a synonym. It is also commonly known in places as

the “red morel,” a term we do not like to use since it is not a true morel. Gyromitra caroliniana can attain massive sizes; we have read of specimens weighing as much as seven pounds each. Where their ranges overlap, it can be difficult to separate Gyromitra caroliniana from Gyromitra brunnea (p. 146).

Pezizom ycetes

148

MACROSCOPIC FEATURES:

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Gyromitra esculenta (Persoon) Fries fruitbody consisting of a cap and stalk; cap 3.5–10 cm wide, 4–10 cm high, brain-like to irregularly lobed, deeply wrinkled to convoluted, moist to dry; margin undulating to contorted, often curved toward the stalk; upper surface pinkish tan to dark reddish brown or orange-brown, lubricous when fresh; lower surface pale pinkish tan to yellowish tan; interior chambered; flesh very britt le; stalk 2–7 cm long × 2–3 cm thick, enlarging downward or nearly equal, stuffed with cottony hyphae, sometimes hollow or chambered, surface smooth and waxy to slightly granular, dingy white to pinkish tan or tan, often ribbed near the base. MICROSCOPIC FEATURES: spores 16–28 × 7–13 μm, elliptic, smooth, bigutt ulate, uniseriate, hyaline; asci up to 350 × 16–20 μm, 8-spored; paraphyses cylindrical, branched, with clavate tips. MACROSCOPIC FEATURES:

Br a in Mushroom, Fa lse Mor el

solitary, scattered, or in groups on the ground under conifers, especially white pine, or in mixed woods; spring and early summer; widely distributed in North America and Europe; common. EDIBILIT Y: poisonous; this species contains hydrazine derivatives that can cause serious illness or death. COMMENTS: Many common names have been assigned to this mushroom, including Conifer False Morel, Beefsteak Morel, and Lorchel. Helvella esculenta Persoon is a synonym. A cream-colored variant is known as Gyromitra esculenta var. alba Pilát. We have found miniature versions of this species only 1 cm tall. Gyromitra esculenta, Gyromitra ambigua (P. Karsten) Harmaja, and Gyromitra infula (p. 150) are in the subgenus Gyromitra. OCCURRENCE:

Gyromitra esculenta

Gy rom itr a

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Gyromitra infula (Schaeffer: Fries) Quélet

Sa ddle-Sh a ped Fa lse Mor el

fruitbody consisting of a cap and stalk; cap 2.5–10 cm wide × 2–10 cm high, usually saddle-shaped or sometimes trilobate, margin incurved; upper surface wrinkled to convoluted or sometimes nearly smooth, moist when fresh, reddish brown to dark brown or sometimes yellow-brown, lacking distinct violet to lavender tints; lower surface paler, minutely velvety; interior hollow or chambered, flesh britt le; stalk 2– 6 cm long × 2–2.5 cm thick, dry, hollow, fi nely granular, whitish to pinkish buff. MICROSCOPIC FEATURES: spores 18–23 × 7–10 μm, elliptic, smooth, with 2 large oil drops when mounted in water, uniseriate, nonapiculate, hyaline; asci 250–330 × 14–15 μm, 8-spored; paraphyses cylindrical, septate, forked, and enlarged apically. OCCURRENCE: solitary, scattered, or in groups on decaying wood or humus; summer and fall, also winter and spring in coastal California; widely distributed in North America; occasional to locally common. EDIBILIT Y: poisonous. COMMENTS: Th is false morel is unusual for the genus Gyromitra in that it typically fruits in the summer and fall rather than spring, except for coastal California where it fruits in the spring. We have found versions of this species less than 1 cm tall in the spring. Helvella infula Schaeffer is a synonym. Gyromitra ambigua (P. Karsten) Harmaja is very similar and also fruits in the summer and fall, but its cap and stalk have distinct violet to lavender tints, with the cap a dark red-brown, nearly chestnut colored, and its spores are larger (21–30 × 7– 12 μm) and indistinctly apiculate (Harmaja 1969a). Some authors consider Gyromitra ambigua and Gyromitra infula to be synonyms, but the combination of fruiting body color and spore characteristics appear to be consistent diagnostic features (Abbott and Currah 1997). Sphaeronaemella helvellae

(P. Karsten) P. Karsten grows on the living fruitbodies of Gyromitra infula and Gyromitra ambigua. Sphaeronaemella helvellae gives Gyromitra fruitbodies a velvety, withered appearance but Gyromitra ascospores are always present, indicating that the infection does not occur until host maturity. Sphaeronaemella helvellae fruitbodies are superficial to semi-immersed in the spore-bearing area of the host. They are composed of perithecia that measure 0.09– 0.25 mm in diameter, are nearly spherical to ovoid, bald, smooth, and have a long neck through which the spores emerge. They are colored bright yellow-orange and are densely gregarious. Their spores are 8–10.5 × 3–4.5 μm, unilaterally flattened-elliptic in side view, elliptic in face view, smooth, hyaline, and yellowish in mass.

Pezizom ycetes

150

MACROSCOPIC FEATURES:

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Gyromitra korfii (Raitviir) Harmaja fruitbody consisting of a cap and stalk; cap 5–14(–25) cm wide × 4–15 cm high, typically brain-like, wrinkled and deeply folded or sometimes nearly smooth, moist and lubricous; margin undulating to contorted, usually free from the stalk or sometimes fused with it; upper surface yellow-brown to reddish brown; lower surface somewhat granular, pale yellow-brown to whitish; stalk 4–7 cm long × 4–7.5 cm thick, stout, enlarging downward or nearly equal, chambered and usually stuffed with cottony white hyphae; stalk surface whitish, smooth to slightly granular, with thick rounded ribs extending from the apex to the base; flesh whitish, britt le. MICROSCOPIC FEATURES: spores 24–36 × 10–15 μm, broadly elliptic to fusiform, smooth at fi rst and becoming fi nely roughened to somewhat reticulate, with blunt appendages on each end that appear only on the spores of mature specimens, containing 1 large central oil drop and 1–2 smaller ones at each end, hyaline; asci 280–300 × 18–25 μm, MACROSCOPIC FEATURES:

Bu ll-Nose Fa lse Mor el

8-spored; paraphyses cylindrical, branched, septate, with clavate tips. OCCURRENCE: solitary, scattered, or in groups on the ground in hardwoods, coniferous forests or mixed woods; spring; throughout eastern North America and the Midwest, although probably not found much south of Tennessee; occasional to locally common. EDIBILIT Y: poisonous when raw, although eaten by some people when thoroughly cooked; not recommended. COMMENTS: Some authors consider Gyromitra gigas (Krombholz) Cooke, named from European material, to be the appropriate name for both the eastern species we recognize as Gyromitra korfii and the western species we recognize as Gyromitra montana (p. 153). Gyromitra korfii and Gyromitra montana are closely related to species formerly placed in the genus Discina and are in subgenus Discina of Gyromitra (Methven, Zelski, and Miller 2013).

Gyromitra korfii

Gy rom itr a

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Gyromitra melaleucoides (Seaver) Pfister fruitbody 1–11 cm wide × 1–5 cm tall, saucer- to cup-shaped or lobed, with the margin typically turned down; upper surface smooth and undulate, pale gray-brown to dark brown, drying dark brown to dark red-brown; underside smooth to pubescent, cream to pale yellow-brown, turning very white on drying; stalk indistinct or up to 1–5 cm tall × 0.5–3 cm thick, distinctly flared at the apex, often enlarged at the base, internally infolded and fluted, with hollow chambers, concolorous with the underside. MICROSCOPIC FEATURES: spores 11.5–14.5 × 8– 9.5 μm, elliptic to broadly elliptic, rough to warted, nonapiculate, normally bigutt ulate but sometimes having 1 oil drop; asci 200–240 × 11–14.5 μm, 8-spored; paraphyses 5.5– 9 μm at apex, clavate, brown in mass. OCCURRENCE: solitary to numerous, sometimes subcespitose on soil or rotted wood in both coniferous and mixed woods; soon after snowmelt; Rocky Mountains westward in both montane and coastal sites; common. EDIBILIT Y: not recommended, defi nitely poisonous raw or if not thoroughly cooked. MACROSCOPIC FEATURES:

COMMENTS: Gyromitra recurva (Snyder) Harmaja is a synonym (Pfi ster 1980). Th is species has a distinctive spore morphology that quickly sets it apart from similar species such as Gyromitra ancilis (p. 144), Gyromitra leucoxantha (Bresàdola) Harmaja, and Gyromitra olympiana (Kanouse) Harmaja. The spores of Gyromitra melaleucoides are far shorter (average 12–13 μm), warted, lack apiculi, and spore prints are easy to obtain. G. melaleucoides typically has a gray cast, while the other similarly shaped species are yellow-brown, pinkish buff, or red-brown. However, the colors do overlap and are highly variable within each species mentioned. The most reliable macroscopic feature is that the underside of G. melaleucoides quickly turns white as it starts to dry out. The other species tend to retain more color on the underside as they start to dry out. One location is known in Washington State (near Salmon la Sac) where there is an unnamed cream white variety that otherwise macroscopically and microscopically matches G. melaleucoides. Gyromitra melaleucoides is the only member of subgenus Melaleucoides (Methven, Zelski, and Miller 2013).

Gyromitra melaleucoides

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Gyromitra montana Harmaja

Snow Mushroom, Wa lnut

fruitbody a cap and stalk, 3–20 cm broad × 3–15 cm high; cap upper surface brain-like with compact folds and wrinkles that become smoother in old age, yellowish brown to butterscotch-brown or reddish brown in age, edges blackening; underside usually whitish; stalk massive, often nearly as wide as the cap, often completely covered by the cap margin, convoluted to folded with several anastomosing interior channels, white. MICROSCOPIC FEATURES: spores (21.5–)24–36(–38) × (9–)10–16 μm, elliptic, some with a broad apiculus at each end, smooth to fi nely roughened with 1 large central oil drop and smaller oil drops at the ends; asci 350–400 × 18–24 μm; paraphyses cylindric-capitate, 4–12 μm at widest part, golden granules in the upper cells. MACROSCOPIC FEATURES:

solitary to gregarious under both conifers and hardwoods; near melting snow banks or shortly after snowmelt; mountainous areas of western North America; common. EDIBILIT Y: not recommended, but commonly eaten, poisonous when raw or not thoroughly cooked. COMMENTS: Some authors have considered Gyromitra montana to be a synonym of Gyromitra gigas (Krombholz) Quélet, a European species. The two are morphologically very similar. Gyromitra montana is a part of the unique snow bank flora of western North America but has been shown to be genetically distinct. See also Gyromitra korfii (p. 151), an eastern North American species. OCCURRENCE:

Gyromitra montana

Gy rom itr a

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Gyromitra sphaerospora (Peck) Saccardo Rou nd-Spor ed Gy romitr a, Rou nd Spor ed Fa lse Mor el

fruitbody up to 15 cm high, consisting of a cap and stalk; cap 5–12 cm wide, 3.5–8 cm high, saddle-shaped or irregular, wrinkled to convoluted, dry; margin undulating to contorted, strongly incurved when young and typically remaining so at maturity; upper surface yellow-brown to grayish brown; lower surface slightly tomentose, whitish; flesh thin, britt le; stalk 3.5– 9 cm long, 2–5 cm wide, stout, variable, typically enlarged toward the base and apex, sometimes nearly equal; surface purplish red to rosy pink with whitish areas when young, becoming pinkish brown to pale brownish yellow in age, deeply pitted and ribbed, smooth, typically with a fi ne, white basal tomentum; ribs markedly extending onto the lower surface of the cap.

MACROSCOPIC FEATURES:

MICROSCOPIC FEATURES: spores 9–12 μm, globose, smooth, uniseriate, hyaline; asci up to 150 × 10– 12 μm, cylindric to subcylindric; paraphyses cylindric, clavate to abruptly enlarged at the tips. OCCURRENCE: solitary, scattered, or in groups on decaying logs and stumps in hardwoods or mixed woods or on sawdust piles; spring and early summer; east of the Rocky Mountains in Canada, New York, Vermont, New Hampshire, Michigan, and Montana; rare. EDIBILIT Y: unknown. COMMENTS: Pseudorhizina sphaerospora (Peck) Pouzar and Helvella sphaerospora Peck are synonyms. See comments under Gyromitra californica (p. 147).

Gyromitra sphaerospora

Pezizom ycetes

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Helvella acetabulum (Linnaeus) Quélet fruitbody 2–7 cm wide, cup-shaped; upper surface pale to dark brown; lower surface pale to dark brown near the margin, becoming whitish near the base; stalk 1–7 cm long × 1–2.5 cm thick, conspicuously ribbed, white; ribs rounded, branched upward, extending over the outer surface. MICROSCOPIC FEATURES: spores 16–22 × 11–14 μm, broadly elliptic, smooth, with 1 oil drop, uniseriate, hyaline; asci 270–400 × 15–20 μm, cylindric, 8-spored; paraphyses septate and branched at the bases, with clavate tips. OCCURRENCE: scattered or in groups on the ground in woodlands, usually under hardwood trees; spring and early summer, also winter and early spring in the Southwest; widely distributed in North America, Europe, and Asia; occasional. EDIBILIT Y: unknown. COMMENTS: Acetabula sulcata (Persoon) Fuckel, Peziza sulcata Persoon, and Paxina acetabulum (Linnaeus) Kuntze are synonyms. Helvella costifera Nannfeldt, Helvella griseoalba N. S. Weber (not illustrated) and Helvella hyperborea Harmaja (not illustrated) are all similar to Helvella acetabuMACROSCOPIC FEATURES:

R ibbed-Sta lk ed Cup, Elfin Cup

lum. Both Helvella griseoalba and Helvella costifera have a pale to dark gray apothecium, and Sean Abbott (personal communication) considers them to be synonymous. Compared to Helvella acetabulum, Helvella costifera is less ribbed, with bluntedged ribs, a nonlacunose stalk, and a more pubescent apothecium. Mushrooms with intermediate features raise the possibility that Helvella costifera, Helvella griseoalba, and Helvella acetabulum are all synonyms (Abbott and Currah 1997). Helvella hyperborea Harmaja (not illustrated), a rare species from northern areas in northeastern North America, northern Europe, and one collection from Alaska, has a medium brown, strongly ribbed cup with dark pubescence (Harmaja 1981). Helvella robusta S. P. Abbott (not illustrated), known only from Alberta, Canada, is similar to all of these species but is larger, has a robust stalk, an autumnal fruiting pattern, and a unique habitat in dry, hilly grassland with poplars and shrubs (Abbott and Currah 1988). Helvella solitaria (p. 169) is similar in color but has a relatively narrower stalk with less prominent ribbing, less ribbing on the underside of the cap, and a generally smaller size.

Helvella acetabulum

H e lv e l l a

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Helvella albella Quélet fruitbody consisting of a cap and stalk; cap 1–2.5 cm wide, weakly to moderately saddle-shaped when young, becoming strongly saddle-shaped to trilobate at maturity; margins curving over the upper surface when young, expanding and flaring in age, entire or lacerate; upper surface gray-brown to brown, smooth to slightly wrinkled; lower surface creamy white to white, pubescent when young, becoming nearly glabrous in age; stalk 2–5 cm long × 2– 6 mm thick, rounded, somewhat enlarged downward, glabrous to fi nely pruinose; whitish. MICROSCOPIC FEATURES: spores 20–24 × 12–15 μm, oblong to elliptic, with 1 large central oil droplet and up to 4 small droplets at each end, smooth, uniseriate, hyaline; asci 280–330 × 14.4–20.8 μm, cylindric to subcylindric, 8-spored; paraphyses cylindric, tips clavate to abruptly clavate or variably enlarged, contents fi nely granular, pale brown in mass. MACROSCOPIC FEATURES:

scattered or in groups on the ground in woodlands; summer, fall, and early winter; widely distributed in North America; occasional to locally common. COMMENTS: Leptopodia albella (Quélet) Boudier is a synonym. Helvella ephippium Léveillé (= Helvella murina [Boudier] Saccardo and Traverso), found in eastern North America and boreal Canada, is brown to dark brown, with a paler underside due to pubescent to villose hairs and spores measuring 17–21 × 10–13 μm. Other similar species are Helvella pezizoides (p. 168) and Helvella atra (p. 157), which have black rather than brown upper surfaces, and Helvella latispora (p. 164), which is not villose on the underside of the cap. Helvella elastica (p. 162) is very similar but its cap is typically paler, margin never inrolled, and its lower surface is smooth when young. OCCURRENCE:

Helvella albella

Pezizom ycetes

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Helvella atra J. König

Da r k Elfin Sa ddle, Black Helv ella

fruitbody consisting of a cap and stalk; cap 0.3–3 cm wide × 0.3–2 cm high, saddle-shaped to irregularly lobed, smooth to undulate-rugose, typically laterally compressed; margin appressed, usually free from stalk, initially incurved, soon reflexed; upper surface black to blackish gray-brown; underside bald to somewhat pubescent, pale to medium gray-brown; stalk 0.5– 5 cm tall × 0.1– 0.8 cm thick, round or shallowly fluted at base, solid, sometimes hollow at base, pubescent, gray-brown to blackish except for yellowish gray to white at base; flesh thin, britt le. MICROSCOPIC FEATURES: spores 15–21 × 10–13 μm, broadly elliptic, smooth, unigutt ulate, uniseriate, hyaline; asci 190–250 × 13–17 μm, 8-spored; paraphyses clavate, 5– 9 μm at apex, terminal cell 11–20 μm long, contents fi nely granular, dark brown in mass. OCCURRENCE: gregarious to scattered and numerous on the ground or on rotten wood in deciduous or mixed woods; summer and fall; western, eastern, and southern North America, as well as Europe and Asia; common. MACROSCOPIC FEATURES:

Helvella atra Oeder, Helvella atra J. König, and Helvella nigricans var. atra (Afzelius) Persoon are all listed by MycoBank as currently used legitimate names, and as far as we can determine all refer to the same fungus. Abbott and Currah (1997) give Helvella atra Holmskjøld: Fries as the preferred name. Helvella subglabra N. S. Weber (not illustrated) was separated based partly on a fuzzier, grayish fruitbody but Abbott and Currah (1997) agree with Häff ner (1987) that the differences are insufficient to separate the two. Helvella pezizoides (p. 168) is very similar, and although Häff ner (1987) considers the two to be synonymous, Abbott and Currah (1997) tentatively retain Helvella pezizoides as a distinct species due to the following characteristics: it occasionally becomes irregularly cup- to disc-shaped, it has a very involute cap margin, the cap underside is brown and densely pubescent to villose, and the stalk is concolorous with the underside. Helvella albella (p. 156) is similar, but the cap is brownish, the stalk is white, and the spores are longer and wider. COMMENTS:

Helvella atra

H e lv e l l a

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Helvella compressa (Snyder) N. S. Weber MACROSCOPIC FEATURES: fruitbody consisting of

a cap and stalk; cap 0.5–5 cm wide × 0.5–3 cm tall, saddle-shaped to irregularly lobed, margin strongly inrolled initially and often flaring when mature but still remaining somewhat inrolled, free from stalk; upper surface smooth, brown, grayish brown or dark brown, drying dark brown; underside minutely hairy, whitish; stalk 0.5–10 cm × 0.1–1.5 cm, equal or widening at base, round to somewhat flattened or pitted near base, solid except in large specimens, pubescent to villose, especially near apex, white to pale cream; flesh thin, rather britt le. MICROSCOPIC FEATURES: spores 19.5–24 × 11.5– 15 μm, broadly elliptic, with 1 large central oil drop; asci 280–349 × 15–22 μm, 8-spored; paraphyses 4– 9 μm at apex, clavate, contents fi nely granular, pale brown to brown in mass.

Compr essed Elfin Sa ddle

solitary to subcespitose on soil, litter, or burnt woody debris in mixed woods; generally only found in spring; known only west of the Rocky Mountains; regionally fairly common. COMMENTS: Helvella latispora (p. 164) is a similar species that is common in eastern North America and Europe, but rare in the West. Helvella albella (p. 156), a widely distributed species, is colored very much like Helvella compressa, but is glabrous to fi nely pubescent to pubescent on the underside of cap and on the upper stalk, as opposed to pubescent to villose like Helvella compressa. Helvella elastica (p. 162) has bent-down margins (rather than initially inrolled upward), a hollow stalk, and a bald underside. OCCURRENCE:

Helvella compressa

Pezizom ycetes

Beug-final.indb 158

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Helvella corium (O. Weberbauer) Massee fruitbody consisting of a cap and stalk; cap 0.3–4 cm wide × 0.2–1.2 cm tall, deeply to shallowly cup-shaped, margin initially inrolled, expanding at maturity; upper surface black; underside sometimes with ribs on the basal portion, black or blackish brown with a whitish marginal pubescence; stalk 0.2–4 cm × 0.05– 0.8 cm, equal or enlarged at base, round or slightly fluted to occasionally strongly ribbed, solid, whitish at base, dark brown to black on upper stalk. MICROSCOPIC FEATURES: spores 16–21 × 9–15 μm, broadly elliptic, smooth, unigutt ulate, hyaline; asci 225–250 × 12–17 μm, 8-spored; paraphyses 4–8 μm at apex, clavate, contents granular, brown in mass. OCCURRENCE: solitary to numerous, usually on bare soil, often near willows, in mixed woods, and in tundra; spring through summer; throughout MACROSCOPIC FEATURES:

Bl ack Scur f y Fa iry Cup

northern North America, Europe, and Asia; widespread but uncommon. COMMENTS: Helvella arctica Nannfeldt is a synonym. Helvella verruculosa (Saccardo) Harmaja (not illustrated) and Helvella solitaria (p. 169) are morphologically somewhat similar. Abbott and Currah (1997) postulate an affi nity with the subgenus Leucomelaenae. Dark specimens of the arctic and alpine species Helvella verruculosa are characterized by a strongly ribbed stalk with ribs extending to the undersurface of the cap. Other black Helvella species, such as Helvella atra (p. 157), Helvella pezizoides (p. 168), Helvella vespertina (p. 170), and Helvella dryophila Vellinga and N. H. Nguyen (see H. vespertina) are all characterized by having a saddle-shaped to irregularly lobed cap.

Helvella corium

H e lv e l l a

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Helvella crispa (Scopoli) Fries Fluted W hite Helv ella, W hite Elfin Sa ddle

fruitbody consisting of a cap and stalk; cap 2– 6 cm wide × 1–4cm high, saddle-shaped to irregularly lobed, dry; margin rolled inward when young then gradually unrolling and expanding at maturity, entire or somewhat lacerated; upper surface pale cream to pale buff, smooth to somewhat wrinkled; lower surface pale cream to buff, with tiny short hairs; stalk 2.5– 9 cm long × 1–3 cm thick, typically enlarged in the middle and tapered toward the base and apex, sometimes nearly equal; surface whitish to pale buff or pinkish buff, deeply pitted and ribbed, internally chambered; ribs branching and anastomosing and extending onto the lower surface of the cap. MICROSCOPIC FEATURES: spores 16–22 × 10–14 μm, broadly elliptic, smooth, with 1 large oil drop, uniseriate, hyaline; asci 250–300 × 14–18 μm, cylindric to subcylindric, 8-spored; paraphyses cylindric, enlarged at the tips. OCCURRENCE: solitary, scattered, or in groups on the ground in hardwoods or conifer forests; summer, fall, and early winter; widely distributed in North America; common. EDIBILIT Y: not recommended. Although this mushroom is consumed in Europe, the edibility of North American collections has not been established. COMMENTS: Costapeda crispa (Scopoli) Falck is a synonym. The pale cream to buff colors of the cap and stalk, and a cap margin that is not attached to the stalk, are diagnostic field characters. It sometimes fruits on moss-covered decaying wood. Helvella maculata (p. 167) is morphologically the most similar species, but is readily distinguished MACROSCOPIC FEATURES:

Pezizom ycetes

Beug-final.indb 160

Helvella crispa

by its medium brown cap. The Helvella lacunosa Fries complex is also similar (see Helvella vespertina, p. 170), but their caps are usually pale to dark gray or nearly black, and if cream colored they have grayish tones. The white to cream-colored Helvella lactea Boudier, which some authors consider to be a synonym of Helvella lacunosa, is distinguished from the similarly colored Helvella crispa by a smooth to at most fi nely pubescent cap underside and a cap margin that is att ached to the stalk rather than free.

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Helvella cupuliformis Dissing and Nannfeldt fruitbody consisting of a cap and stalk; cap 0.9–3.5 cm wide × 0.5–2 cm tall, deeply cup-shaped with an inrolled margin or somewhat saddle-shaped when young, becoming shallowly cup-shaped to flat and plate-like with the margin sometimes split in age; upper surface smooth, ochre-brown, drying dark brown to graybrown; underside pubescent to villose, gray-brown but paler than the upper surface; stalk 0.2–2.5 cm long × 0.2– 0.8 cm thick, indistinct to prominent, solid, round, slightly fluted at the base, white to cream, often light gray-brown near the top, densely villose to pubescent, especially at the apex. MICROSCOPIC FEATURES: spores 18–21 × 11.5– 13.5 μm, broadly elliptic, with 1 large oil drop, smooth, hyaline; asci 300–350 × 17–20 μm, MACROSCOPIC FEATURES:

8-spored; paraphyses 4.5–7 μm at the apex, clavate, contents fi nely granular, pale brown to brown. OCCURRENCE: single to gregarious on soil or humus in hardwood and coniferous forests; spring; eastern North America, also found in Oregon, Europe, and Asia; uncommon. COMMENTS: Helvella fibrosa (p. 163) is closely related but has a gray to gray-brown upper surface, somewhat narrower spores, and a longer, less robust stalk. Helvella rivularis Dissing and Sivertsen (not illustrated), a boreal and arctic species found in North America, Europe, and Asia, is also closely related and is differentiated by having a dark graybrown to dark brown hymenium that dries blackish brown. Helvella solitaria (p. 169) is easily distinguished by its ribbed stalk.

Helvella cupuliformis

H e lv e l l a

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Helvella elastica Bulliard

Com mon Elfin Sa ddle, Smooth-Sta lk ed Helv ell a

fruitbody consisting of a cap and stalk; cap 2–4 cm wide, depressed saddle-shaped to irregularly lobed, lobes typically bent downward toward the stalk, margin entire or somewhat lacerated; upper surface tan to pale grayish tan or dark grayish brown, smooth to somewhat wrinkled or sometimes pitted, dry or moist; lower surface white to pale tan, smooth; stalk 3–10 cm long × 3–10 mm thick, rounded, somewhat enlarged downward, whitish to buff, smooth overall or slightly roughened near the apex. MICROSCOPIC FEATURES: spores 17–20 × 10–14 μm, elliptic to oblong, smooth, with 1 large central oil drop, uniseriate, hyaline; asci 200–280 × 14– 20 μm, cylindric to subcylindric, 8-spored; paraphyses cylindric, with strongly enlarged tips, hyaline. OCCURRENCE: solitary, scattered, or in groups on the ground or on extremely rotted wood in conifer or hardwood forests; summer and fall in eastern North America, also winter and spring in the Southwest; widely distributed in North America; common. COMMENTS: Leptopodia elastica (Bulliard) Boudier is a synonym. Helvella albella (p. 156) is very similar, but its cap is typically darker and its lower surface is pubescent when young. MACROSCOPIC FEATURES:

Pezizom ycetes

Beug-final.indb 162

Helvella elastica

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Helvella fibrosa (Wallroth) Korf fruitbody composed of a cap and stalk; cap 1–3 cm wide × 0.5–1 cm tall, shallowly cup-shaped, undulating, planar, initially laterally compressed; outer surface smooth, dark gray to gray-brown (both fresh and dried); margin and underside paler, villose to densely pubescent; stalk 1–4 cm tall × 0.1– 0.5 cm wide, more or less equal, round and solid, densely pubescent, gray-brown. MICROSCOPIC FEATURES: spores 16–20 × 10–12 μm, broadly elliptic, smooth, unigutt ulate, uniseriate, hyaline; asci 240–260 × 17–19 μm, 8-spored; paraphyses 5–7.5 μm at apex, clavate, hyaline to pale brown. MACROSCOPIC FEATURES:

OCCURRENCE: solitary to gregarious on litter in mixed woods; spring through fall; boreal forests, montane habitats, and coastal regions of eastern and western North America as well as Norway, Sweden, Asia, and Australia; widely distributed. COMMENTS: Helvella chinensis (Velenovský) Nannfeldt and L. Holm, Helvella villosa (Hedwig) Dissing and Nannfeldt, Helvella pallidula N. S. Weber, and Helvella dissingii Korf are all synonyms (Korf 2008). It is important to note that Helvella villosa Schaeffer is a different species and is not a synonym of H. fibrosa. Helvella solitaria (p. 169) is easily differentiated by its ribbed stalk. See also Helvella cupuliformis (p. 161) and H. macropus (p. 166).

Helvella fibrosa

H e lv e l l a

Beug-final.indb 163

163

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Helvella latispora Boudier fruitbody consisting of a cap and stalk; cap 0.3–1.6 cm wide × 0.5–1.5 cm high, saddle-shaped to irregularly lobed, margin initially inrolled, then reflexed; upper surface smooth to rugose, undulate, pale gray-brown to brown, drying yellow-brown to brown; underside somewhat pubescent, cream to pallid gray-brown; stalk 0.6–2 cm long × 0.1– 0.5 cm thick, slightly thickened at base, round but base may be slightly fluted, solid, somewhat pubescent, white, drying cream to pallid grayish yellow-brown. MICROSCOPIC FEATURES: spores 17–20.5 × 11– 12.5 μm, broadly elliptic, with 1 large oil drop, hyMACROSCOPIC FEATURES:

aline; asci 290–330 × 13–15.5 μm, 8-spored; paraphyses 5–8.6 μm at apex, clavate, contents fi nely granular, hyaline. OCCURRENCE: solitary to gregarious on soil or litter in all types of woods; late spring through summer; eastern North America and rarely in boreal and montane regions of the west. COMMENTS: Helvella stevensii Peck and Helvella connivens Dissing and M. Lange are both synonyms. Helvella compressa (p. 158) is closely related and is found mainly in the west. See also Helvella albella (p. 156) and Helvella elastica (p. 162).

Helvella latispora

Pezizom ycetes

Beug-final.indb 164

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Helvella leucomelaena (Persoon) Nannfeldt fruitbody 3–7 cm high, 4– 6 cm wide, somewhat globose and closed at fi rst, becoming deeply cup-shaped with a short stalk, and shallowly cup-shaped at maturity; upper surface dark brown to gray-brown, smooth to slightly undulate-rugose; lower surface gray to dark brownish gray near the margin, becoming creamy white below and often white at the base; margin often split and somewhat reflexed; stalk up to 2 cm long, up to 1.5 cm thick, conspicuously ribbed to folded, or sometimes rudimentary. MICROSCOPIC FEATURES: spores 18–24 × 11– 15 μm, broadly elliptic to elliptic, smooth, uniguttulate, hyaline; asci 225–330 × 14.6–20 μm; paraphyses with clavate tips, brown, contents fi nely granular. OCCURRENCE: solitary, scattered, or in groups on leaf litter, among mosses, or on the ground in conifer or mixed woods; spring, summer, and fall; westMACROSCOPIC FEATURES:

ern North America, also reported in northeastern North America; occasional. EDIBILIT Y: unknown, not recommended. COMMENTS: Helvella confusa Harmaja and Helvella pedunculata Harmaja are synonyms. Helvella crassitunicata N. S. Weber (not illustrated), a similar species known on the west coast from Washington to Alaska, is unique in the genus by having the largest spores (22.5–28 × 12.5–15 μm) and distinctive thick-walled paraphyses. The two are best differentiated macroscopically by the more pronounced ribbing and stronger pubescence on the stalk and cup underside of Helvella crassitunicata (Abbott and Currah 1997). Helvella aestivalis (R. Heim and Remy) Dissing and Raitviir (not illustrated) is a similar species from circumpolar arctic tundra regions that can be easily differentiated based on the reddish brown colors of the cups and densely pubescent lower surface.

Helvella leucomelaena

H e lv e l l a

Beug-final.indb 165

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Helvella macropus (Persoon: Fries) P. Karsten fruitbody consisting of a cap and stalk; cap 1.5–4 cm wide × 4–12 mm high, deeply cup-shaped at fi rst, becoming shallowly cup-shaped at maturity; margin inrolled initially then expanded; upper surface dark brown to graybrown, smooth to slightly wrinkled, dry or moist; lower surface gray to gray-brown, sometimes with olivaceous tints, densely pubescent to villose especially near the margin; stalk 4– 6 cm long × 1.5– 4 mm thick, terete to slightly sulcate, somewhat enlarged downward, dry, solid, gray to gray-brown with a whitish base, pubescent to villose. MICROSCOPIC FEATURES: spores 18–25 × 10.3– 12.2 μm, fusiform-elliptic to subfusiform, with 1 large central oil drop and 2 smaller droplets, one at each end, usually fi nely punctate, hyaline; asci 175–250 × 14–18 μm; paraphyses with clavate tips, pale brown, contents fi nely granular. OCCURRENCE: solitary, scattered, or in groups on the ground, among mosses, or on decaying wood, in hardwoods or conifer forests; summer and fall; widely distributed in North America; occasional. COMMENTS: Cyathipodia macropus (Persoon: Fries) Dennis, Helvella brevis (Peck) Harmaja, and Paxina subclavipes (W. Phillips and Ellis) Seaver are all synonyms. Helvella macropus var. brevis Peck (not illustrated) is characterized by a much

Long-Sta lk ed Gr ay Cup

MACROSCOPIC FEATURES:

Pezizom ycetes

Beug-final.indb 166

Helvella macropus

shorter stalk (0.8–1.6 cm long). The distinctive subfusiform, fi nely roughened spores easily separate Helvella macropus from other cupulate Helvella species such as Helvella cupuliformis (p. 161) and Helvella fibrosa (p. 163).

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Helvella maculata N. S. Weber

Fluted Brow n Elfin Sa ddle

fruitbody with a cap and stalk; cap 1–5 cm wide × 0.5–3 cm tall, saddle-shaped to irregularly lobed, margin initially strongly inrolled but flaring out and often splitting in age; upper surface smooth to undulate, buff to rich medium brown or gray-brown, sometimes mott led; underside densely pubescent to villose, white to pallid gray; stalk 1–12 cm tall × 0.3–3 cm thick, fi nely pubescent, ribbed (with ribs extending to one third of cap underside), internally chambered, white with localized regions of brown to gray-brown pigmentation. MICROSCOPIC FEATURES: spores 18–23.5 × 11–14 μm, broadly elliptic, smooth, with 1 large central oil drop, hyaline; asci 268–300 × 15– 19 μm, 8-spored; paraphyses 5.5– 9 μm thick at MACROSCOPIC FEATURES:

apex, clavate, contents fi nely granular, brown in mass. OCCURRENCE: solitary to gregarious on soil or duff under both conifers and hardwoods; normally appearing late summer through fall but can appear early spring in coastal areas; reported only from western North America and China. EDIBILIT Y: unknown, undoubtedly poisonous if consumed raw or undercooked. COMMENTS: Helvella vespertina (p. 170) can have a mott led cap but has a gray-brown to black hymenium. Its margin is turned down when young and is fused to the stalk, while the margin of Helvella maculata is free. Helvella crispa (p. 160) is quite closely related to Helvella maculata (Abbott and Currah 1997).

Helvella maculata

H e lv e l l a

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Helvella pezizoides Afzelius: Fries fruitbody consisting of a cap and stalk; cap 0.25–1 cm wide × 0.2– 0.7 cm tall, variably saddle-shaped, irregularly lobed, cupshaped, or discoid, margin strongly inrolled initially, remaining somewhat incurved as it matures; upper surface smooth to undulate, black; underside pubescent to villose, dark brown to dark graybrown; stalk 0.3–2.5 cm tall × 0.1– 0.3 cm thick, base enlarged, apex tapered, round, sometimes with fluted base, solid, pubescent to villose, brown to blackish brown. MICROSCOPIC FEATURES: spores 17–21 μm × 10.5– 13.5 μm, broadly elliptic, smooth, with 1 large oil drop, hyaline; asci 210–315 × 14–19 μm, 8-spored; paraphyses 4– 9 μm, clavate, terminal cell 40– MACROSCOPIC FEATURES:

60 μm long, contents fi nely granular, brown to dark brown in mass. OCCURRENCE: solitary to gregarious on soil, humus, or rotted wood; summer; found in much of North America as well as Europe and Asia; uncommon. COMMENTS: Helvella ephippium Léveillé is related but has a paler hymenium and more cupulate cap. Note that Helvella ephippium sensu Cooke = Helvella pezizoides. Helvella atra (p. 157) is similar but is normally saddle-shaped to irregularly lobed, is not pubescent to villose on the underside of the cap, is not strongly involute when young, and has a lighter underside and stalk coloration (Abbott and Currah 1997).

Helvella pezizoides

Pezizom ycetes

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Helvella solitaria P. Karsten

R ibbed Elfin Saucer

fruitbody consists of a cap and stalk; cap 1.5–12 cm wide × 0.5–5 cm high, cupulate to shallowly cupulate, often laterally compressed, margin somewhat inrolled when young; upper surface undulate, smooth to rugose, gray-brown to dark brown, drying blackish brown; underside weakly to distinctly pubescent, ribs when present extending only slightly past the stalk, gray to dark gray-brown at margin, paler or white where attached to the stalk; stalk 0.5–12 cm tall × 0.2–5 cm thick, equal or enlarged downward, weakly to distinctly pubescent, sparsely ribbed to conspicuously ribbed, ribs sometimes anastomosed, solid, or with a few chambers, white to dark gray-brown, especially at apex, sometimes tinged buff to ochre in places. MICROSCOPIC FEATURES: spores 17–21 × 11–13.5 μm, broadly elliptic, smooth, with 1 prominent central oil drop, hyaline; asci 240–300 × 14–18 μm, with a lateral beak at base, 8-spored; paraphyses 7–8 μm at apex, clavate, terminal cell 90–145 μm long, contents granular, brown. OCCURRENCE: solitary to gregarious on the ground, humus, or rott ing wood under conifers or MACROSCOPIC FEATURES:

hardwoods; spring through fall, winter to spring in California; widely distributed in North America and Europe; fairly common. COMMENTS: Helvella queletii Bresàdola is a synonym (Harmaja 1977a). Helvella solitaria sensu Dissing = Helvella leucomelaena (p. 165). Helvella leucomelaena has larger spores, and has a stalk that is usually entirely below the ground surface. The similarly colored Helvella costifera Nannfeldt has more prominent ribbing on a broader stalk, and the ribbing extends farther on the underside of the cup. Helvella hyperborea Harmaja (not illustrated), a rare species from Alaska, is differentiated by its dark brown color and by prominent ribbing that extends farther on the underside of the cup, similar to the ribbing of Helvella costifera. Helvella unicolor (Boudier) Dissing is rare, found in Alberta, and similar in color and morphology to Helvella solitaria but is distinguished by its very broadly elliptic spores (17.4–20.8 × 13.1–15.2 μm). Small specimens of Helvella solitaria can resemble Helvella cupuliformis (p. 161). See also Helvella fibrosa (p. 163) and Helvella maculata (p. 167).

Helvella solitaria

H e lv e l l a

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Helvella vespertina N. H. Nguyen and Vellinga Bl ack Elfin Sa ddle, Fluted Bl ack Helv ell a

fruitbody 5–30 cm tall consisting of a cap and stalk; cap 2–5(–12) cm wide × 1–7(–15) cm high, mitre-shaped to convex or saddle-shaped, dry; margin typically wavy and often incurved, entire or somewhat lacerated, fused with stalk in several places; upper surface pale gray to black, wrinkled; lower surface grayish to black, glabrous, with ribs extending from the stalk apex; ribs often forked and interconnected; stalk 6–14(–25) cm long × 1–3(–10) cm thick, nearly equal or enlarged downward, dry, hollow or chambered; stalk surface dull white to gray or black, typically darkest on the ribs and toward the apex, sometimes ochraceous along lower portions, deeply pitted and conspicuously ribbed; ribs branching, anastomosing, and extending onto the lower surface of the cap. MICROSCOPIC FEATURES: spores 14.5–21.5 × 10– 13.5 μm, broadly elliptic, smooth to slightly wrinkled, unigutt ulate, hyaline; asci 238–268 × 13– 17 μm, 8-spored, not amyloid; paraphyses cylindric, with clavate tips, contents fi nely granular, brown. OCCURRENCE: solitary, scattered, or in groups on the ground or on decaying wood under conifers; fall, winter, and early spring; western North America, also reported from Michigan and Ontario; common. EDIBILIT Y: not recommended. COMMENTS: According to Nancy Weber (personal communication), “this species complex is in need of serious work,” a sentiment echoed by Else Vellinga, who is part of a team working on this complex in California. In western North America, Vellinga has not found the European Helvella lacunosa. Instead she fi nds two common species: Helvella vespertina N. H. Nguyen and Vellinga (= H. lacunosa sensu Morse [1945] and sensu other American authors) and Helvella dryophila Vellinga and N. H. Nguyen (Nguyen et al. 2013). MACROSCOPIC FEATURES:

Pezizom ycetes

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Helvella vespertina

The fall-fruiting conifer associate, Helvella vespertina, closely resembles H. lacunosa macroscopically and microscopically but sometimes can be much larger than European H. lacunosa, which is rarely more than 14 cm tall or 5 cm wide. Helvella dryophila is also strikingly similar to H. lacunosa macroscopically and microscopically but never gets to be larger than European H. lacunosa. Helvella dryophila is associated with oaks and fruits mid-December through April (June at higher elevations). It is reported from southern Oregon to southern California, though we have also found it as far north as Seatt le, Washington. It has been identified on root tips of Arbutus menziesii, and we have found it in a mixed fruit and nut tree orchard in Washington State. The cap is gray-brown to fre-

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quently black; the underside is bald to pubescent; the margin is att ached to the stalk in places; the stalk is typically quite short, blackish to whitish at the base, frequently with brown stains; and the spores are 17.5–21.5 × 11–12.5 μm. These two species are easily confused in mixed conifer-oak stands when their fruiting seasons overlap. There are similar rare species with other hosts (Else Vellinga, personal communication). Helvella sulcata Fries is a name sometimes given to pale to dark gray fruitbodies of eastern members of this complex that have distinctly ribbed stalks that lack pits. Some

authors accept a broad species concept and consider Helvella sulcata = Helvella lacunosa, while others consider it to be a distinct species. Helvella palustris Peck (not illustrated) is an eastern species, smaller than Helvella lacunosa, with a slender solid stalk with parallel ribs with litt le anastomosis, no lacunae, and a saddle-shaped or loosely lobed gray to black cap that remains free from the stalk. Häff ner (1987) considers it a variety of Helvella lacunosa, and it is microscopically very similar to H. lacunosa.

H e lv e l l a

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Humaria hemisphaerica (F. H. Wiggers) Fuckel Brow n-H a ir ed W hite Cup

fruitbody 1–3 cm wide × 1–2 cm high, cup-shaped; inner surface whitish to pale gray, smooth; outer surface brownish yellow, covered by a dense layer of brownish hairs that project from the margin over part of the upper surface. MICROSCOPIC FEATURES: spores 22–27 × 10–13 μm, broadly elliptic, fi nely to coarsely verrucose, bigutt ulate, obliquely uniseriate, hyaline; asci 230– 270(–350) × 19–23 μm, 8-spored; paraphyses fi liform, septate, with clavate tips. OCCURRENCE: scattered or in groups on soil, among mosses, or on decaying wood; late spring, summer, and fall; widely distributed in North America; fairly common. COMMENTS: Lachnea hemisphaerica (F. H. Wiggers) Gillet, Mycolachnea hemisphaerica (F. H. Wiggers) MACROSCOPIC FEATURES:

Maire, and Peziza hemisphaerica F. H. Wiggers are all synonyms. See also Geopora sepulta (Fries) Korf and Burdsall (in comments under Geopora arenosa, p. 89) if the species is found buried or partially buried, or Peziza vesiculosa (p. 223) if it is on manure piles. Trichophaea hemisphaerioides (p. 261) is a very similar species that also has 2 small oil drops, one at each end of a spore. Tricharina gilva (p. 259) spores lack oil drops. Hypomyces stephanomatis Rogerson and Samuels is a parasite on Humaria hemispherica. It produces a white cottony covering over the host, aseptate ascospores that are naviculate and 8–12.5 × 2–4.5 μm, and aseptate conidia that are 8–18.5 × 3.5–5.5 μm and cylindrical to elliptical with a flat basal abscission scar.

Humaria hemisphaerica

Pezizom ycetes

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Jafnea semitosta (Berkeley and Curtis) Korf fruitbody 2–5 cm wide, × 2–7 cm high, deeply cup-shaped; inner surface whitish to creamy yellow when young, becoming tan to brownish in age, smooth; outer surface pale creamy yellow to pale yellowish brown, covered by a dense layer of soft brown hairs; base of the cup crimped to form a short stalk, 1–2.5 cm long × 1– 2 cm wide, yellowish to brownish, covered with soft brown hairs. MICROSCOPIC FEATURES: spores 25–35 × 10–12 μm, elliptic to fusiform, verrucose when mature, biguttulate, obliquely uniseriate, hyaline; asci 300–325 × 14–15 μm, cylindric to clavate, 8-spored; paraphyses fi liform, septate, with clavate tips. MACROSCOPIC FEATURES:

solitary, scattered, or in groups on soil or decaying wood in conifer or hardwood forests; summer and fall; northeastern North America; infrequent. COMMENTS: Paxina semitosta (Berkeley and Curtis) Seaver, Peziza semitosta Berkeley and Curtis, and Sepultaria semitosta Morgan are all synonyms. Jafnea fusicarpa (W. R. Gerard) Korf is nearly identical but smaller, measuring 2–3 cm wide × 1 cm high, and has somewhat longer spores that measure 30–45 × 10–12 μm. OCCURRENCE:

Jafnea semitosta

Jafnea

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Lamprospora crechqueraultii var. crechqueraultii (P. Crouan and H. Crouan) Boudier MICROSCOPIC FEATURES: fruitbody 2–5 mm wide, shallowly cup-shaped to scutellate, inner surface smooth, pale orange to pale orange-yellow, margin indistinct; outer surface smooth, very pale orange; stalk rudimentary or absent. MICROSCOPIC FEATURES: spores 20–25 μm, globose, smooth at fi rst but soon conspicuously echinate, uniseriate; asci 300–325 × 25–27 μm, cylindric to subcylindric, 8-spored; paraphyses rather stout, with enlarged tips, hyaline. OCCURRENCE: scattered or in groups, on the ground or among mosses; summer and fall; eastern North America; uncommon.

Octospora crechqueraultii (P. Crouan and H. Crouan) Caillet and Moyne, and Ramsbottomia crechqueraultii (P. Crouan and H. Crouan) Benkert and Schumacher are synonyms. Lamprospora crechqueraultii var. macrantha Boudier (= Lamprospora macrantha [Boudier] Seaver) is nearly identical but has larger and longer spores that measure 30–35 × 20 μm excluding spines that reach a length of 10 μm, and a diameter of 3–4 μm at the base. COMMENTS:

Lamprospora crechqueraultii var. crechqueraultii

Pezizom ycetes

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Melastiza chateri (W. G. Smith) Boudier fruitbody 1.5– 6 mm wide, cup-shaped to shallowly cup-shaped, often distorted, stalkless; inner surface bright orange to reddish orange, smooth, shiny; outer surface orange, covered toward the margin by clumps of short reddish brown hairs that resemble small granules. MICROSCOPIC FEATURES: spores 17–20 × 9–13 μm, elliptic, ornamented by a coarse reticulum, sometimes with one or more spine-like projections at each end, often gutt ulate, hyaline; asci up to 300 × 12–15 μm, cylindric to subcylindric, 8-spored; paraphyses fi liform with clavate tips, hyaline; hairs 60–150 × 12–18 μm with 1–4 septa, walls 1–2 μm thick. OCCURRENCE: in groups or clusters on moist, sandy soil, sometimes among mosses, also reMACROSCOPIC FEATURES:

ported to occur on sawdust; summer and fall; eastern North America, west to Colorado, Idaho, and Alaska; infrequent. COMMENTS: In some field guides and reference works, the epithet is sometimes misspelled as “charteri.” Humaria miniata Fuckel and Melastiza miniata (Fuckel) Boudier are synonyms of an almost identical species, Melastiza cornubiensis (Berkeley and Broome) J. Moravec. Melastiza cornubiensis has identical spore size and ornamentation to M. chateri, but the hairs are paler brownish and the hymenium is orange. Some authors consider the two species to be identical, in which case the older name, Melastiza cornubiensis, would take precedence.

Melastiza chateri

Melastiza

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Microstoma floccosum (Schweinitz) Raitviir fruitbody 1–4.5 cm high, 3–10 mm wide, cup-shaped with a distinct stalk; inner surface bright red, smooth; outer surface and margin pinkish red to bright red, covered by conspicuous white hairs; stalk 2–3.5 cm long, 1.5–3 mm thick, nearly equal, smooth to fi nely roughened, white. MICROSCOPIC FEATURES: spores 20–35 × 15–17 μm, elliptic to fusiform, smooth, uniseriate, hyaline; asci 300–325 × 20 μm, cylindric to subcylindric; paraphyses fi liform, with slightly enlarged tips. MACROSCOPIC FEATURES:

Sh aggy Sca r let Cup

in groups or clusters on decaying hardwood branches; summer and early fall; eastern North America; occasional to fairly common, especially in the southern part of its range. COMMENTS: In some field guides and reference works, the epithet is sometimes misspelled as “floccosa.” Anthopeziza floccosa (Schweinitz) Kanouse, Geopyxis floccosa (Schweinitz) Morgan, and Plectania floccosa (Schweinitz) Seaver are synonyms. OCCURRENCE:

Microstoma floccosum

Pezizom ycetes

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Microstoma protractum (Fries) Kanouse (as protracta) fruitbody at fi rst funnelshaped, later cup-shaped then flattened and torn to lobed at the margin, with a stalk; cup up to 2 cm wide, opening by a small pore that is surrounded by a row of stiff, short, hyaline hairs; interior vivid rose-red to almost scarlet; upper portion of underside smooth; lower portion of underside clothed with soft, white hairs that are flexuous, forming a delicate but abundant tomentum, orange to bright orange-red; stalk 3–4 cm long × 0.1– 0.2 cm thick, gradually expanding into the cup, aerial part long, slender, often branched, giving rise to as many as eleven cups, upper half of stalk whitish, downy with hyaline hairs, lower half dark-colored; stalk arising from a buried, hard, elongate pseudorhiza attached to buried wood or roots of trees. MICROSCOPIC FEATURES: spores 25–50 × 10– 16 μm, elliptic-fusoid, narrowed toward either end, smooth, usually containing 1 large oil droplet and 1–2 smaller ones, uniseriate but slightly overlapped, hyaline to slightly yellowish; asci 200– MACROSCOPIC FEATURES:

275 × 20–23 μm, operculum somewhat laterally placed, 8-spored; paraphyses slender, sometimes extending beyond the asci, not flexuous, dichotomously branched, only slightly enlarged at the tips, with red contents which turn green with Melzer’s; hairs 6–7 μm, blunt, hyaline; stalk and cup contain a middle layer of gelatinous hyphae. OCCURRENCE: on buried sticks and roots in both hardwood and coniferous forests; fruits right after snowmelt, sometimes on frozen ground; montane regions or northern latitudes, western North America, northern Europe and the Alps, Russia, and Japan; uncommon. COMMENTS: A unique feature of this fungus is the presence of a pseudorhiza. Interestingly, the pseudorhiza appears to be perennial, and older pseudorhizae give rise to aerial stalks with more branching and therefore more aerial cups than younger pseudorhizae. Two additional unique features are a gelatinous layer in both the stalk and the cups, and the dichotomously branched paraphyses.

Microstoma protractum

M icrostom a

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Midotis irregularis (Schweinitz) Cooke fruitbody 5–7 cm wide × up to 3 cm tall, a somewhat ear-like to split cupshaped clusters arising from a common stalk-like base; individual fruitbodies up to 3 cm wide × 3 cm tall, at fi rst closed then opening and becoming much elongated on one side, irregularly lobed or lacerated; fertile upper surface almost black; exterior scurfy, dark chestnut-brown; base up to 2 cm long, partially below ground. MICROSCOPIC FEATURES: spores 8–10 × 3–4 μm, smooth, with 2–3 oil drops, uniseriate to partly biseriate, subhyaline; asci 50–70 × 4–5 μm, cylindricclavate with a nearly truncate to slightly rounded apex; paraphyses terminated by a lanceolate, 1– 3 septate head, 18–30 × 3–4 μm, that projects beyond the hymenium and is easily detached. OCCURRENCE: solitary or in cespitose clusters on rotten wood and branches on the ground; sumMACROSCOPIC FEATURES:

mer and fall; eastern North America, Oregon; uncommon. COMMENTS: Ionomidotis irregularis Durand, Peziza irregularis Schweinitz, and Otidea doratophora Saccardo are synonyms for this litt le-known fungus. Midotis species typically start out subglobose and closed, expanding and usually becoming vertically elongated on one side when mature. The fruitbodies can be yellow, brown, dark blue, or olivaceous, with an exterior that is covered with short hair-like hyphae that typically turn momentarily violet with KOH. There are several named species in North America, but most are only known from the type locality. Midotis irregularis differs from Otidea species and Wynnea species in having relatively small inoperculate asci and smaller spores (versus operculate asci). See also Chlorencoelia versiformis (p. 366).

Midotis irregularis

Pezizom ycetes

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Morchella americana Clowez and C. Matherly Com mon Mor el, Y ellow Mor el

fruitbody 4–22(–40) cm tall, composed of a large sponge-like head and a stalk; head 2–11(–22) cm high × 1.5–4(–13) cm wide, oval to conic or somewhat cylindric, hollow, divided into pits and ridges of variable color, continuous with the stalk below, without a substantial rim at the point of att achment; with 12–30 vertical ridges and numerous horizontal and oblique ridges, along with scattered, sunken transecting ridges; pits round to vertically elongated and irregular, glabrous or very fi nely tomentose, grayish or brownish or almost black when young, becoming gray to yellowish at maturity; ridges glabrous or nearly so, anastomosing, whitish to pale yellowish when young, yellow or yellow-brown at maturity; stalk 2–12(–24) cm long, 2– 9(–14) cm thick, nearly equal or enlarged (sometimes massively) to-

MACROSCOPIC FEATURES:

ward the base, hollow, surface whitish, granular, often ribbed. MICROSCOPIC FEATURES: spores (17–)18–22(–24) × 11–13(–15) μm, elliptic, smooth, egutt ulate, uniseriate, hyaline, creamy yellow to orange in mass; asci 225–325 × 15–25 μm, cylindric-clavate, 8-spored; paraphyses 75–180 × 5–15 μm, clavate or enlarged toward the tips, nearly hyaline; elements on sterile ridges 75–160 μm long × 10–27.5(–37.5) μm wide, septate; terminal cell subclavate to clavate or subfusiform to widely cylindric. OCCURRENCE: solitary, scattered, or in groups on soil in a variety of habitats, often associated with white ash but also found near dead elms, in old apple orchards, riparian areas, burned areas, or mixed hardwood forests, sometimes under conifers, often on steep slopes or along old railroad beds; spring

Morchella cf. americana

Morch ell a

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and early summer; widely distributed throughout North America; occasional to locally abundant. EDIBILIT Y: edible and choice, but must be thoroughly cooked; one of the most highly prized and sought-after mushrooms for the table. COMMENTS: Morchella americana is the most common yellow morel in North America. Morchella populina Clowez and R. Lebeuf is a synonym. (Morchella populina var. lata Clowez and R. Lebeuf is a synonym of the late-fruiting M. americana var. lata). Morchella rigida (Krombholz) Boudier sensu Clowez (synonym Morchella californica Clowez and D. Viess) is so closely related to Morchella americana that a decision may be made to group them together, in which case the name Morchella rigida will take precedence. Morchella esculentoides M. Kuo, Dewsbury, Moncalvo, and S. L. Stephenson will be declared a synonym in the M. americana/M. rigida group. When found with a stout and strongly enlarged stalk base, members of the yellow morel clade have been known as Morchella crassipes (Ventenat) Persoon = Morchella esculenta var. crassipes (Ventenat) Kreisel, but DNA evidence has shown that the large-stalked examples are simply older classi-

cal yellow morels (Morchella americana and lookalikes) that have grown under optimal conditions (Jung et al. 1993). Without doing DNA analysis, Morchella ulmaria Clowez (synonym Morchella cryptica M. Kuo and J. D. Moore) cannot be reliably differentiated from M. americana/M. rigida, although it is frequently somewhat paler and its ridges are usually more flattened. Morchella ulmaria is found in midwestern hardwood forests, especially in association with ash and elm trees. M. ulmaria is most common under American elms the fi rst year after they have died. Morchella americana var. lata, found in Quebec, has a head that is dirty gray ochraceous, soon fading to ochraceous, and is often as wide as it is tall, having the appearance of an equilateral triangle. The edges of the ridges are sometimes tinged reddish, and the pits are more numerous and less regular than in M. americana. It is differentiated by the presence of a distinct sinus and a late fruiting season, after the main fruiting of M. americana. See also Morchella prava (p. 187) and Morchella rufobrunnea (p. 190).

Pezizom ycetes

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Morchella angusticeps Peck

Com mon Easter n Black Mor el

fruitbody up to 18 cm high, composed of a fairly large conical to oval head and a long stalk; head 2– 6 cm wide, 2.5– 9 cm high, sponge-like, hollow, divided into pits and ridges, continuous with the stalk below, typically attached to the stalk with a slight overhang or rim, hollow; pits elongated and irregular, yellow-brown to gray-brown, with 16–24 vertical ridges, with occasional shorter secondary ridges and frequent sunken transecting ridges, anastomosing, dark brown to brownish black; head att ached to stalk with a 2–5 mm deep × 2–5 mm wide sinus; stalk 2–10 cm long, 1–4 cm thick, nearly equal or sometimes enlarged near the base, hollow; surface granular, sometimes ribbed, white to dingy yellow. MICROSCOPIC FEATURES: spores 22–28 × 11–15 μm, broadly elliptic, smooth, sometimes with small, polar, exterior oil drops, uniseriate, hyaline, cream to yellowish or orangish in mass; asci 250–400 × 17– 30 μm, cylindric-clavate, 8-spored; paraphyses cylindric, multiseptate, branched near the base, with slightly enlarged tips; elements on sterile ridges 100–200 μm long × 7.5–35 μm wide, septate; terminal cell widely cylindric with a rounded to clavate or irregular apex. OCCURRENCE: solitary, scattered, or in groups on soil in a variety of habitats, including mixed woods, associated with cherry, poplars, or pines, and mixed hardwood forests; spring; widely distributed in eastern North America; fairly common. EDIBILIT Y: edible and choice, but must be thoroughly cooked. COMMENTS: Th is member of the Morchella elata clade is generally the fi rst true morel to appear in eastern North America in spring. Although choice, they have caused gastrointestinal distress in some individuals, sometimes when consumed with al-

cohol. The European Morchella pulchella Clowez and Petit (not illustrated; synonyms Morchella angusticeps sensu Clowez and Morchella septentrionalis M. Kuo, J. D. Moore, and Zordani) is found in northern portions of eastern North America (from about 44° N latitude and northward). It is characterized by its small size (4–7.5 cm tall), frequent association with the deadwood of hardwoods, and its slightly smaller spores (19–)20–22(–25) × 11– 15 μm. (Note: Slight differences between European M. pulchella and M. septentrionalis may justify the retention of M. septentrionalis as a distinct taxon.) See also Morchella elata Fries (p. 184).

Morch ell a

181

MACROSCOPIC FEATURES:

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Morchella angusticeps

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Morchella diminutiva M. Kuo, Dewsbury, Moncalvo, and S. L. Stephenson W hite Mor el, Tu lip Mor el MACROSCOPIC FEATURES:

fruitbody a ridged and pitted head on a stalk; overall 3–10 cm tall; head 1.5–5 cm high × 1–3 cm broad, variable in shape but generally conical and somewhat pointed, without a substantial rim at the point of att achment to the stalk, hollow; ribs more or less vertically oriented, with 8–16 primary ridges and occasional secondary ridges, irregularly anastomosing, with scattered to frequent, sunken, transecting horizontal ridges; ridges rounded, about 1 mm thick, glabrous, whitish, tan to yellow at maturity; pits vertically elongated, smoky to grayish brown when young, tan to yellow at maturity; stalk characteristically as long as or longer than the head, 1–7 cm long × 0.5–2 cm wide, nearly equal, sometimes enlarged at the base, smooth or with granules, whitish to cream or pale tan, hollow. MICROSCOPIC FEATURES: spores (18–)20–24(– 26) × (10–)11–16(–18) μm, elliptic, smooth, eguttulate, uniseriate to partially overlapping, hyaline, orangish yellow in mass; asci 175–325 × 12.5–25 μm long, cylindric or subcylindric, 8-spored; paraphyses 125–250 × 7.5–20 μm, at most slightly enlarged at tips, nearly hyaline; elements on sterile ridges 75–175 μm long × (7.5–)10–30 μm wide, septate; terminal cell usually subfusiform at maturity. OCCURRENCE: solitary to gregarious in grassy places, usually on the edges of wooded areas in

a wide variety of hardwood forest habitats, often found with tulip trees, hickories, and ash, also associated with other hardwoods, especially old apple trees, not associated with conifers; late spring; widely distributed outside the coastal plain of eastern North America. EDIBILIT Y: a fi ne edible when thoroughly cooked; never eat raw. COMMENTS: Morchella sceptriformis Clowez and C. Matherly (synonym Morchella virginiana O’Don nell and S. A. Rehner) is slightly larger, 5– 12.5 cm tall × 2–3.5 cm wide at the widest point, and is found only under tulip trees in river bottoms, drainage areas, or coastal plains from Virginia to Northern Mississippi in April and May. Elements on sterile ridges in M. sceptriformis are variably shaped (cylindrical to subcapitate and subfusiform), scattered, and difficult to locate. In contrast, elements on sterile ridges of M. diminutiva are primarily widely fusiform and are easy to locate. Both Morchella diminutiva and M. sceptriformis can be difficult to differentiate from a small yellow morel, Morchella americana (p. 179), but both have vertically arranged pits rather than the more or less random arrangement of the pits in M. americana. In the past, both M. diminutiva and M. sceptriformis have been known as Morchella deliciosa Fries sensu N. S. Weber and A. H. Smith (1985).

Pezizom ycetes

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Morchella diminutiva

Morch ell a

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Morchella elata Fries sensu Clowez fruitbody a sponge-like head on a stalk, 6–20 cm tall; head 3–15 cm tall × 2– 9 cm wide, conic to ovoid, pitted, and ridged with 12–20 primary vertical ridges and numerous horizontal ridges, giving a laddered appearance; ridges glabrous to very fi nely tomentose, pale to dark gray when young, aging dark grayish brown to nearly black, bluntly rounded when young, sharpened and eroded in age; pits vertically elongated, glabrous or very fi nely tomentose, gray when young, becoming grayish olive to brownish yellow at maturity; head att ached to stalk with a 2–5 mm deep × 2–5 mm wide sinus; stalk 3–10 cm tall × 2– 6 cm wide, often basally clavate to subclavate, glabrous to fi nely mealy with whitish granules, with longitudinal ridges and channels at maturity. MICROSCOPIC FEATURES: spores 18–24 × 10–13 μm, elliptic, smooth, hyaline; asci 220–300 × 12–25 μm, 8-spored; paraphyses 150–250 × 7–15 μm, apices rounded, subclavate, subacute, or subfusoid; elements on sterile ridges 125–300 long × 10–30 μm wide, septate; terminal cell rounded to clavate or subfusiform. MACROSCOPIC FEATURES:

L a ndsca pe Mor el, Mu lch Mor el

solitary to gregarious where ground bark chips have been applied the previous fall to landscaped new construction alongside highways, shopping malls, or homes, occasionally on bare soil surrounding new construction, and occasionally after bark chips were applied to an older landscaped area; early spring; widespread; common. EDIBILIT Y: choice, but possible lead contamination if near a highway, or possible herbicide uptake in manicured areas. COMMENTS: Morchella elata (synonyms Morchella importuna M. Kuo, O’Donnell, and T. J. Volk and Morchella costata [Ventenat] Persoon) has a worldwide distribution. In North America it is most common in landscape beds the year after bark mulch has been placed in a newly landscaped bed. It can appear in stupendous abundance. Morchella hotsonii Snyder (not illustrated; so far known only from the 1935 type collection) is very similar. It was known from landscaped beds in the Pacific Northwest. It is distinguished by DNA and its fi nely tomentose ridges and pits. We have found a similarappearing morel under oaks in Washington, as well as under conifers in the Pacific Northwest. OCCURRENCE:

Morchella elata

Pezizom ycetes

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Morchella frustrata M. Kuo Mou nta in Blond Mor el, W ester n Blond Mor el

fruitbody 6–20+ cm tall × 4–10 cm wide, consisting of a ridged and pitted head on a stalk; head 4– 6+ cm tall, hollow, narrowly to broadly conic when young, expanded at maturity but typically remaining relatively narrow in relation to its height, or sometimes becoming globose; ridges vertically arranged with 16–22 primary vertical ridges, very few shorter secondary ridges, and several sunken transecting horizontal ridges; cross-ribs form vertically elongated pits in the manner of a typical black morel; ridges nearly glabrous when young, dry and waxy in age; sterile edges typically remain intact, initially pale grayish tan, becoming ivory to pale tan in age and often with rusty ochre stains; pits primarily vertically elongated, initially light smoky gray, be-

MACROSCOPIC FEATURES:

coming straw-yellow in age; stalk 2–4 cm tall × 1– 2.5 cm wide, hollow, flared out at the apex, swung down somewhat and fused to the lower edge of the head, creating a shallow rim 2–4 mm deep × 2–4 mm wide. MICROSCOPIC FEATURES: spores 20–29 × 14–19 μm, elliptic, smooth, hyaline; asci 225–300 × 15–25 μm, 8-spored; paraphyses 100–225 × 10–25 μm, apices usually rounded; elements on sterile ridges 100– 175 long × 12.5–20 μm wide, septate; terminal cell clavate or subclavate. OCCURRENCE: solitary to gregarious in pine forests, fi r and Douglas fi r forests, and mixed stands of conifers and hardwoods, under oaks, but not in burns; late spring to early summer; California to Washington; common.

Morchella fr ustrata

Morch ell a

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EDIBILIT Y:

choice, must be thoroughly cooked, never eat raw. COMMENTS: DNA testing has shown the western blond morel to be a genetically distinct species in the Morchella elata clade of black morels. It has ridges and pits on the head that are aligned like those on a black morel, with the color of a yellow morel. The flavor is milder than a black morel and not as complex as a yellow morel, but it still is wonderful to eat. Morchella fr ustrata differs from the

yellow morels in that the western blond has vertically elongated, generally vertically aligned pits, and a sinus where the stalk attaches to the head (a feature that Kuo has described as “an ant racetrack”). Morchella snyderi (p. 191) can sometimes have pale Morchella esculenta-like coloring and is distinguished by the distinctly lacunose stalk and fi nely tomentose pits. Morchella fr ustrata has glabrous pits, and an equal to subclavate nonlacunose stalk.

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186

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Morchella prava Dewsbury, Moncalvo, J. D. Moore, and M. Kuo fruitbody 5–10 cm tall, a ridged and pitted sponge-like head on a stalk; head 3– 6 cm tall × 2–5 cm wide, irregular in shape, often somewhat ovoid, subconic at apex, bluntly attached to stalk; with 12–18 vertical ridges, numerous horizontal and oblique ridges, and scattered, sunken, transecting ridges; ridges glabrous to very fi nely tomentose, thick, flattened or widely rounded when young, sometimes sharpened and eroded in age, whitish to very pale yellowish or tan when young, pale brownish yellow at maturity with yellowish brown to reddish brown in places; pits asymmetrical and irregular in outline and size, glabrous or very fi nely tomentose, gray to nearly

MACROSCOPIC FEATURES:

black when young, often remaining dark for a long time before turning pale brownish yellow at maturity; stalk 2.5–4 cm tall × 1–3 cm wide, nearly equal above a subclavate base, smooth or nearly so. MICROSCOPIC FEATURES: spores (16–)17–21(–24) × (8–)10–12(–13) μm, elliptic, smooth, hyaline; asci 200–300 × 15–25 μm, 8-spored; paraphyses 100– 175 × 5–12.5 μm; cylindric, apices rounded to subclavate, septate; elements on sterile ridges difficult to locate or distinguish from the paraphyses. OCCURRENCE: habitat highly variable; found from 43° to 50° N latitude across North America (known from Montana, South Dakota, Michigan, Wisconsin, Saskatchewan, Ontario, and Quebec); spring.

Morchella prava

Morch ell a

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EDIBILIT Y:

edible and choice but must be thoroughly cooked. COMMENTS: Morchella prava resembles a thickfleshed and contorted Morchella americana (p. 179). Microscopically the only difference is that Morchella prava lacks distinctive sterile elements on the ridges, while the sterile elements on both M. americana and M. ulmaria Clowez are easily located. Kuo (2005) called this species the “Classic North American Morel III” and it appears to be the same as Morchella deliciosa sensu Miller (Miller and Miller 2006). There is a sand morel, anchored by a large network of sand-encrusted hyphae, found in dunes in coastal eastern North America as well as along

the Great Lakes that is morphologically similar to Morchella prava. To our knowledge, it has not been sequenced. There is a morphologically similar morel, Morchella vulgaris (Persoon: Fries) Boudier, known at least from Europe. When growing in sand dunes, it is anchored by a large network of sand-encrusted hyphae and has been known as Morchella dunensis (Castañera and G. Moreno) Clowez. The European sand morel has the general shape and characteristics of a morel in the Esculenta clade (a yellow morel) but has a peculiar odor of morels, cooked crustaceans, stagnant water, and old mop. It is found on the coasts of France and Spain, where it is restricted to some well-known spots.

Pezizom ycetes

188

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Morchella punctipes Peck

E aster n H a lf-Fr ee Mor el

fruitbody up to 18 cm high, composed of a conic head and a long stalk; head 1.5–4.5 cm wide × 1–4 cm high, broadly conic with a round to blunt apex, sponge-like, hollow, divided into pits and ridges, with 15–26 primary vertical ridges and infrequent shorter secondary transecting horizontal ridges; pits elongated and irregular, grayish tan to yellow-brown; ridges anastomosing, brownish, darkening to grayish brown or brownish black in age; att ached to the stalk about midway and flaring below with a sinus 0.8– 2 cm deep; underside of flaring head whitish and granular; stalk 1.5–15 cm long × 1–4.5 cm thick, enlarging downward or nearly equal overall, hollow, often fragile, surface whitish, slightly granular, commonly ribbed. MICROSCOPIC FEATURES: spores 22–25(–30) × (10–)14–18 μm, broadly elliptic, smooth, sometimes with small polar external oil drops, uniseriate, hyaline, cream to yellowish or orangish in mass; asci 200–350 × 15–25 μm, cylindric-clavate, 8-spored; paraphyses fi liform, septate, with clavate tips. OCCURRENCE: solitary or scattered on the ground in hardwood forests or old apple orchards; spring; MACROSCOPIC FEATURES:

widely distributed in eastern North America; common. EDIBILIT Y: edible. COMMENTS: The DNA of the western collections of half-free morels is distinct from eastern collections, and both North American taxa are different from Mitophora semilibera (DeCandolle) Léveillé of Europe. Morchella punctipes Peck typically fruits before the common or yellow morel and is easily recognized because of its half-free head and hollow stalk. Morchella populiphila M. Kuo, M. C. Carter, and J. D. Moore (Kuo et al. 2012), the western halffree morel, can be found under black cottonwoods along river bottoms and is macroscopically and microscopically very similar to the eastern halffree morel. Verpa bohemica (p. 267) is similar, but its head is att ached only at the top of the stalk, it has extremely large spores that measure 55–85 × 15–22 μm, and its stalk is usually stuffed with pithy material. Members of the Verpa conica (p. 268) group have a smooth to fi nely wrinkled head that is rounded to flat at the center and att ached to the stalk only at the top of the head.

Morchella punctipes

Morch ell a

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Morchella rufobrunnea Guzmán and F. Tapia R ed-Brow n Blushing Mor el

fruitbody a ridged and pitted head on a stalk; head 6–12 cm tall × 3–5 cm wide, usually conical when young, remaining conical or becoming ovoid at maturity, att ached at the bottom to the stalk; ribs typically more or less vertically arranged, anastomosing, whitish to gray, maturing yellowish, brownish, or brownish-yellow, bruising reddish orange to salmon when young; pits vertically elongate when young, irregularly shaped in age, dark brown when young and contrasting sharply with the pale ridges, maturing dull yellow and nearly concolorous with the ridges; stalk 3– 9 cm high × 1–2.5 cm wide, nearly equal or enlarged at the base, irregularly wrinkled near the base, with minute dark granules towards the top, whitish to cream or pale gray to gray-brown, bruising in spots or becoming almost completely reddish brown in age. MICROSCOPIC FEATURES: spores (19–)20–24(–25.5) × (13–)14–16(–17) μm, smooth, elliptic, egutt ulate, uniseriate, hyaline, pale orange or yellowish orange in mass; asci 300–360 μm long × 16–20 μm wide. MACROSCOPIC FEATURES:

solitary to gregarious, sometimes in large clusters; under subtropical oak, sweet-gum, or white-alder, sometimes in dirt roads and landscaping areas; winter and spring, year-round in Hawaii; known from the Gulf Coast of Mexico, Israel, and Australia, in landscaping in coastal California and Hawaii and possibly other warm, moist environments; common and abundant at times. EDIBILIT Y: good to choice if well cooked; do not sample raw. COMMENTS: Michael Kuo has suggested that this is the mushroom that has been known as Morchella esculenta (Linneaus) Persoon in California. Morchella rufobrunnea is found in California in the winter, usually the year following ground disturbance. At full maturity it closely resembles a classical yellow morel (see M. americana, p. 179). Morchella guatemalensis Guzmán and F. Tapia and Morchella herediana Guzmán and F. Tapia are two morel species of Mexico and Central America that have not spread northward (Guzmán and Tapia 1998). OCCURRENCE:

Morchella rufobrunnea

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Morchella snyderi M. Kuo and Methven fruitbody 6–14 cm tall, composed of a ridged and pitted head on a stalk; head 3.5–8 cm tall × 3–5 cm wide, conic to broadly rounded, often broadening near the base as it matures, att ached to the stalk with a rim, hollow, minutely velvety when young, dry and smooth at maturity, edges sterile and not eroding in age, steely gray to dark brownish or greenish or burgundybrown when young, becoming black or blackish at maturity; ribs 16–22 with frequent shorter secondary ribs and occasional sunken, transecting horizontal ridges; pits fi nely tomentose, steely gray to dull grayish or greenish or reddish tan when young, grayish tan or light brown in age; stalk 3.5– 7 cm tall × 2.5–4 cm wide, flared out at the apex and swung down a bit to fuse to the lower edge of the head, creating a shallow rim 2–4 mm deep × 2– 4 mm wide, at fi rst fi nely mealy with whitish gran-

MACROSCOPIC FEATURES:

Com mon W ester n Black Mor el

ules, becoming prominently granulated, usually prominently ribbed or lacunose at maturity, ivory to tan with a rose flush, aging tan or rosy tan. MICROSCOPIC FEATURES: spores (23–)26–37 × 15– 23 μm, elliptic, smooth, hyaline; asci 225–300 × 17– 33 μm, 8-spored; paraphyses 100–200 × 7.5–20 μm, subclavate to subcapitate, septate; elements on sterile ridges 75–175 × 10–20 μm, septate; terminal cell subclavate to subcapitate. OCCURRENCE: solitary to gregarious to cespitose on sand, soil, litter, and duff, in burns no sooner than the second spring after an intense wildfi re (except for fruiting in unburned islands); spring, also early summer at higher elevations; western North America; common and sometimes abundant. EDIBILIT Y: choice when cooked thoroughly; never eat raw.

Morchella snyderi

Morch ell a

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COMMENTS:

Th is black morel has gone by several names, including Morchella elata Fries, Morchella angusticeps Peck, and Morchella conica (Persoon: Fries) Persoon. Morchella snyderi is a highly variable fungus that can have the coloring of a yellow morel, or can be gray, tan, reddish brown, or greenish. The color of its ridges varies from only slightly darkening to jet-black in age. It is one of about ten western black morels in the Morchella elata clade. Other western black morels from unburned

woods include Morchella brunnea M. Kuo (natural black morel, Kuo et al. 2012) and an unnamed species (Mel-8, O’Donnell et al. 2011), which are both found associated with hardwoods. The base of the stalk of Morchella brunnea is equal to subclavate and is not ridged or lacunose, which are distinctive features of the stalk of Morchella snyderi. See also Morchella elata (p. 184) and Morchella angusticeps (p. 181).

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192

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Morchella tomentosa Kuo Gr ay Mor el, Ghosts, Black Foot Mor el, Fuzzy Foot

fruitbody a ridged and pitted head on a stalk; head 3–11 cm tall × 2–5 cm wide, elongate-ovoid when young, expanding variously in age, hollow; ribs lacking fertile tissue, conspicuously velvety to somewhat hairy only when young, appearing bald in age, silvery gray when young, maturing gray to black but soon cracking and breaking away to expose white to ivory underlying tissue; pits deep gray to black when young and increasingly lighter as the mushroom matures, eventually becoming gray to tan to dark ivory at maturity; stalk hollow, 2– 6 cm high × 1–4 cm wide, flared out at the apex and swung down a bit to fuse

MACROSCOPIC FEATURES:

to the lower edge of the head, creating a shallow rim, often swollen at the base, dark gray to nearly black and densely velvety when young, becoming pale gray to tan or ivory with tuft s of brown hyphal tips as it matures. MICROSCOPIC FEATURES: spores 18–20 × 8–12 μm, elliptical, without oil droplets, smooth; asci 8-spored; paraphyses cylindric to subclavate, with rounded or subacute tips, 2–5 septae; hairs abundant on sterile ridges and stalk surface, 275 × 15 μm, walls brown in KOH. OCCURRENCE: solitary to gregarious on burnt soil; conifer forests the fi rst year following a burn and

Morchella tomentosa

Morch ell a

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to a lesser degree the second year after a burn; late spring, summer; western North America; abundant at high elevation and northern latitudes. EDIBILIT Y: choice when thoroughly cooked; never eat raw. COMMENTS: Distinguishing features of Morchella tomentosa are a stalk and ribs that are blackish and densely velvety when young, and the eroded look of the edges of the ribs when old. In age, M. tomentosa is colored much like a western blond morel or a yellow morel, but neither the western blond nor the yellow morels are burn species. Morchella carbonaria Clowez and R. Chesnaux (not illustrated; found under burned cedar) also fruits late and is strikingly similar. These two species sometimes appear as late as August or even September at high elevations when there has been sufficient moisture. There are at least four species of burn morels that fruit earlier in the season, right after fruiting of the natural black morels found in non-burned areas. These species are Morchella anthracophila Clowez and D. Winkler, Morchella eximia Boudier (synonym Morchella septimelata M. Kuo), Morchella exuberans Clowez, Hugh Smith, and Sandi Smith (synonym Morchella capitata M. Kuo and M. C. Carter), and Morchella sextelata M. Kuo. These four species appear in burns the fi rst year after a fi re, rarely two years after a fi re. Commercial pickers know these four burn morel species as “pinks” and “greens,” but the coloration of each of these species is variable, and color cannot be used to distinguish between them. Morchella exu-

berans is easily distinguished by elements on the sterile ridges that have a terminal cell that is usually capitate, often dramatically so. We are aware of no macroscopic or microscopic way to reliably distinguish M. anthracophila, M. eximia, and M. sextelata, none of which have capitate cells on their sterile ridges. The image of Morchella exuberans Clowez, H. Smith, and S. Smith (p. 22) is from the type collection found at a burn in California. The highly contorted form and exceptionally thick flesh are not typical of M. exuberans, which typically is macroscopically indistinguishable from M. anthracophila, M. eximia, and M. sextelata.

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Neottiella vivida (Nylander) Dennis fruitbody up to 1.5 cm broad, closed at fi rst, expanding to turbinate, then cup-shaped, fi nally flat, sessile, or with a short stalk; upper surface reddish orange, the margin crenate and fringed with a few delicate hairs; underside whitish or pale orange, clothed with undulating, entangled, downy hair; stalk absent or buried in soil or sand. MICROSCOPIC FEATURES: spores 22–25 × 13–15 μm, broadly elliptic, ornamented with a more or less continuous hyaline reticulum, containing 1–2 large oil drops, uniseriate, obliquely arranged in the ascus, often with the ends overlapping; asci reaching 300–350 × 20 μm; paraphyses enlarged at tips, up to 6 μm wide, fi lled with orange granules that turn green in Melzer’s; hairs up to 300 × 10–12 μm, thinwalled, septate, hyaline. MACROSCOPIC FEATURES:

solitary to scattered or gregarious on soil among mosses, especially Polytrichum, apparently growing on dead remnants and often hidden by the living mosses; known coast to coast in the northern forests and in northern Europe; fall and early winter; common but seldom noticed. COMMENTS: Leucoscypha vivida (Nylander) Dennis and Rifai is a synonym. Neottiella rutilans (Fries) Dennis is virtually identical except for having ascospores with a more or less continuous hyaline reticulum. Neottiella hetieri Boudier (not illustrated) has a preference for moss-covered old burns, and its ascospores are smaller, 14–17 × 8– 9 μm. Some authors synonymize Octospora with Neottiella, but Octospora species have a glabrous underside while Neottiella species are clothed in downy hairs on the underside. OCCURRENCE:

Neott iella vivida

Neottiella

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Neournula pouchetii (Berthet and Riousset) Paden Rose Goblet, W ester n U r nu la

fruitbody 2.0–3.0 cm wide × 4.0 cm tall, consisting of a cup and stalk; club-shaped and hollow in the upper part when very young, expanding and spreading to urnshaped or margin splitt ing and spreading when old; upper surface pale pink-gray to purplish gray, becoming gray at maturity; margin with sharp, pointed to rounded teeth; underside when fresh with a pale gray felted layer, at fi rst nearly white, darkening to brown-gray to gray when old, bruising brown or brown-black, paler to nearly white at the base; stalk 2–4 cm tall × 0.5–1.5 cm thick, usually buried, with basal rhizomorphs, paler than the underside; flesh tough, leathery. MICROSCOPIC FEATURES: spores 23–32 × 8–10.5 μm, narrowly elliptic to somewhat cylindrical, smooth when young, then with low random ridges and warts that dissolve in dilute KOH, sometimes with a few small oil drops, uniseriate, hyaline; asci 290– MACROSCOPIC FEATURES:

390 × 12–15 μm, cylindrical, suboperculate, opercula eccentric, inamyloid, relatively thick-walled, base narrow and curving, 8-spored; paraphyses 3– 3.5 μm wide, narrow, septate, freely branched and anastomosing to form a three-dimensional network with tips embedded in a pale brown, amorphous material. OCCURRENCE: scattered to gregarious in conifer stands; spring and summer; Pacific Northwest, eastern Canada, France, and North Africa; locally common. COMMENTS: Neournula pouchetii is a European species, possibly genetically distinct from North American material. Neournula nordmanensis Paden and Tylutki, named from the Pacific Northwest (Paden and Tylutki 1968) and later synonymized with N. pouchetii (Paden 1972a), may be resurrected as the accepted name for Pacific Northwest material (Carbone, Agnello and Alvarado 2013).

Neournula pouchetii

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Octospora humosa (Fries) Dennis fruitbody 5–10 mm wide, shallowly cup-shaped, stalkless; inner surface smooth, shiny or dull, bright orange; outer surface whitish to pale brownish yellow; margin wavy, weakly toothed. MICROSCOPIC FEATURES: spores 19–22 × 11–13 μm, elliptic-cylindric, smooth, with 1 large oil drop and usually several smaller drops, uniseriate, hyaline; asci 220–250 × 19–20 μm; paraphyses cylindrical, septate, with curved clavate tips. OCCURRENCE: scattered or in groups on sandy soil among mosses; summer and fall; northeastern North America; infrequent. COMMENTS: Humaria humosa (Fries) Saccardo, Octospora leucolomoides (Rehm) K. B. Khare and V. P. Tewari, and Peziza humosa var. humosa Fries MACROSCOPIC FEATURES:

are synonyms. Octospora leucoloma Hedwig ex Gray (not illustrated) is an uncommon but widespread and att ractive yellowish orange to bright orange cup fungus that gets its name from a distinctive white, fi nely toothed margin. Its spores measure 18–24 × 11–15 μm, are broadly ellipticfusoid, smooth, and usually with 1 large oil droplet and several small ones. There are undoubtedly a dozen, and probably two dozen, species of Octospora in North America, often associated with burns and sometimes with mosses. The genus is badly in need of revision. Byssonectria tetraspora (Fuckel) Korf (= Octospora tetraspora [Fuckel] Korf) is an orange species that grows on the ground and is distinguished by 4-spored asci (spores 22–26 × 9.5–11.5 μm).

Octospora humosa

Octospor a

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Otidea alutacea (Persoon) Massee MACROSCOPIC FEATURES:

fruitbody a lopsided cup with litt le, if any, stalk; 2–4cm wide × 2– 6cm high; shorter side of cup split lengthwise or open and semierect, split edges usually closely inrolled, margin often irregularly wavy or contorted from growing in clusters, apex often truncate when growing erect, not ear-shaped; interior surface smooth, tan or light brown to grayish brown or brown; exterior surface often slightly scurfy, colored like the interior surface or a bit paler, yellowish in one variety; stalk absent or present as a narrowed, whitish, downy base; bases of several fruitbodies sometimes joined; flesh britt le, rather thick, yellowish. MICROSCOPIC FEATURES: spores 12–17.5 × 6– 9 μm, elliptic, smooth, typically bigutt ulate, uniseriate or obliquely arranged, hyaline to slightly yellowish; asci 250–300 × 8–15 μm, inamyloid; paraphyses slender, 3–4 μm wide, tips bent over, slightly lobed, septate and forked at the base, occasionally branched in the lower part one or two times, hyaline. OCCURRENCE: scattered to densely clustered in humus, usually under conifers (in hardwood forests in Europe); summer and fall; western North America and Europe. EDIBILIT Y: unknown, not recommended. COMMENTS: We fi nd it of note that in North America, Otidea alutacea is associated with conifers, while in Europe it is calcium-loving and associated with deciduous trees. It appears worth investigating whether or not we are dealing with the same species in North America as that which occurs in Europe. Otidea microspora (Kanouse) Harmaja (not illustrated), found on the west coast of North America, was considered by Kanouse (1949) to be a

small-spored variety (var. microspora) of Otidea alutacea. It has smaller spores (9–11 × 5– 6.5 μm) and a pale, clear yellow upper surface. Otidea alutacea and Otidea microspora both have truncate (rather than ear-shaped) fruitbodies, and paraphyses tips that are bent or hooked. Two other similar species that are truncate and have bent paraphyses are Otidea unicisa (Peck) Harmaja and Otidea cantharella var. minor Boudier (not illustrated). Otidea unicisa is an eastern species most commonly known as Otidea grandis sensu Kanouse and is frequently misidentified as being either Otidea leporina (Batsch) Fuckel or Otidea onotica (p. 200). Otidea cantharella var. minor is a small (1–1.5 cm high × 1–2.5 cm wide), pale yellow western species with spores measuring 10–12 × 6–7 μm. Otidea concinna (Persoon) Saccardo is an uncommon western hardwood associate with a lopsided cup slit laterally and somewhat inrolled, truncate and not earlike, strongly folded into convolutions, with a stout stalk-like base. Its interior and exterior surfaces are concolorous, clear lemon-yellow with a hint of red when fresh, becoming pinkish cinnamon with a hint of yellow when dried. Spores are 10–12(–13) × 5– 6 μm, elliptic, bigutt ulate, smooth, and yellowish. Paraphyses are fi liform, sometimes forked below, with apices hooked or bent, and with yellow encrustations. The spores of Otidea leporina var. minor Rehm are essentially the same size as O. concinna. The two are differentiated based on the fruitbody color, which is dull yellowish brown for Otidea leporina var. minor versus clear yellow with a hint of red for O. concinna. Otidea leporina (Batsch) Fuckel spores are larger (the same size as Otidea onotica, p. 200) and the fruitbody is a dull yellowish brown.

Pezizom ycetes

198

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Otidea alutacea

Ot i de a

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Otidea onotica (Persoon) Fuckel

Donk ey Ea rs, Or a nge Otidea

MACROSCOPIC FEATURES:

fruitbody 2– 6 cm wide, up to 10 cm high, irregularly cup-shaped and erect, slit down to the base on one side, resembling elongated ears or somewhat spoon-shaped; inner surface smooth, dull orange to orange-buff or ochreyellow, often rosy or pinkish tinged when fresh; outer surface slightly scurfy, colored like the inner surface but lacking the rosy or pinkish tinge; flesh britt le; stalk rudimentary, typically encrusted with debris, whitish or paler than the cup. MICROSCOPIC FEATURES: spores 10–13 × 5–8 μm, elliptic, smooth, hyaline, bigutt ulate; asci up to 250 × 10 μm, cylindric to subcylindric; paraphyses fi liform, strongly hooked at the tips. OCCURRENCE: solitary or in groups or clusters on soil and debris, or among mosses, in hardwoods, also reported in conifer woods in California; summer and fall, also winter and spring in California; widely distributed in northern North America, south along the Rocky Mountains and California; occasional. EDIBILIT Y: unknown, not recommended. COMMENTS: Aleuria abietina (Persoon) Gillet, Discina abietina (Persoon) Rehm, Otidea abietina (Persoon) Fuckel, Pseudotis abietina (Persoon) Boudier, and Scodellina onotica (Persoon) Gray are all synonyms. Otidea leporina (Batsch) Fuckel, a widespread conifer associate known commonly as hare’s ear and yellow rabbit ears, is nearly identical but has a pale yellow to brownish yellow cup and lacks a rosy or pinkish tinge on its inner surface. Otidea leporina spores are 12–15 × 6–8 μm, elliptic, smooth, bigutt ulate, uniseriate, and hyaline, and the paraphyses are fi liform and strongly hooked at the tips. Helvella leporina (Batsch) Franchi, L. Lami, and M. Marchett i, Scodellina leporina (Batsch) Gray, and Wynnella auricula (Schaeffer) Boudier are synonyms.

Otidea unicisa (Peck) Harmaja (synonyms Sowerbyella unicisa [Peck] J. Moravec and Otidea grandis sensu Kanouse) is commonly misidentified as Otidea leporina or O. onotica (Kanouse 1949; Moravec 1985; Harmaja 2009). It differs from O. onotica and O. leporina in its truncate (rather than ear-like) stature, and the outside surface is a bit darker and more reddish-brown when dried. All Otidea species except O. unicisa have smooth spores. Otidea unicisa is a common eastern species 1–2 cm high × 1–4 cm broad, with a fleshy-leathery truncate cup that is not ear-shaped and is turned up on one side and split to the base, on a short stalk. The margin of the split edges and top of fruitbody is inrolled when fresh, becoming deeply inrolled when dried. The fertile inner surface is pale vinaceous fawn, the outer surface brown to reddish brown, frequently with patches of orangered color when dry. The stalk is thick, up to 1 cm long, and yellowish. Otidea unicisa spores are 14– 17 × 6–7 μm, long-elliptic to slightly fusoid, biguttulate, with a smooth outer wall and an inner wall minutely roughened in age, and very careful examination under oil immersion reveals a delicate thin ornamentation of winding ridges and warts which form patches of reticulum in places and are coarsest at the spore ends (ornamentation part of a cyanophilic secondary wall) and inamyloid. Otidea tuomikoskii Harmaja is ear-shaped, clearly longer than wide, with a fi nely echinulatepapillate outer surface, which is observable without a hand lens. It appears to be associated mainly with spruce, has slightly brownish basal mycelium, and the spores are 8.5–11 × 5– 6 μm with encrustations that partly melt into amber drops with Melzer’s.

Pezizom ycetes

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Otidea onotica

Ot i de a

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Otidea smithii Kanouse fruitbody 3– 9 cm tall, typically taller than broad, an ear-shaped cup, split on one side, arising from a solid foot-like base of mycelium mixed with soil; margin even; inner surface brown to deep purple-brown when fresh, drying wood-brown to vinaceous-buff; exterior somewhat darker purple-brown shading to vinaceous toward the base; stalk subsessile, covered with offwhite hyphae. MICROSCOPIC FEATURES: spores 10–13.5(–15.5) × 6–7(–8)μm, narrowly elliptic, smooth, bigutt ulate, hyaline to faintly yellowish; asci 100–160 × 12– 14 μm, 8-spored, inamyloid; paraphyses with large hooked or bent tips, hooks sometimes ornamented by small irregular protuberances, hyaline. MACROSCOPIC FEATURES:

solitary to gregarious on exposed soil, duff, or moss under black cottonwood, Douglas fi r, and western hemlock; summer, fall, and early winter; western North America; uncommon. COMMENTS: Otidea smithii is distinguished by its large size, rabbit-ear-like shape, and deep vinaceous-brown coloration. Otidea rainierensis Kanouse is a rare Pacific Northwest species with large rabbit-ear-like fruitbodies that have a drab gray to vinaceous-buff interior and a cinnamon-buff to wood-brown exterior (Kanouse 1948). It is distinguished by straight clavate paraphyses, sometimes with globose heads up to 10 μm diameter. Its spores are indistinguishable from Otidea smithii. See also Midotis irregularis (p. 178). OCCURRENCE:

Otidea smithii

Pezizom ycetes

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Pachyella babingtonii (Berkeley and Broome) Boudier fruitbody up to 2 cm wide, shallowly cup-shaped, stalkless, broadly attached to the substrate; inner surface glabrous, uneven, and often wrinkled or fluted, typically shiny, pale yellow-brown to dull brown; outer surface pale grayish brown; flesh translucent, watery, and gelatinous. MICROSCOPIC FEATURES: spores 17–23 × 9–16 μm, broadly elliptic, thick-walled, smooth or very delicately punctate, usually bigutt ulate, hyaline; asci 250–325 × 15–20 μm, cylindric; paraphyses fi liform, with clavate tips. OCCURRENCE: scattered or in groups on wet decaying wood, cones, woody debris, or decaying stems of herbaceous plants; summer and fall, also winter in parts of the Pacific Northwest and California; widely distributed in North America; occasional. COMMENTS: Th is species was named in honor of Mr. Babington, collector of the holotype. Pachyella depressa (Phillips) Boudier and Peziza babingtonii Berkeley and Curtis are synonyms. Pachyella clypeata (Schweinitz) Le Gal is also similar but has a larger (0.5–4 cm broad) reddish brown fruitbody, MACROSCOPIC FEATURES:

later tinted olive then greenish-black and without luster. Pachyella clypeata spores are smooth with 1– 2 oil drops and measure 25–35 × 12–14 μm. Pachyella adnata (Berkeley and Curtis) Pfister is macroscopically nearly identical to P. clypeata but has smaller spores (18–20 × 10–12[–14] μm) that are elliptic, distinctly verrucose, thin-walled, hyaline, and with 1–2 oil drops. The epithet adnata means “broadly att ached,” referring to the broad att achment of the fruitbody to the substrate. Pachyella punctispora Pfister (not illustrated) is another nearly identical species with intermediate-sized spores (21–25 × 12–15 μm) stippled with tiny warts when mature. Miladina lecithina (Cooke) Svrček, found on water-soaked wood in water and streams in northern areas, is similar in size and shape to Pachyella babingtonii but is more brightly colored (pale yellow to orange, drying deep orange to blackish) and has elliptic, sometimes asymmetric, spores that measure 18–25 × 8–14 μm. The spores of Miladina lecithina have small, sometimes anastomosing, cyanophilic warts and usually numerous small oil droplets.

Pachyella babingtonii

Pa c h y e l l a

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Peziza ammophila Durieu and Montagne fruitbody up to 4 cm wide × 3 cm high, cup-shaped at fi rst but soon splitting in a star-shaped manner and becoming flattened on the sand surface; margin lobed and often split; inner surface smooth, dark brown; outer surface dark ochre to brown, usually coated with sand; stalk absent. MICROSCOPIC FEATURES: spores 14–18 × 8–10 μm, elliptic, smooth, pale brown; asci 200–275 × 12–15, strongly amyloid; paraphyses straight, slightly enlarged at the tips. OCCURRENCE: solitary, scattered, or in groups partially immersed in sand and anchored by a tuft of mycelial threads; widely distributed in North MACROSCOPIC FEATURES:

Sa nd-Lov ing Cup

America, particularly along coastal areas and along the Great Lakes region; fall and early winter; common but very seldom collected. COMMENTS: Geopyxis ammophila (Durieu and Montagne) Saccardo, Sarcosphaera ammophila (Durieu and Montagne) Moesz, and Tarzetta ammophila (Durieu and Léveillé) Theodorowicz are all synonyms. The epithet ammophilus means “sand loving.” When viewed from above, the fruitbodies resemble brown holes in the sand with lobed edges. Th is fungus has been reported from coastal eastern Canada, Florida, Texas, California, and the Great Lakes region.

Peziza ammophila

Pezizom ycetes

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Peziza arenaria Osbeck fruitbody cup-shaped to disc-shaped with litt le or no stalk, 2–5 cm broad; margin smooth to somewhat crenulate, slightly inrolled; inner surface violaceous to vinaceous; outer surface very slightly tomentose and slightly granular to nearly smooth, whitish to ochraceous. MICROSCOPIC FEATURES: spores 18–22 × 8–11 μm, smooth, with 2 large gutt ules; asci 8-spored; paraphyses erect with violaceous contents. OCCURRENCE: solitary to clustered on soil in forests; spring to early summer; eastern North America and Europe. EDIBILIT Y: not reported, difficult to identify correctly. COMMENTS: Peziza ampelina Quélet is a synonym, although Peziza ampelina Passerini is a different European species that is similar in appearance but exudes bluish latex when cut. Seaver (1978b) lists MACROSCOPIC FEATURES:

Peziza ampelina as a synonym of the very similar Peziza sublilacina Svrček (p. 219) (= Peziza violacea [Hedwig]). Peziza sublilacina is associated with burned ground, as is Peziza praetervisa (p. 216). Peziza arenaria was found to be closely related to P. praetervisa, Peziza howsei Roze and Boudier (not illustrated), and Peziza lobulata (Velenovský) Svrček (not illustrated) (Hansen, LoBuglio, and Pfister 2005). Peziza howsei differs from P. arenaria by having fi nely warted spores (17–22 × 8–11 μm). Peziza lobulata (= Peziza violacea Persoon ss. Dennis) is differentiated by having smaller, smooth, egutt ulate spores (13–15.5 × 7–8.5 μm) and curved paraphyses. Both Peziza howsei and P. lobulata grow on damp soil in woods and on wood in hardwood forests and, like P. arenaria, are not burn associates.

Peziza arenaria

Peziza

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Peziza arvernensis Boudier

Fa iry Tub

fruitbody a large cup, 2– 11 cm wide, often irregularly folded from the sides, expanding to saucer-shaped in age, undulating, without a stalk; margin often incurved; inner surface yellowish brown to chestnut-brown; outer surface fi nely hairy to nearly smooth, whitish to buff; flesh thin and fragile. MICROSCOPIC FEATURES: spores 16–19(–20) × 7.5– 10 μm, broadly elliptic, smooth, becoming fi nely punctate at maturity with irregularly spaced, distinctly rounded, and elongated warts, egutt ulate, hyaline; asci 180–240 × 11–13 μm, tips amyloid, 8– spored; paraphyses straight, septate, tips slightly clavate reaching 5–7 μm. OCCURRENCE: solitary to gregarious on soil (most often in association with leaves, small branches, or sawdust) or rott ing wood in both coniferous and hardwood (especially beech) forests; spring and summer; widely distributed in North America and Europe. EDIBILIT Y: unknown, not recommended. MACROSCOPIC FEATURES:

Peziza silvestris (Boudier) Saccardo, Aleuria silvestris Boudier, and Aleuria amplissima Boudier are all synonyms. There are many terrestrial brown Peziza species, as well as other somewhat similar brown cup fungi. Differentiating them requires microscopic study. The core Peziza species are all somewhat fragile and have asci tips that turn blue in Melzer’s. The spores are smooth until maturity. Therefore, correct identification requires a spore print to determine whether the specimen has spores that remain smooth or spores that develop ornamentation. Peziza varia (p. 221) is not easily distinguished from Peziza arvernensis. Peziza varia often has a lighter-colored, crenulate margin, the flesh is not especially fragile, and five layers are usually distinctly identifiable using a hand lens. See also Peziza domiciliana (p. 208). Tarzetta species are highly similar except that some have a distinct stalk and none have asci tips that are amyloid.

COMMENTS:

Peziza arvernensis

Pezizom ycetes

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Peziza badia Persoon

Pig’s Ea rs

fruitbody 2–10 cm wide, cup-shaped to shallowly cup-shaped, stalkless; inner surface pale to dark olive-brown, smooth; outer surface reddish brown, scurfy. MICROSCOPIC FEATURES: spores (15–)17–22 × 8– 10(–11) μm, elliptic, with a delicate irregular reticulum, often bigutt ulate, uniseriate, usually obliquely arranged, hyaline; asci 300–330 × 12–15 μm, amyloid at apex, cylindric, 8-spored; paraphyses cylindrical, septate, enlarged at the tips, yellowish. OCCURRENCE: in dense clusters or groups on sandy soil or sawdust, usually under conifers; late summer and fall, also winter and spring in California; widely distributed in North America; fairly common. MACROSCOPIC FEATURES:

unknown, not recommended. Galactinia badia (Persoon) Arnould, Helvella cochleata Bolton, Plicaria badia (Persoon) Fuckel, and Scodellina badia (Persoon) Gray are all synonyms. Peziza phyllogena (p. 215) is very similar, but has fi nely verrucose spores. Peziza alaskana Cash (not illustrated) is a small (0.3–1.8 cm wide) northern species closely related to P. badia. Peziza alaskana has a deep purplish black upper surface with a distinct margin, grows on naked calcareous soil of lakes and riverbeds, and has ornamented bigutt ulate spores (19–22 × 8.5–10 μm) with short ridges and warts up to 1.4 × 1 μm.

EDIBILIT Y:

COMMENTS:

Peziza badia

Peziza

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Peziza domiciliana Cooke

Domicile Cup Fu ngus

fruitbody consisting of an irregular cup, often with a short stalk, especially when young; cup up to 10 cm wide, concave and rounded at fi rst, soon becoming turned down to flattened, often retaining a depression at the center; margin typically irregular, wavy, and somewhat angular in age, sometimes splitt ing; inner surface smooth, whitish at fi rst, becoming dingy buff to tan or brown at maturity; outer surface smooth, whitish; flesh thin, britt le, often staining golden yellow when broken; stalk up to 1 cm long, irregular, whitish, becoming rudimentary in age. MICROSCOPIC FEATURES: spores (12–)14–16 × (6–) 8– 9(–10) μm, elliptic, smooth to slightly roughened, sometimes with 2 small oil droplets; asci 200–250 × 11–12 μm, cylindric, amyloid at apex, MACROSCOPIC FEATURES:

8-spored; paraphyses slender, septate, slightly enlarged at the tips. OCCURRENCE: solitary, in groups or clusters on a wide variety of domestic substrates where moisture is present, including shower stalls, carpets, clothing, cracks in concrete, plaster, greenhouses, and others too numerous to mention; year-round, widely distributed in North America; common. EDIBILIT Y: unknown, not recommended. COMMENTS: Th is fungus has a highly variable morphology and sometimes develops quite slowly. It is usually easy to identify because of its growth habit. However, it is not the only Peziza species known to grow in shower stalls, on wet carpets, etc. See also Peziza varia (p. 221).

Peziza domiciliana

Pezizom ycetes

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Peziza echinospora P. Karsten fruitbody 1–5(–10) cm broad, cup- to saucer-shaped, often irregularly compressed, lacking a stalk; inner surface dark brown to yellowish-brown; margin notched, inrolled when young, expanding in age; outer surface delicately to coarsely roughened or pimpled, lighter brown than the inner surface to almost whitish; flesh fragile, thin, brownish. MICROSCOPIC FEATURES: spores 14–17 × 6– 9 μm, elliptic, egutt ulate, with isolated regular warts or spines (0.5–1 μm high) that are often denser at the poles; asci 250–280 × 11–12 μm, tips amyloid, 8-spored; paraphyses cylindric-fi liform, straight, tips clavate (reaching 10 μm wide), with hyaline or pale yellowish vacuoles. OCCURRENCE: solitary or in fused clusters on burnt ground 10–120 weeks after a fi re; spring, summer, MACROSCOPIC FEATURES:

and fall; widespread in North America and Europe; not common. EDIBILIT Y: unknown, not recommended. COMMENTS: Aleuria umbrina Boudier and Peziza anthracophila Dennis are synonyms. It has also been spelled Peziza echinispora. Several other Peziza species—see comments under Peziza praetervisa (p. 216) and Peziza petersii (p. 213)—and three Plicaria species are associated with burnt ground. None of the three Plicaria species react to Melzer’s to give a blue ring at the tip of the ascus, and Plicaria spores are globose, not elliptic. The other Peziza species associated with burnt ground all have smooth to minutely warted spores. The spores of Peziza echinospora are unique in possessing distinct isolated spines and warts. The species is known only from burnt ground or charcoal.

Peziza echinospora

Peziza

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Peziza fimeti (Fuckel) E. C. Hansen fruitbody 0.5– 1.5(–2.5) cm wide, disc-shaped to shallowly cupshaped, stalkless or with a rudimentary stalk; inner surface concave to nearly flat, smooth; margin crenulate, pallid brown to yellowish brown; outer surface smooth to furfuraceous with pustules of globose cells, concolorous with the inner surface. MICROSCOPIC FEATURES: spores 18–22.5 × 9.5– 12.5 μm, elliptic, smooth, egutt ulate; asci up to 280 μm × 18 μm, cylindric, amyloid tips, 8-spored; paraphyses abundant, irregularly swollen up to 8 μm at the tips, tips straight or bent at a sharp angle, hyaline or with distinct yellow gutt ules. OCCURRENCE: on dung of ungulates, clayey soil, soil mixed with wood chips, or burnt ground; summer to fall; widespread in North America and Europe. EDIBILIT Y: unknown, not recommended. COMMENTS: Peziza bovina W. Phillips and Humaria fimeti Fuckel are synonyms. There are at least four named, and seven unnamed, small, shallowly cupshaped, smooth-spored Peziza species that are closely related. Peziza alcis Harmaja (not illusMACROSCOPIC FEATURES:

trated) is macroscopically similar to Peziza fimeti. It grows only on elk dung and has small spores (15– 16.5 × 7.5– 9 μm) and paraphyses that are bent at a sharp angle at the apex. Peziza nivalis (Heim and Remy) M. M. Moser (not illustrated) grows on soil and decaying grasses, often on burnt ground, near melting snow (Pfister 1992). Peziza nivalis can be separated from Peziza fimeti by habitat and DNA, although it is macroscopically and microscopically highly similar. Peziza ampliata Persoon (not illustrated) is microscopically indistinguishable from Peziza fimeti. It is recognized by its relatively thick flesh (0.1– 0.3 cm thick) and growth on hardwood trunks where it is sessile and broadly att ached or broadly substipitate. Peziza ampliata is also found on rich soil mixed with wood chips and on chalk soils. Peziza domiciliana (p. 208) is found on mortar, damp walls, and plaster. While Peziza domiciliana is usually 1–2 cm broad, it can be up to 5 cm broad. Peziza domiciliana is readily distinguished by its small bigutt ulate spores (14.5–15 × 7–8 μm) with fi ne regular warts.

Peziza fimeti

Pezizom ycetes

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Peziza gerardii Cooke (as gerardi) fruitbody 0.4– 0.5(–1.5) cm wide, initially closed and subglobose, becoming nearly discoid, margin thin, stalkless or with a rudimentary stalk; inner surface violaceous, or dark violet to dark violaceous brown; outer surface smooth to scurfy, concolorous with the inner surface or pale violaceous to bluish below. MICROSCOPIC FEATURES: spores 25–35 × 9–12 μm, narrowly fusoid, with low longitudinal ribs, with one to several oil drops, uniseriate or partially biseriate, hyaline; asci clavate, amyloid, 8-spored; paraphyses rather stout, clavate. OCCURRENCE: gregarious on naked, rich soil; summer to autumn; eastern North America and Europe. COMMENTS: Peziza ionella Quélet, Humarina gerardii (Cooke) Seaver, and Peziza violacea W. R. Gerard are synonyms. The violaceous color is common in the genus Peziza; see comments under Peziza praetervisa (p. 216). However, this very small thick-fleshed species is found on bare soil and is distinctive in North America. Smardaea planchonis (Dunal ex Boudier) Korf and W. Y. Yang MACROSCOPIC FEATURES:

(synonyms Peziza atroviolacea Delile and Marcelleina atroviolacea [Delile ex De Seynes] Brummelen) is genetically closely related and is another small (0.5–1 cm wide) thick-fleshed purplish species typically found on the ground, but is distinguished by smooth to slightly warted, 9–11 μm, round spores. Marcelleina persoonii (P. Crouan and H. Crouan) Brummelen (not illustrated), which occurs on calcareous soils and burnt ground, can only be distinguished from Smardaea planchonis by the presence of an almost complete network of ridges on its 10–12 μm globose spores. Smardaea planchonis and Marcelleina persoonii are also distinguished from Peziza species by nonamyloid asci. It is worth noting that the genus Peziza does not appear to be monophyletic, and Peziza gerardii is quite unusual for a Peziza: the amyloid reaction is not a distinct ring zone at the tip of the ascus, as in the core group of Peziza, but involves a bluing of nearly the entire ascus. Peziza azureoides Donadini is very similar in size to Peziza gerardii but is distinguished by its more intense purple coloration.

Peziza gerardii

Peziza

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Peziza ostracoderma Korf Anamorph: Chromelosporium fulvum (Link), Hennebert and Korf Peat Mould

fruitbody of apothecial state 0.5–2 cm in diameter, initially deeply cupshaped, flattening in age, often with an irregular shape; upper surface fulvous to dark brown; underside pruinose, more or less translucent, sometimes with a violaceous cast; interior tissues pallidtranslucent; anamorph colonies velvety to floccose, white or yellow, turning to cinnamon when actively growing and sporulating. MICROSCOPIC FEATURES: ascospores 11–13.5 × 6– 7.5 μm excluding warts, elliptical, at fi rst smooth, later warted and eventually with a delicate cyanophilic reticulum, bigutt ulate, hyaline; asci 200– 320 × 10–15 μm, operculate, amyloid at tip, 8-spored (rarely 4-spored), arising from croziers; conidia 5– 13 μm, globose, fulvous, single on 3 μm long × 1 μm thick stalks, produced densely and irregularly on the surface of the conidiogenous cell. OCCURRENCE: on burned ground, sterilized soil, mushroom compost; year-round; throughout MACROSCOPIC FEATURES:

North America, Europe, and Asia; common, especially in greenhouses. COMMENTS: The anamorph, Chromelosporium fulvum, has been known for more than 200 years and has gone by many different names, appearing most often in herbaria labeled as Botrytis species or Phymatotrichum species. Chromelosporium ollare (Person) Hennebert is a frequently encountered synonym (Hennebert and Korf 1975). Chromelosporium coerulescens (Bonorden) Hennebert is another commonly encountered Peziza anamorph. Chromelosporium coerulescens is found in large patches on decayed wood and mosses in forests. It has white, brown, crystal blue, and rose-violet growth phases. The small globose conidia are similar in size to the conidia of Chromelosporium fulvum and are produced on distinctive broom-like conidiophores. A teleomorphic state of Chromelosporium coerulescens is unknown.

Peziza ostracoderma

Pezizom ycetes

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Peziza petersii Berkeley fruitbody 2–5 cm broad, initially spherical, gradually expanding into a regular or contorted stalkless cup that typically has a scalloped margin; inner surface pale brown to dark brown or reddish brown, sometimes with a grayish to bluish tinge; outer surface nearly glabrous at fi rst, becoming scurfy to densely pustulate with bran-like particles when mature, brownish towards the margin, bluish to lead-gray towards the base; flesh thin, gray. MICROSCOPIC FEATURES: spores 10–12 × 5.5– 6 μm, fi nely warted, usually bigutt ulate, uniseriate; asci approximately 200 × 10 μm, amyloid, 8-spored; paraphyses clavate, 7–8 μm at tips. OCCURRENCE: solitary or more typically clustered to cespitose on burnt ground, charcoal, and stumps MACROSCOPIC FEATURES:

in hardwood forests with rich soil; summer and fall; widespread in North America and Europe. EDIBILIT Y: inedible. COMMENTS: Peziza limbricalis Cooke and Galactinia sarrazini Boudier are synonyms. There are at least nine Peziza species and three Plicaria species that are found on burned wood or burned ground. Several of these could sometimes be confused with Peziza petersii. The Plicaria species are differentiated by their globose spores and inamyloid asci. Peziza petersii does not have the violaceous colors of several of the burn-site Peziza species; see Peziza praetervisa (p. 216). The similarly colored Peziza echinospora (p. 209) has much larger spores (14– 17 × 7–8 μm) that are egutt ulate and fi nely warted. Peziza nivalis (R. Heim and L. Remy) M. M. Moser

Peziza petersii

Peziza

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(not illustrated) is a burnt-ground species that has still larger spores (18–21 × 9–10 μm) that are eguttulate. Peziza nivalis is an early spring fruiter associated with snow banks. Peziza ostracoderma (p. 212) is another burn species. Its spores measure 12–15 × 7–10 μm and are irregularly reticulate and bigutt ulate. Peziza proteana f. proteana (p. 217) is a burnt-ground species that has spores very similar to those of Peziza petersii, but it is differentiated by an upper surface that is white when

young and rosy, and pale lilac or slightly brownish in age. Peziza subisabellina (Le Gal) Blank, Häffner, and Hohmeyer (not illustrated) has a reddish to purplish brown upper surface (1–10 cm broad), grows on burnt ground, and has smooth spores (12–28 × 9–14 μm) with granulose content. Peziza sepiatra Cooke (not illustrated), another smoothspored burn-site species, is more blackish brown, measures less that 2 cm broad, and has egutt ulate spores (19–22 × 12–13 μm).

Pezizom ycetes

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Peziza phyllogena Cooke

Com mon Brow n Cup

fruitbody 3–15 cm wide, cup- to shallowly cup-shaped, stalkless; inner surface smooth, purple-brown to reddish brown or olive-brown; outer surface scurfy or granular, pale reddish brown to purple-brown; flesh thin, britt le, brownish; stalk absent. MICROSCOPIC FEATURES: spores 16–21 × 8– 10 μm, elliptic, fi nely verrucose, often with thickened polar walls, bigutt ulate, hyaline; asci about 140 × 12 μm, amyloid tips; paraphyses enlarged at the tips. OCCURRENCE: scattered, in groups, or in clusters on soil or decaying wood; spring and early summer, also winter and spring in California; widely distributed in North America; fairly common. EDIBILIT Y: reported as edible but not recommended. COMMENTS: As of 2012, Peziza badioconfusa Korf was listed as a synonym of Peziza phyllogena in Species fungorum, but it is considered a distinct species by MycoBank. The two are difficult to impossible to distinguish microscopically or macroscopically. Peziza badioconfusa is typically gregarious, up to 5 cm in diameter, broadly attached with white cottony mycelium, and is found from Quebec to British Columbia, Alaska, and the Northwest territories from May to September on well-rotted hardwood (alder, willow, and poplar). Peziza badia (p. 207) is very similar but its spores have a delicate irregular reticulum versus the fi nely warted spores of P. badioconfusa and P. phyllogena. Peziza brunneoatra Desmazières (not illustrated) is a widespread species measuring up to 2.5 cm broad. It is

brown to brownish black, sometimes with an olive tinge, has smooth elliptic spores (16–22 × 8–12 μm) that become roughened or partially reticulate when mature, with 1–2 oil droplets. The paraphyses have oily contents and are rather strongly widened in the upper part, reaching a width of 7–8 μm at the tips. The soft-fleshed, purplish brown Discinella boudieri (Quélet) Boudier, at 0.5–1.5 cm broad, might be mistaken for a small, somewhat planar Peziza until it is examined microscopically. Discinella boudieri is an inoperculate fungus with an amyloid pore, not an operculate fungus where the ascus is capped with a lid, and its spores measure 10– 15 × 4–5 μm.

Peziza

215

MACROSCOPIC FEATURES:

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Peziza phyllogena

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Peziza praetervisa Bresàdola

Pur ple Fa iry Cup

fruitbody 1–3(– 6) cm broad, saucer- to cup-shaped, lacking a stalk; inner surface smooth, light to dark violet, often with brownish tints; outer surface fi nely scurfy, pale purple; flesh thin, fragile, pinkish vinaceous. MICROSCOPIC FEATURES: spores 11–15 × 6–8 μm, elliptic, fi nely warted, bigutt ulate; asci 170–300 × 9– 12 μm, tips strongly amyloid, 8-spored; paraphyses slightly clavate, up to 7 μm at tips, tips bent slightly, fi lled with violet to violet-brown granules. OCCURRENCE: solitary to gregarious or fused in a cluster on burned soil, most often on remains of old campfi res in the fi rst twelve months after the fi re, sometimes on sandy paths or sawdust without any relation to burned wood; fall and spring; widespread in North America and Europe; common. EDIBILIT Y: unknown, not recommended. COMMENTS: Galactinia praetervisa (Bresàdola) Boudier is a synonym. Peziza praetervisa sensu Dennis = Peziza sublilacina Svrček. Peziza praetervisa is the common violet cup found at burn sites. Pfister (1982) noted that the description of Peziza violacea Persoon in Seaver (1978b) is actually PeMACROSCOPIC FEATURES:

ziza praetervisa and that most herbarium collections labeled Peziza violacea are actually P. praetervisa. It appears that Pfister thinks that the two can best be separated based on spore ornamentation and contents. Peziza praetervisa has lightly ornamented spores with 2 oil drops, while Peziza sublilacina (p. 219) has smooth spores without oil drops. Svrček (1976, 1977) erected Peziza subviolacea Svrček (= Peziza praetervisa) for the common North American and European species having fi nely warted spores. Peziza arenaria (p. 205) (as Peziza ampelina), Peziza howsei Roze and Boudier (not illustrated), and Peziza lobulata (Velenovský) Svrček (not illustrated) were found to be closely related to P. praetervisa (Hansen, LoBuglio, and Pfister 2005), but these three species are not associated with burn sites. Peziza howsei and P. arenaria both have larger spores (17–22 × 8–11 μm) than P. praetervisa. Peziza arenaria has smooth spores while P. howsei has fi nely warted spores. Peziza lobulata (= Peziza violacea Persoon ss. Dennis) is differentiated by smaller, smooth egutt ulate spores (13–15.5 × 7–8.5 μm) and having curved paraphyses.

Peziza praetervisa

Pezizom ycetes

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Peziza proteana f. proteana (Boudier) Seaver fruitbody 3– 6 cm broad, cup-shaped at fi rst then expanding, lacking a stalk; inner surface concave, becoming plane or convex, usually umbilicate, white when young, soon rosy to pale lilac or slightly brownish, darkening with age; outer surface often slightly scurfy, white, becoming faintly reddish or lilac; flesh thin, britt le. MICROSCOPIC FEATURES: spores 10–13 × 5– 7 μm, elliptic, usually with 2 small oil drops, smooth when young, becoming minutely roughened with warts or papillae when mature, uniseriate; asci 225–250 × 10 μm, amyloid tips, 8-spored; paraphyses slender, septate, clavate, reaching 7–8 μm at the tips, hyaline or with yellowish drops. OCCURRENCE: on old burnt places overrun with mosses; spring; eastern North America, occaMACROSCOPIC FEATURES:

sionally western North America and Europe; uncommon. EDIBILIT Y: edible if well cooked, but not recommended. COMMENTS: Aleuria proteana Boudier and Galactinia proteana (Boudier) Saccardo and P. Sydow are synonyms. Peziza proteana f. sparassoides (p. 218) is exceptionally similar microscopically, but very distinctive macroscopically. For a discussion of other burn-site Peziza species, see the microscopically similar Peziza praetervisa (p. 216) for violaceous species and Peziza petersii (p. 213) for brownish species.

Peziza proteana f. proteana

Peziza

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217

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Peziza proteana f. sparassoides (Boudier) Korf Fa lse Spa r assis, Bonfir e Cau liflow er

fruitbody 10–30 cm broad × up to 25 cm high; a cauliflower-like mass of contorted small cups lacking a stalk; inner surface smooth or wrinkled, individual cups distinctly contorted, whitish to tan, often tinged pinkish or lilac, aging brownish; outer surface concolorous or lighter than inner surface, often tinged lilac near the base; flesh thin, britt le. MICROSCOPIC FEATURES: spores 10–13 × 5– 7 μm, elliptic, bigutt ulate, smooth when immature, becoming minutely ornamented with fi ne, irregular, oblong warts at maturity, uniseriate, hyaline; asci 250–300 × 10–11 μm, amyloid tips, 8-spored; paraphyses clavate, 5–7 μm at the tips. MACROSCOPIC FEATURES:

on burnt ground; spring, also winter in California; widespread in North America and Europe; rare. EDIBILIT Y: reportedly choice if thoroughly cooked. COMMENTS: Aleuria proteana var. sparassoides Boudier, Peziza proteana f. campbellii (Saccardo) Korf, and Underwoodia campbellii Saccardo are synonyms. Th is distinctive species has no ascomycete look-alikes although it does bear a vague resemblance to Basidiomycete species in the genus Sparassis. OCCURRENCE:

Peziza proteana f. sparassoides

Pezizom ycetes

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Peziza sublilacina Svrček

V iolet Fa iry Cup

fruitbody 0.5–3 cm wide, cup- to saucer-shaped, stalk absent or rudimentary; inner surface smooth, lilac or violet to reddish violet; outer surface paler than the inner surface, grayish or light gray-violet to light brown, pruinose to mealy; flesh thin, fragile, pale violet. MICROSCOPIC FEATURES: spores 13–17 × 7–10 μm, elliptic, smooth, egutt ulate, uniseriate, hyaline; asci 300–340 × 10–11 μm, tips amyloid, 8-spored; paraphyses clavate (to 7 μm at tips), tips bent, fi lled with brown to purplish-brown granules. OCCURRENCE: scattered to gregarious on burnt soil; spring, summer, and fall, but mainly in the spring; widespread in North America and Europe but not frequently encountered. EDIBILIT Y: unknown, not recommended. COMMENTS: Peziza violacea Persoon, Aleuria violacea (Persoon) Gillet, and Galactinia violacea (Persoon) Svrček and Kubička are synonyms. While MACROSCOPIC FEATURES:

Peziza violacea is the name accepted by many authors, several species have carried the epithet Peziza violacea: Peziza violacea sensu Relhan = Ascobolus stercorarius (Bulliard) J. Schröter; Peziza violacea sensu auct. Dennis = Peziza pseudoviolacea Donadini; Peziza violacea W. R. Gerard = Peziza gerardii Cooke. See further comments under Peziza praetervisa (p. 216). Peziza griseorosea W. R. Gerard is a similar petite species (up to 2 cm broad) with prominently ornamented spores measuring 14–18 × 7–10 μm. Peziza azureoides Donadini is most easily differentiated from both P. violacea and P. praetervisa by its growth on unburned ground and its beautiful coloration. Peziza saccardoana Cooke (also spelled saccardiana) is distinguished from both P. violacea and P. praetervisa by its growth on unburned ground and by the reticulations that form a partial reticulum on its multigutt ulate spores (16–17 × 8– 9 μm).

Peziza sublilacina

Peziza

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Peziza succosa Berkeley

Y ellow Sa p Cup

fruitbody 2–5 cm wide, cup-shaped when young, becoming shallowly cupshaped to nearly flat in age, stalkless; inner surface smooth, pale grayish brown, often with an olivaceous tint; outer surface scurfy, whitish to pale grayish or yellowish, staining yellow when bruised; flesh thin, britt le, whitish, staining yellow from sap that exudes from broken flesh. MICROSCOPIC FEATURES: spores 16–22 × 8–12 μm, elliptic, coarsely ornamented with conspicuous verrucae and ridges, bigutt ulate, obliquely uniseriate, hyaline; asci 200–225 × 12–15 μm, cylindric to subcylindric, 8-spored, amyloid apex; paraphyses fi liform, septate, with enlarged tips. OCCURRENCE: solitary or in groups on the ground in woods, parklands, and along stream beds, often on bare soil; late spring, summer, MACROSCOPIC FEATURES:

and fall; widely distributed in North America; occasional. EDIBILIT Y: unknown, not recommended. COMMENTS: Galactinia succosa (Berkeley) Saccardo is a synonym. The name succosa means “full of sap.” Compare with Peziza vesiculosa (p. 223), which is similar but grows on manure piles or manured soil, has a pale yellow-brown fruitbody, and has thicker flesh that does not exude yellow-staining sap when broken. Peziza michelii (Boudier) Dennis exudes a watery juice when injured and the flesh at the injured spot fi nally becomes yellowish. The upper surface is reddish brown and may have a lilac tint, while the lower surface is lighter colored to yellowish brown and minutely scurfy, with or without a short stalk, and with spores that are smaller (16– 17.5 × 8 μm) and less ornamented than P. succosa.

Peziza succosa

Pezizom ycetes

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Peziza varia (Hedwig) Fries

R ecurv ed Cup, Spr ea ding Brow n Cup Fu ngus

fruitbody 2–15 cm broad, cup-shaped, expanding to nearly flat in age, stalk absent to rudimentary; inner surface light brown to golden brown or gray-brown, aging darker; margin often lighter colored, wavy and weakly notched; outer surface white to gray-brown and fi nely hairy, giving a scurfy appearance; flesh fairly fi rm, watery gray to light brown, often with a dark line in cross section, up to 5 layers visible with a hand lens. MICROSCOPIC FEATURES: spores 14.5–17.5 × 8– 10.5 μm, elliptic, smooth to slightly punctate, eguttulate, hyaline; asci 250–300 × 12–14 μm, amyloid tips, 8-spored; paraphyses septate, moniliform; lower and middle cells up to 20 μm wide (in a highMACROSCOPIC FEATURES:

moisture microenvironment); apical tips slender and slightly clavate. OCCURRENCE: solitary or fused together in clusters; in hardwood forests and gardens on buried or rott ing wood, on soil rich in humus, on sawdust, occasionally in wet basements on plaster or brick walls, in sand and gravel; late spring, summer, or fall; widespread in North America and Europe; common. EDIBILIT Y: unknown, not recommended. COMMENTS: Hansen, Læssøe, and Pfi ster (2002) placed Peziza cerea Sowerby ex Merat and Peziza micropus Persoon in synonymy, although Index fungorum as of 2013 still lists both as valid species. Hansen, Læssøe, and Pfister (2002) also found that

Peziza varia

Peziza

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Peziza repanda Persoon does not differ enough macroscopically, microscopically, or molecularly from P. varia to be separable as a species. The name Peziza repanda is particularly troublesome since there is no type collection, and various authors have interpreted the species differently, leading Hansen, Læssøe, and Pfister (2002) to reject the name P. repanda. Those who maintain Peziza repanda, Peziza micropus Persoon, and Peziza cerea Bulliard as separate species point to features such as the shape of the paraphyses, the consistently smooth spores, different habitats, different degree of development of a stalk, and different coloration of the upper surface and the underside. However, all of these features have proven to be plastic. Kullman (1995) studied the fruitbodies of one population on plaster over a six-week period. He observed that the presence or absence of a stalk and the color of the upper and lower surfaces varied during that time frame, with the features sometimes be-

ing characteristic of the classic Peziza cerea, other times resembling P. micropus, and at still other times matching the description of P. varia—thus supporting the DNA evidence for a single species, P. varia. We believe that the evidence for one variable species, rather than for four separate species, is very convincing, though as of this writing Peziza cerea, P. micropus, and P. repanda are all still in current use. Peziza cerea is used for species lacking inflated paraphyses and lacking five different layers in the fruitbody, and having an upper surface that is pale yellowish to saff ron. Peziza repanda is considered to be pale brownish on the upper surface and grayish on the lower surface. Peziza micropus is considered to be yellowish brown on the upper surface and greyish white underneath, with spores on the shorter end of the range, at 14–17 × 8–10 μm. Peziza ampliata Persoon: Fries (not illustrated) is very similar to Peziza micropus except for spores that are 17–20 × 9–11 μm.

Pezizom ycetes

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Peziza vesiculosa Bulliard

Bla dder Cup

fruitbody 2–8 cm wide × 1–5 cm high, cup-shaped and often distorted, margin frequently incurved, stalkless; inner surface pale to dark yellowish brown, nearly smooth; outer surface pale yellowish brown, scurfy; flesh brittle, brownish, forming tiny blisters in the center of the cup. MICROSCOPIC FEATURES: spores (18–)19–23(–24) × (10–)11–13(–14) μm, elliptic, smooth, egutt ulate, hyaline; asci 320–370 × 17–24 μm, cylindric, amyloid tips, 8-spored; paraphyses slender, septate, highly constricted at the septa, with slightly enlarged tips and granular contents. MACROSCOPIC FEATURES:

scattered or in clusters on manure piles and manured soil; late spring, summer, and fall, year-round in parts of the Pacific Northwest and California; widely distributed throughout North America; common. EDIBILIT Y: unknown, not recommended. COMMENTS: Galactinia vesiculosa (Bulliard) Le Gal is a synonym. Peziza succosa (p. 220) is similar, but it grows on non-manured soil and has thinner flesh that exudes yellow-staining sap when broken. OCCURRENCE:

Peziza vesiculosa

Peziza

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Peziza waltersii Seaver fruitbody 2–3 cm wide, subturbinate, constricted at the base into a short, stout stalk; inner surface slightly depressed at the center, dull yellow; outer surface nearly concolorous with the inner surface but darker dull yellow; flesh very soft . MICROSCOPIC FEATURES: spores average 20 × 10 μm, elliptic, coarsely tuberculate at maturity, uniseriate, hyaline; asci up to 340 × 15 μm, cylinMACROSCOPIC FEATURES:

dric, 8-spored; paraphyses enlarged to 5 μm at the apex, fi lled with granules. OCCURRENCE: on much-rotted wood of sugar maple and beech; early summer; northeastern North America; rare. EDIBILIT Y: unknown, not recommended. COMMENTS: Th is is a very distinctive but rare species.

Peziza waltersii

Pezizom ycetes

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Pithya cupressina (Fries) Fuckel fruitbody 1–2(–5) mm wide, at fi rst nearly spherical, becoming circular or elongated and irregular, flat or slightly concave at maturity; inner surface deep bright orange; outer surface lighter in color than the inner surface; stalk if present is short, stout, downy, with white mycelial threads that att ach the fruitbody to the substrate. MICROSCOPIC FEATURES: spores 9–12 μm, globose, smooth, granular within and usually containing one large oil droplet, uniseriate; asci 210–250 × MACROSCOPIC FEATURES:

12–15 μm, 8-spored; paraphyses branched in lower part, only slightly enlarged in upper part, reaching a width of 2–4 μm at the tips. OCCURRENCE: gregarious on recently killed foliage of various species of juniper, cyprus, hemlock, and some other conifers; spring and summer; North America and Europe; common. COMMENTS: Pithya cupressina is generally similar to some species of Hymenoscyphus and Bisporella. Pithya vulgaris Fuckel is larger and grows on wet, dead branch tips (with needles), twigs, and bark

Pithya cupressina

Pith ya

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of Abies and Sequoia, in montane areas, often near snow banks, in western North America and Europe. Pithya vulgaris is 0.1–1.5 cm wide, at fi rst somewhat cylindric, becoming cushion- to discshaped, flat to slightly convex, irregular in form when old. Its stalk is very short or absent. Its inner surface is bright orange and bald or slightly scurfy. Its outer surface is tinged with the color of the inner surface at the margin, and is paler and whiter towards the base, with white anchoring mycelium at the base. Its spores are 12–14 μm wide, round, at fi rst granular within, later with one large oil drop, uniseriate, and hyaline. Pseudopithyella minuscula (Boudier and Torrend) Seaver, another inopercu-

late species, is very similar in habitat and general appearance to species of Pithya. Pseudopithyella minuscula, found on decaying foliage of cedar and known from California, Bermuda, and Europe, is minute, being less than 2 mm in diameter, and is usually stalked, with a scarlet fertile upper surface and a whitish somewhat tomentose exterior, and has 15–17 × 10–11 μm spores each with 2 oil drops. Hymenoscyphus epiphyllus (p. 404) is also bright orange and sometimes is found on conifer foliage, but is quickly distinguished by its narrowly elliptic spores (16–20 × 4–5 μm) versus the round spores of Pithya species. See also comments under Gelatinodiscus flavidus (p. 380).

Pezizom ycetes

226

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Plectania melastoma (Sowerby) Fuckel fruitbody 1–3 cm wide, 1–2.5 cm high, rounded when young, soon becoming cup-shaped, stalkless or with a rudimentary stalk up to 1 cm long and thick; inner surface smooth, blackish brown to black; outer surface blackish brown to black, coated with matted hairs that are encrusted with reddish orange granules, especially near the margin; margin short-toothed and sometimes split, reddish orange; flesh gelatinous; stalk when present is att ached to the substrate by dense, wiry, black hairs. MICROSCOPIC FEATURES: spores (20–)21–24(–28) × 8–10(–12) μm, elliptic to fusiform, smooth, containing many small oil drops when young, egutt ulate at maturity, uniseriate, hyaline; asci 380–450 × 11–13 μm, long-cylindric, inamyloid, 8-spored; paraphyses fi liform, repeatedly forked, brown, tips slightly enlarged. OCCURRENCE: solitary, in groups, or in clusters on conifer debris; spring and early summer; widely distributed in North America; occasional. COMMENTS: Bulgaria melastoma (Sowerby) Seaver and Peziza melastoma Sowerby are synonyms. Th is is a very distinctive species. Some authors say that the spores are fi nely warted and gutt ulate at maturity, rather than smooth and egutt ulate. Similar species of Pseudoplectania have globose spores. See also Urnula padeniana (p. 265).

Jelly lik e Bl ack Ur n

MACROSCOPIC FEATURES:

Plecta ni a

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Plectania melastoma

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Plectania milleri Paden and Tylutki fruitbody 1–4 cm broad × 5–10 mm high, cup- to disc-shaped, sessile or with a short thick stalk; inner surface dark brown to purple-brown when fresh, drying black; margin stellate; underside hirsute, black, covered with appressed dark brown to black hyphae; flesh thin and gelatinous at maturity. MICROSCOPIC FEATURES: spores 21–31(–37) × (8–) 9–11 μm, elliptic, smooth, egutt ulate but fi lled with granular content, unripe spores globose to subelliptic, thick-walled with some oil drops, hyaline; asci 380– 600 × 12–15 μm, long-cylindric, operculum slightly eccentric, 8-spored; paraphyses 1.5–3 μm wide, thread-like, septate, occasionally branched, with numerous small oil drops. OCCURRENCE: gregarious to crowded; saprotrophic in plant debris, associated with mixed conifers; spring; western North America. COMMENTS: There are a great many small to medium (and in one case large), black, cup- to MACROSCOPIC FEATURES:

turban-shaped ascomycetes. The stellate margin of Plectania milleri is distinctive and its thin gelatinous flesh should remove any doubt about its identification (Paden and Tylutki 1969). Plectania melastoma (p. 227), a similarly sized black cup, is characterized by orange granules near the margin. Plectania melaena (Fries) Paden and Pseudoplectania nigrella (p. 236) have round spores. Plectania nannfeldtii (p. 229) has a long stalk, the exterior is less hairy, and the spores are slightly larger. Pseudosarcosoma latahense (Paden and Tylutki) M. Carbone, Agnello, and P. Alvarado (= Sarcosoma latahense Paden and Tylutki = Plectania latahense [Paden and Tylutki] M. Carbone) is much more gelatinized, although in dry weather it can lose most of its moisture, flatten out, and resemble a very large Plectania milleri; see Urnula padeniana (p. 265).

Plectania milleri

Pezizom ycetes

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Plectania nannfeldtii Korf fruitbody 0.5–3.0 cm wide × 1 cm high, shallowly cup-shaped, usually circular in outline from above to somewhat wavy, margin straight or slightly inrolled, with a long, usually buried, stalk; inner surface smooth, deep brownish black; outer surface clothed with minute, inconspicuous black hairs, wrinkled when dry, brownish black to black; stalk 1.5– 6 cm long × 1– 4 mm thick, nearly equal, widening somewhat toward the base, oval to flattened or fluted in cross section, clothed with minute black hairs, with black mycelium surrounding the base; flesh fibrous, tough. MICROSCOPIC FEATURES: spores 22–32(–37) × 10– 15 μm, elliptic, walls 1–2 μm thick (in 3 percent KOH the wall may rupture and may appear sculptured), usually with 2 small oil drops, one at each end, uniseriate, hyaline with yellow granular contents in Melzer’s reagent and in KOH; asci 280– 500 × 16–20 μm, narrowly clavate tapering gradually toward base, operculate, 8-spored, hyaline in MACROSCOPIC FEATURES:

KOH and in Melzer’s reagent; paraphyses abundant, 380–420 × 3.5–5(–12) μm, light brown in KOH, light reddish brown in Melzer’s reagent, septate with a hypha-like end cell culminating in a series of irregular swollen cells; hairs reaching 10 μm wide, septate, thin-walled, flexuous, pale brown. OCCURRENCE: solitary or in a cluster of two to several on conifer sticks, usually in or near melting snow, at 3,000–8,000 feet elevation; late spring to early summer; western North America; common. COMMENTS: Originally named Paxina nigrella Seaver in 1928, this species is currently placed in section Donadinia of Plectania. As a result of recent genetic work, Carbone, Agnello, and Alvarado (2013) proposed raising section Donadinia to species rank, giving this species a new name, Donadinia nigrella (Seaver) M. Carbone, Agnello, and P. Alvarado. The new name has not been accepted (as of July 2013). Helvella corium (p. 159) is larger, not as tough, and often shows ribs on the stalk.

Plectania nannfeldtii

Plecta ni a

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Plicaria carbonaria Fuckel fruitbody 0.5–3 cm wide, cup-shaped at fi rst, becoming flattened and shallowly cup-shaped at maturity; inner surface fi nely roughened to uneven and folded, dark brown to blackish, sometimes exuding droplets when injured; outer surface dark grayish brown, fi nely roughened; margin entire when young, becoming wavy to folded and sometimes ragged in age. MICROSCOPIC FEATURES: spores 11.5–13(–14) μm (excluding the spines), round, with coarse, blunt, wart-like spines, uniseriate, pale brown; asci 270 × 20 μm, cylindric, amyloid, 8-spored; paraphyMACROSCOPIC FEATURES:

ses cylindric, septate, with thickened, sometimes fi nely encrusted, rounded tips. OCCURRENCE: scattered or in groups in grass or on burned ground; spring and summer; widely distributed in North America; occasional. COMMENTS: Plicaria anthracina (Cooke) Boudier and Plicaria trachycarpa var. muricata Grélet are synonyms. The round spores with coarse, blunt, wart-like spines are most distinctive. Plicaria trachycarpa (Currey) Boudier (not illustrated) is nearly identical macroscopically, but its spores (10–11 μm) are ornamented with low rounded 1 μm warts.

Plicaria carbonaria

Pezizom ycetes

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Plicaria endocarpoides (Berkeley) Rifai fruitbody 1–7 cm wide, at fi rst spherical, expanding to shallowly cupshaped, occasionally incised and Otidea-like, soon becoming flattened or undulating, stalk rudimentary or absent; inner surface smooth or rough, sometimes wrinkled, dark reddish brown with an olive tinge to black; outer surface smooth or roughened, concolorous with or paler than the inner surface; flesh britt le, thin, when cut becoming yellowish and exuding a yellow juice. MICROSCOPIC FEATURES: spores 8–10 μm, globose, smooth, typically with several small oil droplets or one large central oil drop, rather thick-walled, biseriate becoming uniseriate, hyaline or slightly yellowish; asci 190–200(–350) × (10–)15–18 μm, cylindric or subcylindric in upper part, tapering in lower part into a stalk-like base, amyloid, 8-spored; paraphyses, slender, rather abruptly widened in upper part, densely granular within, reaching 4– 7 μm wide, showing a tendency to stick together in bundles, covered by an amorphous yellow-brown encrustation. MACROSCOPIC FEATURES:

solitary to gregarious on burnt ground, 20–130 weeks after fi re; all through the year, but usually spring; western North America, northern Europe, and New Zealand; common. EDIBILIT Y: unknown, not recommended. COMMENTS: Plicaria leiocarpa (Currey) Boudier is a synonym. Plicaria trachycarpa (Currey) Boudier (not illustrated) and Plicaria carbonaria (p. 230) are smaller (up to 2.5 cm across), blackish species with a roughened upper surface and warted (rather than smooth) spores. There are also similar burn-associated species in the genus Peziza, but these species are characterized by elliptic (rather than globose) spores; see Peziza petersii (p. 213). Ascobolus carbonarius (p. 129) is also similar, but is much smaller (0.2– 0.5 cm broad) and is characterized by fruitbodies that are yellowish green before turning brownish to blackish, an upper surface that is dotted with protruding asci, a fi nely mealy or scaly lower surface, a weakly scurfy-notched margin, and warted elliptic spores that are purplish to brownish purple at maturity. OCCURRENCE:

Plicaria endocarpoides

Plica r i a

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Pseudaleuria quinaultiana Lusk fruitbody 0.7–3.5 cm wide × 0.05– 0.2 cm tall; disc-shaped to slightly concave or repand, larger specimens often with a central depression, stalk short or absent; margin often inrolled; inner surface bright reddish orange, drying to a dull reddish brown; outer surface with long flexuous hairs forming a felty layer; flesh fi rm to rubbery, becoming corky when dry. MICROSCOPIC FEATURES: spores 15.5–21 × 7.5– 10.5 μm, elliptic, smooth, egutt ulate, uninucleate, hyaline; asci 200–300 × 8.0–12.5 μm, cylindrical, narrowing toward the base, 8-spored, operculate; paraphyses 240–320 × 2.0–3.5 μm, slightly to greatly and abruptly enlarged in upper part, apices 4.5–11.5 × 4.5– 9.5 μm, straight, septate, rarely branching or branching in top half, containing carotenoid granules; external hairs arise from exterMACROSCOPIC FEATURES:

nal thick-walled cells, nonfasciculate, long (to at least 1,000 μm), and interwoven. OCCURRENCE: scattered or in clusters, near the butt end of fallen conifers, on wood or soil, primarily in old-growth forest with sitka spruce, western hemlock, and Douglas fi r; spring; known only from Washington and Oregon; uncommon. COMMENTS: The single most distinctive feature of this species is the long, thin-walled hairs that are hyaline when fresh (Lusk 1987). Sarcoscypha coccinea (p. 246) is similar but is redder, usually cupshaped, with less conspicuous external hairs and a stalk up to 4 cm long. Aleuria aurantia (p. 126) has thinner, more britt le flesh, and is orange rather than red-orange. It tends to be cup-shaped rather than saucer-shaped, occurs in large groups, has fi ne downy external hairs, and fruits in the fall.

Pseudaleuria quinaultiana

Pezizom ycetes

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Pseudombrophila cervaria (W. Phillips) Brummelen fruitbody 1.5–3 mm wide, cup-shaped with a prominent margin and a rudimentary stalk; inner surface chestnut-brown to purplish brown; margin and outer surface clothed in very pale adnate hairs, darker than the inner surface. MICROSCOPIC FEATURES: spores 14.5–18.5 × 7– 9.5 μm, elliptic, smooth, lacking oil drops but may have de Bary bubbles; asci 150–175 × 10–12.5 μm, abruptly attenuated at apex, inamyloid, 8-spored; paraphyses fi liform, not enlarged at apex. OCCURRENCE: on dung or manured soil; northern portions of North America; common. MACROSCOPIC FEATURES:

COMMENTS: Pseudombrophila theioleuca Rolland (not illustrated) has a pale yellowish gray hymenium, a prominent dark brown margin, an exterior surface with appressed brown hair, and somewhat smaller spores (12.5–16.5 × 6.5–8.5 μm). Pseudombrophila hepatica (Batsch) Brummelen (not illustrated) has a reddish brown to pale purplish brown upper surface, with a darker lower surface that is covered with pale brownish to almost hyaline hairs, and much larger spores (22–35 × 10–13 μm). See Pseudombrophila merdaria (p. 234) for species with spores less than 12 μm long.

Pseudombrophila cervaria

Pseu dom brophi l a

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Pseudombrophila merdaria (Fries) Brummelen MACROSCOPIC FEATURES:

fruitbody 1–5 mm broad, turbinate or disc-shaped to shallowly cup-shaped, with a prominent margin, sessile; inner surface smooth, color variable from pinkish gray to pale reddish brown to brick-red; outer surface concolorous with the inner surface or darker brownish, with network of separate, adnate, flexuous, pale brownish hairs. MICROSCOPIC FEATURES: spores 10.5–13 × 6–8.5 μm, elliptic, smooth, egutt ulate, 8-spored, uniseriate, hyaline and then yellowish; asci inamyloid; paraphyses fi liform but abruptly enlarged to 4 μm at the apex. OCCURRENCE: on dung or herbaceous stems; northeastern North America and northern Europe; common. COMMENTS: Pseudombrophila deerrata (P. Karsten) Seaver (not illustrated) is considered a synonym of Pseudombrophila merdaria by some authors, although Index fungorum and MycoBank recognize both as valid species. Pseudombrophila deerrata can be larger, with longer spores and narrower paraphyses. It has a disc-shaped or shallow cup-shaped fruitbody up to 1 cm across, colored gray-violet to reddish on upper surface, and similarly colored on lower surface but dotted with darker tuft s of

fi ne hairs that appear scale-like, has a short stalk, and grows on rott ing remains of plants. Pseudombrophila deerrata spores are 12–15 × 6–8 μm, elliptic, and hyaline, appearing smooth on superficial examination but revealing an obliquely transverse sinuous structure under oil immersion when stained with cotton blue. The asci are 8-spored, 150–160 × 10–11 μm, and inamyloid. Its paraphyses are uniformly about 2 μm thick, fi liform, branched, and septate, some with dark granular contents. Pseudombrophila porcina (Svrček and Kubička) Brummelen (not illustrated) is similarly colored, grows on dung, and is 0.2–1 cm broad, with distinctive bundles of septate rough hairs on the outside of the apothecium. The spore size is in the same range as Pseudombrophila merdaria. Iodophanus carneus (Fries) Korf is whitish to pale orange or pale pink, also found on dung, but has exterior hairs that are very fi ne and soon disappear. Its spores (17–22 × 10–12 μm) are delicately verrucose and egutt ulate. Dung-inhabiting Cheilymenia species have rooting, sparse to distinctive exterior hairs and fruitbodies that are whitish, yellow, orange, or reddish. Their spores have a loose, irregularly verrucose or echinulate episporium.

Pezizom ycetes

234

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Pseudombrophila merdaria

Pseu dom brophi l a

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Pseudoplectania nigrella (Persoon) Fuckel fruitbody 3–16 mm high, 6–25 mm wide, cup-shaped to shallowly cupshaped, stalkless; inner surface blackish brown to black, smooth; outer surface coated with a dense layer of short, matted hairs, black to blackish brown. MICROSCOPIC FEATURES: spores 10–14 μm, globose or nearly so, with a gelatinous sheath, smooth, often with 1 large oil drop and many small droplets, hyaline or pale brown; asci 250–320 × 12–17 μm, long-cylindric, inamyloid, 8-spored; paraphyses fi li form, multiseptate, tips straight and sometimes forked. OCCURRENCE: scattered or in groups on mosscovered decaying conifer wood; spring and summer; widely distributed in North America; occasional. MACROSCOPIC FEATURES:

H a iry Bl ack Cup

Plectania nigrella (Persoon) P. Karsten is a synonym. Similar species of Plectania have elliptic to fusiform spores. Paden (1983) proposed a transfer of Pseudoplectania nigrella to Plectania nigrella, but this was not widely accepted. Smardaea planchonis (Dunal ex Boudier) Korf and W. Y. Zhuang (synonym Marcelleina atroviolacea [Delile ex DeSeynes] Brummelen) grows on sandy soil, is cup- to disc-shaped and dark purple, with a scalloped margin and a minutely warted exterior surface. Its spores are 10–12 μm, round or nearly round, and smooth or very minutely sculptured, with minute warts at maturity. They are initially hyaline, becoming pale purplish, and contain one to several oil drops. COMMENTS:

Pseudoplectania nigrella

Pezizom ycetes

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Pseudoplectania vogesiaca (Persoon) Seaver fruitbody shallowly cup-shaped to plane with a short stalk, 2–3(–8) cm broad; upper surface smooth, dark olive-brown, blackening in age; margin often wavy, externally very sparingly tomentose, strongly wrinkled when dry, darker than upper surface; flesh subcartilaginous; stalk variable, up to 2–3 cm × 3 mm, attached at base by dense, coarse, dark brown hairs. MICROSCOPIC FEATURES: spores 10–17 μm in diameter, round, smooth, hyaline, with 1 large droplet; asci 200–320 × 13–18 μm, with very long stem-like base, inamyloid, 8-spored; paraphyses fi liform with few septa, tips simple to spirally curved, sometimes forked, up to 3–4 μm wide, with dark contents; hairs up to 120 × 7 μm, cylindric, black-brown, smooth, with few septa, thin-walled, tips blunt, some cells widened and bulging, base with bristlelike outgrowths. OCCURRENCE: single to gregarious on decaying wood in coniferous woods, especially among MACROSCOPIC FEATURES:

Sphagnum; late winter or early spring; widespread in North America; common. COMMENTS: Features include a dark brown to blackish cup sparingly clothed with straight or slightly wavy hairs, a stalk of variable length but never long like Plectania nannfeldtii (p. 229), growth on wood under conifers, smooth round spores, and paraphyses that are hooked or coiled at tips. Pseudoplectania nigrella (p. 236) is typically smaller, has no stalk, and is densely clothed externally with coiled hairs. Most Plectania species have fi nely warted elliptical spores. Plectania melastoma (p. 227) and Plectania milleri (p. 228) have no distinct stalk. Plectania melastoma usually has orange granules on the margin. Plectania melaena (Fries) Paden = Pseudoplectania melaena (Persoon) Saccardo is recognized by both MycoBank and Species Fungorum as distinct from Pseudoplectania vogesiaca, but we have been unable to ascertain any significant macroscopic or microscopic differences.

Pseudoplectania vogesiaca

Pseu dopl ecta ni a

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Pulvinula carbonaria (Fuckel) Boudier fruitbody 0.1– 0.4 cm broad, disc- or cup-shaped, lacking a stalk; inner surface roughened by protruding paraphyses, pale orange; outer surface a litt le paler than the inner surface. MICROSCOPIC FEATURES: spores 15–18 μm, round, smooth, containing 1 oil droplet that almost fi lls the spore, hyaline; asci 225 × 18–20 μm, cylindric or nearly so; paraphyses fi liform, reaching 3–4 μm wide at the tips, significantly longer than the young asci and strongly curved or hooked. MACROSCOPIC FEATURES:

gregarious or crowded on mosscovered soil in places that have been burned; known from western North America and Europe. COMMENTS: Lamprospora carbonaria (Fuckel) Seaver is a synonym. Another small burnassociated species is Pyronema omphalodes (p. 240), which is composed of confluent masses of minute orange cushion-shaped fruitbodies surrounded by cobwebby mycelium. See also Anthracobia melaloma (p. 128) for other small orange species found on burnt ground. OCCURRENCE:

Pulvinula carbonaria

Pezizom ycetes

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Pulvinula convexella (P. Karsten) Pfister fruitbody 0.1–1.2 cm wide, top- to cushion-shaped or disc-shaped, upper surface often drawn together into a short stalk, orange to pink or scarlet; underside lighter than the upper surface or whitish pink. MICROSCOPIC FEATURES: spores 13–18 μm wide, globose, with one large and many smaller droplets, smooth, hyaline; asci 200–250 × 18 μm, 8-spored, inamyloid; paraphyses fi liform, cylindrical, tips bent over like a cane, aseptate, forked. OCCURRENCE: solitary to gregarious in sandy soil, along paths, along streams, on burnt ground, or among mosses; western North America and Europe. COMMENTS: Pulvinula constellatio (Berkeley and Broome) Rehm is given by Index fungorum as a synonym, but Hansen and Knudsen (2000) consider Pulvinula constellatio to be a pink to scarlet species found near burns and on pure limestone. MycoBank gives Pulvinula constellatio as MACROSCOPIC FEATURES:

a synonym for Barlaea constellatio (Berkeley and Broome) Saccardo (not illustrated). Pulvinula convexella is the orange to reddish species found in sandy soil and along streams. Pulvinula archeri (Berkeley) Rifai (= Peziza gemmea W. Phillips = Lamprospora pyrophila Snyder) is a western North American and European species characterized by an orange convex disc (less than or equal to 0.7 cm wide), with much smaller globose spores that measure 9–11.5 μm and generally have 1 large oil droplet. Pulvinula laeterubra (Rehm) Pfi ster has a salmon-red (sometimes yellow) disc with smaller globose, egutt ulate ascospores (11–13.8 μm). It is frequently misidentified as being either Pulvinula constellatio or P. convexella. Coprotus species grow on dung, are yellow to orange with fi liform paraphyses, and have elliptic spores. Coprobia species are also orange and grow on dung, but have a furfuraceous exterior with stout paraphyses.

Pulvinula convexella

P u lv i n u l a

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Pyronema omphalodes (Bulliard) Fuckel MACROSCOPIC FEATURES:

fruitbody 1–2 mm wide, usually convex, lacking a stalk, growing on a dense white mycelial mat; individual fruitbodies becoming confluent and forming congested masses several cm in diameter; upper surface usually pinkish orange, sometimes slightly yellowish to bright orange or reddish orange, purplish tinged or whitish; without a raised margin; underside bald. MICROSCOPIC FEATURES: spores 10–15 × 5–8.5 μm, elliptic, smooth, egutt ulate, uniseriate, hyaline; asci 150 × 10–14 μm, cylindric or nearly so, inamyloid, 8-spored; paraphyses very slightly enlarged above, 4–7 μm at their apices, fi lled with orange granules. OCCURRENCE: on burned places within 1–2 weeks of a burn, in greenhouses on steamed soil or soil that has been sterilized by dry heat; year-round; widely distributed; common. COMMENTS: Pyronema confluens (Persoon) Tulasne and C. Tulasne is a synonym. Moore and Korf (1963) point out that nearly all small, pink discomycetes have at one time or another been identified as this species. Pyronema domesticum (Sowerby) Saccardo and Iodophanus carneus (Persoon) Korf are two common pinkish species that are frequently misidentified as Pyronema omphalodes. The genus Pyronema itself has been made up of a combination of Anthracobia, Ascophanus, Cheily-

menia, Octospora, Neottiella, Psilopezia, Pulvinula, and Pyronemalla. In North America, Pyronema omphalodes has not been found in greenhouses although it has been found in greenhouses in Japan. In North America, the greenhouse invader is Pyronema domesticum, originally described from plaster in a newly plastered house (since new plaster generates heat sufficient to “sterilize” the plaster). It has also turned up in fi replace areas. Pyronema domesticum is highly variable in color, often ranging from red to pink, salmon, ivory, and pure white within a single collection. It is characterized by defi nite hairs on the exterior surface of the fruitbodies, particularly near the margin. They are usually scattered and do not fuse. Pyronema omphalodes and P. domesticum are indistinguishable microscopically, but in nutrient-rich culture only P. domesticum forms sclerotia. Iodophanus carneus (Persoon) Korf is distinguished by amyloid asci that turn blue all over, a feature that separates the genus Iodophanus from almost all other ascomycetes. Iodophanus carneus, which grows in burn areas or on dung, wet rags, etc., is turbinate- to discshaped, measures 0.1– 0.3 cm broad, is whitish, yellowish, or pinkish, has a pubescent exterior that later becomes bald, and has delicately warted or wrinkled spores that measure 17–22 × 10–12 μm.

Pezizom ycetes

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Pyronema omphalodes

Py rone m a

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Rhizina undulata Fries

Pine Fir e Cushion, Crustlik e Cup

fruitbody 2–10 cm wide, disc-shaped to convex, sometimes fused to adjacent fruitbodies, attached to the ground by many short, root-like hairs, fragile and spongy when young, tough when old; upper surface wavy and wrinkled, more or less red-brown, drying blackish brown, usually with a distinct white to yellowbrown margin; underside white to pale brown, fi nely pubescent; stalkless, with numerous white to yellow or gray rhizoid-like projections up to 3 cm long that extend into the ground and are ingrown with debris. MICROSCOPIC FEATURES: spores 30–45 × 8–15 μm, narrowly spindle-shaped, rough, apiculate, multigutt ulate, hyaline; asci 300–400 × 15–20 μm, MACROSCOPIC FEATURES:

8-spored; paraphyses 5.5–8 μm at apex, contents fi nely granular with brown-encrusted pigment; setae thick-walled, aseptate, 6.5–8 μm thick at the base widening to 8.5–11 μm at the tip. OCCURRENCE: solitary to numerous on soil or woody debris, frequently in recently burned areas or clear-cuts under conifers where it is a conifer seed pathogen; spring through fall; across North America but with a northerly distribution, also in northern or montane Europe and Asia; occasional. COMMENTS: Rhizina infl ata (Schaeffer) P. Karsten and Rhizina zonata Berkeley are synonyms. The indeterminate growth and rhizoid-like projections on the underside of this species are a unique combination of features.

Rhizina undulata

Pezizom ycetes

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Rhodoscypha ovilla (Peck) Dissing and Sivertsen fruitbody 1–5 mm broad, disc- to cup-shaped with a prominent shaggy margin, sessile or with a short, fat stalk; inner surface pink, rarely whitish; outer surface whitish with flexuous obtuse hairs that have two-layered walls. MICROSCOPIC FEATURES: spores 33–46 × 10– 16.5 μm, fusoid, at fi rst with 2 oil drops, when maMACROSCOPIC FEATURES:

ture with many small oil drops, smooth or with irregular small warts. OCCURRENCE: on soil or plant debris; autumn. COMMENTS: Leucoscypha ovilla (Peck) Harmaja and Neottiella ovilla (Peck) Saccardo are synonyms.

Rhodoscypha ovilla

R hodosc y ph a

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Rhodotarzetta rosea (Rea) Dissing and Sivertsen fruitbody 3–20 mm wide × 3–15 mm high, nearly round when young, soon becoming cup-shaped, often distorted, usually stalkless or with a rudimentary stalk; inner surface smooth, bright pink to pinkish red; outer surface smooth to slightly roughened or minutely downy, colored like the inner surface; margin entire, sometimes splitt ing; stalk when present is whitish. MICROSCOPIC FEATURES: spores 16–20 × 9–11 μm, broadly elliptic, smooth, hyaline, with 2 large oil MACROSCOPIC FEATURES:

Pink Bur n Cup

drops; asci about 250 × 13 μm; paraphyses cylindric, very slightly enlarged at the tips, with pinkish granular contents. OCCURRENCE: in groups or dense clusters on burned ground, often among mosses; spring and summer; widely distributed in North America; occasional and not often collected. COMMENTS: Tarzett a rosea (Rea) Dennis and Pustularia rosea Rea are synonyms.

Rhodotarzetta rosea

Pezizom ycetes

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Sarcoscypha austriaca (O. Beck ex Saccardo) Boudier fruitbody 2–7 cm wide, rounded and deeply cup-shaped at fi rst, becoming shallowly cup-shaped to ear-shaped or irregular in outline at maturity; inner surface smooth to somewhat wrinkled, shiny or dull, scarlet to red; outer surface granulose to scurfy, whitish to pale ochraceous or pinkish red; margin strongly incurved when immature; flesh thin but not fragile, pinkish; stalk typically short and thick, colored like the outer surface of the cup, sometimes rudimentary or absent. MICROSCOPIC FEATURES: spores 20–38 × 9–16 μm, elongated-elliptic, mostly truncate, smooth, hyaline, multigutt ulate, less than 3 μm in diameter at one or both ends; asci long-cylindrical, gradually tapered at the base, 375–485 × 18–19 μm; paraphyses fi liform, slightly clavate at the tips. MACROSCOPIC FEATURES:

Sca r let Cup

OCCURRENCE: solitary or more often in groups or compressed clusters on fallen or buried branches and twigs in damp hardwoods and mixed woods; spring; northeastern and north-central North America; common. EDIBILIT Y: nonpoisonous, but not recommended. COMMENTS: Th is is one of the fi rst mushrooms to appear in spring, often collected while snow is still on the ground. Sarcoscypha coccinea (p. 246) is nearly identical, but its spores are not mostly truncate. Sarcoscypha dudleyi (p. 247) is also nearly identical, but its spores are not truncate and they have large polar oil drops that measure 5–7.5 μm. Sarcoscypha occidentalis (p. 248) is similar, but has smaller cups and relatively long, slender stalks.

Sarcoscypha austriaca

Sarcoscypha

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Sarcoscypha coccinea (Jacquin: Fries) Lambotte Sca r let Cup, Sca r let Elf Cup

fruitbody up to 4 cm wide, rounded and deeply cup-shaped at fi rst, becoming shallowly cup-shaped to ear-shaped or somewhat irregular in outline at maturity, toughfleshy to rubbery when fresh; inner surface smooth, scarlet to red, fading to reddish orange; outer surface tomentose, at times hoary, colored with an opaque tint of red to reddish orange; margin strongly incurved when immature; stalk typically short and stout, 1.5–4 mm thick, tomentose or nearly smooth. MICROSCOPIC FEATURES: spores 25–40 × 9–14 μm, oblong-elliptic or narrowly elliptic, not truncate, with a mucilaginous sheath, smooth, hyaline, with numerous oil droplets less than 3 μm in diameter aggregating toward the polar ends; asci 375–475 × MACROSCOPIC FEATURES:

13–15 μm, containing numerous small oil droplets when young, inamyloid, 8-spored; paraphyses cylindrical, slender, septate in the lower portion, branched, red granular contents turn green in Melzer’s. OCCURRENCE: solitary or in groups on partially buried wood in conifer or hardwoods; fall, winter, and spring; British Columbia, Vancouver Island, California, Oregon, and Washington; occasional to fairly common, common in Europe. COMMENTS: Peziza coccinea Jacquin is a synonym. Sarcoscypha austriaca (p. 245) is nearly identical, but its fruitbody has a whitish outer surface, it occurs in northeastern and north-central North America, and its spores are mostly truncate. See also Sarcoscypha dudleyi (p. 247).

Sarcoscypha coccinea

Pezizom ycetes

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Sarcoscypha dudleyi (Peck) Baral fruitbody up to 6.5 cm wide, rounded and deeply cup-shaped at fi rst, becoming shallowly cup- to ear-shaped or kidneyshaped at maturity, tough-fleshy to rubbery when fresh; inner surface smooth, scarlet to red, fading to reddish orange; outer surface scurfy or with scattered light brown pustules, colored with an opaque tint of red to reddish orange; margin strongly incurved and crenulate when young but recurved at maturity and becoming laciniate. MICROSCOPIC FEATURES: spores 20–38 × 9–15.5 μm, predominantly oblong-elliptic, not truncate, biguttulate, typically with 1 large oil drop that measures 5–7.5 μm in diameter, and several smaller oil dropMACROSCOPIC FEATURES:

lets at each pole, usually encased in a mucilaginous sheath, hyaline; asci 260–400 × 13–18 μm, longcylindrical, gradually tapered basally. OCCURRENCE: solitary, scattered, or in groups on moss-covered decaying hardwood; fall, winter, and spring; central and eastern United States; uncommon. COMMENTS: Peziza dudleyi Peck is a synonym. Peck named this species in honor of W. R. Dudley, who collected it in 1894. Sarcoscypha austriaca (p. 245) is nearly identical but has mostly truncate spores that lack large oil drops. Sarcoscypha coccinea (p. 246) is also nearly identical but has spores that lack large oil drops.

Sarcoscypha dudleyi

Sarcoscypha

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Sarcoscypha occidentalis (Sowerby) Saccardo fruitbody 1.2–4.5 cm high × 6–16 mm wide, cup- to shallowly cupshaped, with a distinct stalk; inner surface bright red, smooth; outer surface pinkish red to pinkish orange, lacking white hairs; stalk 1–3.5 cm long, 1.5–3 mm thick, nearly equal, smooth, white. MICROSCOPIC FEATURES: spores (17–)20–22 × 10– 12 μm, elliptic, smooth, with 2 or more oil drops, uniseriate, hyaline; asci (200–)240–280(–320) × 12–15 μm, cylindric, 8-spored; paraphyses fi liform, with granular contents. MACROSCOPIC FEATURES:

Sta lk ed Sca r let Cup

scattered, in groups, or in clusters on decaying hardwood branches; late spring, summer, and early fall; eastern and central North America; occasional. COMMENTS: Geopyxis occidentalis (Schweinitz) Morgan and Plectania occidentalis (Schweinitz) Seaver are synonyms. Microstoma floccosum (p. 176) and Microstoma protractum (p. 177) are both similar, but their outer surfaces and margins are clothed in long white hairs. OCCURRENCE:

Sarcoscypha occidentalis

Pezizom ycetes

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Sarcosoma globosum (Schmidel) Caspary fruitbody 4.5–12 cm wide, irregularly rounded, dense and heavy; outer surface incurved over the inner surface, deeply wrinkled and folded, roughened, fibrous-tough and elastic, pinkish brown to yellowish brown or dingy brown when dry, becoming dark reddish brown to blackish brown when wet, often mott led and streaked as the fruitbody dries out; inner surface located in a depression on the upper portion of the outer surface, smooth to fi nely cracked, blackish brown; interior gelatinous with a copious amount of water when fresh.

MACROSCOPIC FEATURES:

MICROSCOPIC FEATURES: spores (20–)23–30 × 7– 12 μm, elliptic, smooth, uniseriate, hyaline; asci cylindric to subcylindric; paraphyses slender, adhering together in masses, dark brown, with enlarged tips. OCCURRENCE: solitary, scattered, or in groups on needle debris or among mosses, in conifer woods; spring and early summer; northeastern North America; rare. COMMENTS: Bulgaria globosa Schmidel is a synonym.

Sarcosoma globosum

Sa rcosom a

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Sarcosphaera coronaria (Jacquin ex Cooke) Boudier Tu lip Cup, Pink Crow n, Crow n Fu ngus

fruitbody 3–20 cm across, at fi rst a barely hypogeous and somewhat flattened ball, the wall splitt ing downward from a central soft spot or rounded hole to form pointed segments or rays that frame a deep cup; stalk absent or a short, thick plug or sometimes long and thick; inner surface smooth or breaking into fi ne scales, initially whitish to creamy, soon pinkish to violet or brownish violet; underside scurfy, usually dirt-encrusted, whitish to grayish; flesh of the stalk and cup rather thick and britt le. MICROSCOPIC FEATURES: spores 14–22 × 7–10 μm, elliptic with blunt or truncate ends, smooth or very slightly roughened, with 1–3 (usually 2) oil droplets, hyaline; asci 300–360 × 10–13 μm, cylindrical, with amyloid tips, 8-spored; paraphyses cylindric, septate, constricted at the septa, some forked at the MACROSCOPIC FEATURES:

base, tips slightly clavate and widening to 5– 6(– 7) μm at the apex, with brownish sap. OCCURRENCE: single to gregarious, usually exposed or partly exposed when mature; under hardwoods and conifers; spring, summer, and fall; North America and Europe; common. EDIBILIT Y: Orson Miller once said it tastes like pink pencil erasers and causes indigestion. It is a hyperaccumulator of arsenic. COMMENTS: Sarcosphaera crassa (Santi) Pouzar and Sarcosphaera eximia (Durieu and Léveillé) Maire are synonyms. Th is species is common and usually quickly recognized. However, sometimes one encounters specimens with extraordinarily long stalks, with the cup color bleached by the sun and the star-like rays folded back, creating a mushroom that can keep even an expert guessing for a while.

Sarcosphaera coronaria

Pezizom ycetes

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Scutellinia crucipila (Cooke and W. Phillips) J. Moravec fruitbody 0.1– 0.4 cm broad, at fi rst subglobose and nearly closed, later saucer- to disc-shaped, broadly att ached to the substrate; upper surface orange to orange-red, fading slightly on drying; margin with inconspicuous stiff hairs; underside with same hairs as on margin plus fi ne stellately branched hairs, paler than the upper surface; hairs light brownish. MICROSCOPIC FEATURES: spores 15–20 × 7– 10 μm, elliptic, fi nely warted, egutt ulate, sometimes with pseudoapiculi, exospore layer separating when heated in lactic acid; asci 180–220 μm long, 8-spored, cylindrical, inamyloid; paraphyses 2–3 μm wide, subclavate (4–10 μm at tips); marginal hairs 100–200(–500) × 8–13 μm, unbranched, 2–5 septate, tapering to a point or rounded, not branched at the base; underside has same hairs as marginal hairs intermixed with 75–150 μm hairs each with 2–5 arms. OCCURRENCE: scattered to crowded on damp bare ground, among mosses, or on bare ground unMACROSCOPIC FEATURES:

der grasses and other herbaceous plants; mostly in summer; reported from the Pacific Northwest, New York, and Europe; uncommon. COMMENTS: Cheilymenia crucipila (Cooke and W. Phillips) Le Gal is a synonym. Cheilymenia stercorea (Persoon) Boudier is similar but grows on dung, is smaller, and has smooth spores. Its bristle-like marginal hairs have branched “roots.” The “root” of the hairs on Scutellinia crucipila is not branched. Scutellinia crucipila is unlike other Scutellinia species in that its spores are egutt ulate and have an exosporium that separates when heated in lactic acid. These features are typical of Cheilymenia species, and DNA places this species in Cheilymenia (Hansen et al. 2013). Scutellinia blumenaviensis Hennings has remarkably ornamented spores (19–23 × 11–14 μm) with a complete and regular reticulum measuring 1–2 μm high and reaching 4 μm high at the poles and thus resembling apiculi.

Scutellinia crucipila

Scutellinia

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Scutellinia erinaceus (Schweinitz) Kuntze fruitbody 1.5–5 mm wide, shallowly cup-shaped, stalkless; inner surface pale orange-yellow when young, becoming dull orange at maturity, smooth, shiny or dull; outer surface and margin dull brownish orange, covered with long, stiff, brown hairs. MICROSCOPIC FEATURES: spores (14–)17–21(–26) × 9–11.5(–15) μm, elliptic, smooth, though cytoplasmic granules can appear as very fi ne markings when viewed under oil immersion, hyaline; asci 270–325 × 12–15 μm, cylindrical, inamyloid, 8-spored; paraphyses fi liform, with clavate tips containing orange granules; rooting hairs (with rootlets at base) abundant, (100–)400– 600(–700) μm long × 20–40 μm wide at base, inconspicuously septate, dark brown, pointed at tips or rarely blunt or with a constriction. OCCURRENCE: in groups or clusters on decaying hardwood; summer and fall, also winter and spring in parts of the Pacific Northwest and California; widely distributed in North America; occasional to fairly common. COMMENTS: Patella setosa sensu Seaver is a synonym. Scutellinia erinaceus is smaller than most MACROSCOPIC FEATURES:

Or a nge Ey el ash Cup

other members of this genus and is characterized by a dull orange coloration with overtones of yellow or brown, rather than orange with overtones of red as found in Scutellinia scutellata (p. 253) and many other species in the genus. Scutellinia setosa (Nees: Fries) Kuntze, with smooth spores (17.5– 20.5 × 10–12.5 μm) and a very small size (1–2.5 mm diameter), is very similar to S. erinaceus but is orange-red. Scutellinia nigrohirtula (Svrček) Le Gal (not illustrated), found on wood and soil in eastern North America, is rare, and larger (4–10 mm). It also has overtones of yellow to brown, and the marginal rooting hairs are grayish black, 3–5-septate, and measure 150–420 × 12–28 μm. The spores are 19.5–26.5 × 14–16.5 μm and have broad, irregular, often confluent warts 0.4 μm high. Tricharina gilva (p. 259) is similarly colored, the margin is covered with brown hairs that are thinner and shorter (150– 190 × 3–5 μm), and the spores are smooth (14–19 × 9–10 μm). It grows on moss-covered burnt ground. Some Cheilymenia species are somewhat similarly colored, but their habitat is dung.

Scutellinia erinaceus

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Scutellinia scutellata (Linnaeus) Lambotte

Ey elash Cup, Ey elash Fu ngus

fruitbody up to 2 cm wide, somewhat rounded at fi rst, soon expanding into a shallow cup or nearly flattened disc; inner surface smooth, shiny, bright red to orange-red or orange; outer surface pale brown, more or less covered with stiff, dark brown to black hairs that form a fringe on the cup margin, with short, pale hyphae-like hairs between the marginal hairs; flesh thin, britt le, reddish. MICROSCOPIC FEATURES: spores 15–23 × 10– 13.5(–17) μm, elliptic, minutely verrucose, lacking a reticulum, multigutt ulate when young, hyaline; asci 240–320 × 15–20 μm, cylindrical, inamyloid, 8-spored; paraphyses with clavate tips; marginal rooting hairs 360–1600 × 22–47 μm, multiseptate, with a multifurcate base, brown. OCCURRENCE: scattered, in groups, or in clusters on decaying wood or on damp soil in woodlands; summer and fall, also winter and spring in parts of the Pacific Northwest, California, and the southern parts of its range; widely distributed in North America; common. COMMENTS: Peziza scutellata Linnaeus: Fries is a synonym. A white variety, Scutellinia scutellata var. leucothecia Le Gal, has been found in Switzerland and eastern Canada. Of the many orange to red Scutellinia species, Scutellinia scutellata is both the most widespread and one of the most distinctive, with its long eyelash-like hairs and orange-red coloration. Scutellinia pennsylvanica (Seaver) Denison, reported from eastern North America on rotting wood and sometimes soil, is very similar but can be pale salmon-orange in low-light conditions, rather than its typical orange-red color. Scutellinia pennsylvanica lacks the superficial hairs on the underside of the disc found in S. scutellata and has distinctive spores that measure (14–)16–19(–21) × (8–)10–13(–14) μm. The spores are strongly ornamented by coarse verrucae that anastomose to form a massive and irregular reticulum similar to

the spores found in the genus Melastiza. Scutellinia verrucipolaris Denison (not illustrated) forms 2– 8 mm wide scarlet, shallow cups on soil and sometimes on wood with much shorter (100–500 μm) septate rooting hairs on the margin. The unusually narrow bigutt ulate spores measure (17–)19– 22(–24) × (7.5–)9–11 μm and are ornamented with discrete rounded or subtruncate verrucae that are larger and more numerous toward the poles, which clearly separates Scutellinia verrucipolaris from all other members of the genus. Scutellinia umbrorum (Fries) Lambotte is a very common species, with a general preference for soil, that forms medium to large, 8–20 mm wide, orange to brownish-red shallow cups. The unusually broad spores measure 18– 23 × 12–15 μm and are ornamented with evenly dis-

Scutellinia

253

MACROSCOPIC FEATURES:

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Scutellinia scutellata

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tributed, moderate to large rounded verrucae. The marginal short rooting hairs measure 200–720 × 25–30 μm and are multiseptate and brown. Superficial hairs, if present, are very short and stout. Scutellinia pilatii (Velenovský) Svrček is bright red, 3–12 mm broad, grows on wood, has pale brownish multiseptate marginal hairs (600–1600 × 25– 43 μm) interspersed with short, pale hyphae-like hairs, and has spores 21–27.5 × 13.5–16.5 μm with

faint isolated warts and ridges 0.3– 0.6 μm broad × 0.3 μm high. Scutellinia erinaceus (p. 252) measures 2–4 mm, grows on wood, and has a pale yellow-orange to dull orange inner cup surface and smooth spores. Scutellinia kerguelensis (Berkeley) Kuntze has stiff, thick-walled brown marginal hairs measuring 300–500 μm long, one-third of them forked at the base, that are 4– 6 septate. Its 19–25 × 12–15.5 μm spores are very delicately ornamented.

Pezizom ycetes

254

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Sowerbyella radiculata (Fries) Nannfeldt fruitbody 1–5 cm broad, disc- to cup-shaped with a rooting stalk; upper surface often wrinkled near the center, greenish yellow; underside paler; stalk prominent, 1–4 cm long, furrowed or depressed, strongly encrusted with needles. MICROSCOPIC FEATURES: spores 12.5–15 × 7–8.5 μm, elliptic, bigutt ulate, with elongated warts that MACROSCOPIC FEATURES:

form a dense, incomplete reticulum; paraphyses curved. OCCURRENCE: solitary to clustered in coniferous forests; summer to late fall; eastern North America and Europe. COMMENTS: Th is distinctive species prefers rich, calcareous soil.

Sowerbyella radiculata

Sower byella

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Sowerbyella rhenana (Fuckel) Moravec Fa lse Or a nge Peel, Sta lk ed Or a nge Peel Fu ngus

fruitbody 1–4 cm wide, cup-shaped with a distinct stalk; inner surface bright orange to yellow-orange, smooth; outer surface pale yellow or whitish, slightly scurfy; margin typically incurved, becoming wavy and torn at maturity; base of the cup compressed and folded; stalk up to 2 cm long, ribbed, covered with a dense white mycelium. MICROSCOPIC FEATURES: spores 18–26 × 9–12 μm, elliptic, smooth at fi rst, soon covered by a coarse reticulum, uniseriate, hyaline; asci 300–350 × 15– MACROSCOPIC FEATURES:

17 μm, cylindric to clavate; paraphyses enlarged at the tips, fi lled with orange granules, fleetingly green with Melzer’s. OCCURRENCE: typically clustered or in groups on soil or among mosses in woodlands, usually associated with conifers; summer and fall, also late fall and early winter in California; widely distributed in North America; uncommon. COMMENTS: Aleuria rhenana Fuckel is a synonym. Aleuria aurantia (p. 126) is very similar but has a rudimentary stalk or is stalkless.

Sowerbyella rhenana

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Sphaerosporella brunnea (Albertini and Schweinitz) Svrček and Kubička fruitbody 2– 6 mm wide, shallowly cup-shaped with a raised margin; inner surface smooth to slightly roughened, reddish brown at fi rst, becoming dark brown at maturity; outer surface brownish to dark reddish brown, fi nely roughened with tuft s of matted fi ne hairs; margin thick, raised, roughened with densely compacted fi ne hairs. MICROSCOPIC FEATURES: spores (13–)14–15(–18) μm, globose, smooth, with 1 large oil drop, uniseriate, hyaline; asci about 200 × 16–20 μm, cylindric, 8-spored; paraphyses cylindric, septate, with clavate tips. OCCURRENCE: scattered, in groups, or in clusters on bare sandy soil near water oaks, or on burnt ground and among mosses; spring and summer; reported from Alabama to Massachusett s MACROSCOPIC FEATURES:

west to Iowa, also west of the Great Plains; rare to uncommon. COMMENTS: Ciliaria brunnea (Albertini and Schweinitz) Boudier, Durandiomyces phillipsii (Massee) Seaver, Daleomyces phillipsii (Massee) Seaver, and Trichophaea brunnea (Albertini and Schweinitz) L. R. Batra are all synonyms. The combination of globose, smooth, hyaline spores and growth on sandy soil is very unusual for a species of cup fungi. Peziza ammophila (p. 204) also grows in sandy habitats and forms dark brown cups with lobed edges. Before being unearthed, only the uppermost portion of the cup rim is exposed, causing the fungus to resemble a brown hole in the sand. Sphaerosporella hinnulea (Berkeley and Broome) Rifai is used by some authors for the form of S. brunnea that grows on clayey soil.

Sphaerosporella brunnea

Sph a erospor ell a

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Tarzetta catinus (Holmskjold) Korf and J. K. Rogers fruitbody 1–5 cm broad, deeply cup-shaped to urn-shaped, sometimes splitting into lobes or becoming expanded when old, with a distinct stalk; margin dentate to slightly notched; inner surface cream to hazelnut-brown; outer surface downy-felty to fi nely granular; concolorous with the inner surface or a bit lighter; stalk very short to somewhat long, buried, gradually widening into the cup, round or with longitudinal furrows; flesh thin, britt le. MICROSCOPIC FEATURES: spores 18–24 × 10– 13.5 μm, elliptic, smooth, bigutt ulate, hyaline; asci 280–350 × 16–20 μm, inamyloid, 8-spored; paraphyses slender, septate, forked toward the base, tips not clavate but often lobed, 3–4 μm wide. OCCURRENCE: several to gregarious on the ground under hardwoods and conifers; spring, summer, and fall during wet weather; North America and Europe; fairly common. COMMENTS: Pustularia catinus (Holmskjold) Fuckel is a synonym. Tarzetta species have also been placed in Geopyxis and Peziza. Tarzetta cupuMACROSCOPIC FEATURES:

laris (Linnaeus) Svrček, the elf cup, is found on soil and in burns in much of North America and is distinguished by its generally smaller size (1–2.5 cm high and wide), shorter stalk, pale yellowish to yellowish gray color, and the absence of lobed tips on the paraphyses. The spores (18–22 × 12–15 μm) are similar to Tarzetta catinus, but have a lower length to width ratio. Tarzetta bronca (Peck) Korf and J. K. Rogers, which is widely distributed in hardwood and conifer forests, is very similar to T. catinus in fruitbody size and color, and the shape of the paraphyses. Tarzetta bronca is differentiated by the absence of a distinct stalk and in having about half of the spores measuring 12–14 × 8– 9 μm with 2 small oil droplets, and the other half measuring 20–24 × 12–13 μm with 1–2 large oil drops. Geopyxis vulcanalis (p. 143) is yellowish, has egutt ulate spores, and is much smaller (0.5–1.2 cm diameter). Peziza species are very similar, generally stalkless, and characterized by having tips of the asci that are amyloid, while Tarzetta asci are inamyloid.

Tarzetta catinus

Pezizom ycetes

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Tricharina gilva (Boudier ex Cooke) Eckblad fruitbody 0.2– 0.5 cm broad, shallowly cup- to saucer-shaped with a flat disc, stalkless; margin with short brown hairs; inner surface yellow-orange when young, paling to light ocher when old; underside weakly furfuraceous or with stiff, light reddish brown hairs, colored like inner surface. MICROSCOPIC FEATURES: spores 14–19 × 8–10 μm, elliptic, smooth, egutt ulate, hyaline; asci 170– 250 × 14–16 μm, inamyloid, 8-spored; paraphyses slender, slightly club-shaped, about 4 μm wide at the tips; hairs 150–200 × 3–5 μm, 10 μm at base, smooth, thin-walled, sparingly septate, tapered to a rounded point. OCCURRENCE: cespitose to clustered on bare and mossy burnt ground; spring and summer; Pacific MACROSCOPIC FEATURES:

Northwest, northeastern North America, and Europe; not common. COMMENTS: Trichophaea gilva (Boudier ex Cooke) Gamundi is a synonym. Over the years this species has also been placed in six other genera. There are probably about a half dozen species of Tricharina in North America but they are not well studied. In general, members of this genus can be found on soil, limestone, or burnt ground. They have a pinkish buff, grayish white, yellowish, or ochraceous upper surface, an underside that is concolorous with the upper surface or brownish and covered with pale brown to reddish brown blunt or pointed septate hairs, and smooth elliptic spores that lack gutt ules and are hyaline. Trichophaea species are found on soil, burnt ground, or plant de-

Tricharina gilva

Tr ich a r ina

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bris. They have a grayish white or grayish brown upper surface, an underside covered in stiff, hyaline, or brownish hairs, and smooth to ornamented gutt ulate spores of various shapes. Cheilymenia species also have hairs on the margin and underside of the fruitbody and grow on dung, rich soil, plant debris, and wood. They have a whitish to yellow-orange or reddish upper surface, marginal underside hairs that are hyaline, yellowish, or brownish, and elliptic egutt ulate spores that have an irregularly verrucose or echinulate episporium.

Scutellinia species are found on soil, limestone, or wood. They have a whitish to ochraceous, orange, or red upper surface, an underside that is concolorous with the upper surface or brownish with stellate hairs on the margin and underside, and gutt ulate spores of many shapes with delicate or coarse ornamentation of warts, spines, or reticulum. If the fruitbody is buried or partially buried, see Geopora sepulta (Fries) Korf and Burdsall in the comments under Geopora arenosa (p. 89).

Pezizom ycetes

260

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Trichophaea hemisphaerioides (Mouton) Graddon fruitbody 0.5–1.5 cm wide, at fi rst hemispherical-bladder-like, then remaining cup-shaped for a long time and later becoming flat, stalk absent; margin with brown hairs and always turned upward; inner surface white to gray-whitish, sometimes with a bluish tint; underside light brown, clothed with short, stiff, light brown hairs. MICROSCOPIC FEATURES: spores 13–18 × (5–)7– 8 μm, narrowly elliptic, some fi nely punctate and rough, with a small oil droplet near each end, hyaline; asci 175–200 × 7–8 μm, 8-spored, inamyloid; paraphyses slender, widening up to 6 μm at the tips, forked at base, septate, tips slightly thickened; hairs 200–400 × 6–17 μm, dark brown, thickMACROSCOPIC FEATURES:

walled (1–2 μm), multiseptate, narrowing to a point. OCCURRENCE: solitary to gregarious on burned ground 20–50 weeks after burning, commonly among the burned-ground moss (Funaria stage of succession) and on burned pieces of wood; found in the Pacific Northwest and Europe and probably in eastern North America, distribution limits yet to be determined; spring, summer, and fall. COMMENTS: Humaria hemisphaerioides (Mouton) Eckblad is a synonym. In general appearance, Trichophaea hemisphaerioides looks very much like a miniature Humaria hemisphaerica (p. 172). The strikingly similar Trichophaea woolhopeia (Cooke and W. Phillips) Boudier has much larger spores

Trichophaea hemisphaerioides

Tr ichoph a e a

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(20–22.5 × 16–18 μm) with 1 large oil drop, often with de Bary bubbles. Trichophaea abundans (P. Karsten) Boudier, which appears very soon after a fi re, is smaller (0.2– 0.5 cm broad) and the hymenium is at fi rst dark grayish brown, maturing to pale grayish brown. Its marginal hairs are almost hyaline, although the underside hairs are brown. The spores are 13–14 × 6–7 μm and biguttulate. Trichophaea gregaria (Rehm) Boudier (not illustrated) is very similar to T. hemisphaerioides but grows solitary to very gregarious on bare sandy ground on the edges of paths and less often on burned places. Its spores are 23–25.5 × 9–10 μm and narrowly elliptic to somewhat fusiform (longer but also much narrower than T. woolhopeia). The margin and underside are both thickly covered with short dark brown hairs and dark brown cells, giving it a mott led appearance. Trichophaeopsis bicuspis (Boudier) Korf and Erb has very small turbinate- to disc-shaped apothecia (0.2– 0.3 cm broad) and a whitish upper surface, a brown-

ish underside with long, stiff, brownish hairs, and is found on soil and twigs under poplars in summer to autumn. Its spores measure 15–17 × 9– 10 μm and contain de Bary bubbles, and it has a peculiar excipulum of vertical rows of cells broader than tall and long thick-walled setae usually with a pronounced basal prong. Trichophaea contradicta (Seaver) H. J. Larsen (= Patella contradicta Seaver), known from burnt ground in New York and British Columbia, has a Dichobotrys Hennebert anamorph (conidia 8– 9 μm, subglobose to globose with 1 μm pedicel), is gregarious, brown, and 1–2.5 mm wide with short brown marginal hairs and sparse long brown hairs on the underside. Its spores measure 12–18 × 8.5–11 μm and are 1–2 gutt ulate, smooth, and broadly elliptical. Its inamyloid asci measure 150–170 × 11–14 μm. Paraphyses are 150–170 μm, clavate to spathulate, 5–8 μm at apex, and the apical cell is brownish with small oil droplets.

Pezizom ycetes

262

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Underwoodia columnaris Peck

Fluted-Sta lk ed Fu ngus

fruitbody 4–12 cm high, 1–3 cm thick, cylindric to fusiform, tapered upward to a rounded tip, with longitudinal grooves or wrinkles, lacking a distinct cap and stalk, cream to tan when young, becoming pale brown in age; interior chambered. MICROSCOPIC FEATURES: spores 25–27 × 12–14 μm, elliptic, coarsely verrucose, uniseriate, hyaline; asci up to 350 × 20 μm, cylindric; paraphyses fi liform, with curved and slightly enlarged tips. OCCURRENCE: in clusters with fused bases or sometimes scattered on the ground in hardwoods; late spring and summer; widely distributed in eastern North America and the Midwest; uncommon to rare. COMMENTS: Helvella columnaris (Peck) Eckblad is a synonym. Th is species is sometimes misidentified as a parasitized mushroom. MACROSCOPIC FEATURES:

Underwoodia columnaris

U n derwoodi a

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Urnula craterium (Schweinitz) Fries fruitbody 4.5–10.5 cm high × 2–7 cm wide, deeply cup-shaped with a distinct stalk; inner surface blackish brown to black, smooth; outer surface black to brownish black, densely matted and wooly, becoming brownish gray, tough and leathery in age; margin toothed and irregularly torn; stalk 2–4 cm long × 5–10 mm thick, tapered downward, black to brownish black. MICROSCOPIC FEATURES: spores 25–35 × 12–14 μm, broadly elliptic, smooth, uniseriate, hyaline; asci up to 600 × 15–17 μm, cylindric; paraphyses fi liform, with slightly enlarged tips, pale brown. MACROSCOPIC FEATURES:

Dev il’s Ur n

in groups or clusters on decaying wood or on the ground attached to buried wood; spring; eastern and midwestern North America; fairly common. COMMENTS: Geopyxis craterium (Schweinitz) Rehm, Peziza craterium Schweinitz, and Sarcoscypha craterium (Schweinitz) Bánhegyi are synonyms. Strumella coryneoidea is probably the anamorph (Davison 1950). Th is is one of the earliest mushrooms to appear in spring. OCCURRENCE:

Urnula craterium

Pezizom ycetes

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Urnula padeniana M. Carbone, Agnello, A. D. Parker, and P. Alvarado Sta rv ing M a n’s Licor ice, Gi a nt Gel Cup

fruitbody 5–10(–20) cm wide × 3–10(–15) cm high, top-shaped, with a raised, often lobed margin, becoming like a thick pancake in age, narrowed below to form a very short, thick stalk; upper surface more or less flat to moderately concave, black or occasionally dark brown; underside often deeply wrinkled, especially in the lower part, fi nely velvety, covered with an extensive dark tomentum with heavily pigmented hairs, often roughened with dark granules, dark gray to black; stalk short, gelatinous, often deeply wrinkled or ribbed or with large pockets; flesh a

MACROSCOPIC FEATURES:

thick, watery-gelatinous mass, collapsed and rubbery in age, clear, gray to black or brownish. MICROSCOPIC FEATURES: spores 23–34 × 10–14 μm, elliptic to somewhat allantoid, smooth, with 1– 3 oil droplets, hyaline; asci 500–570 × 12–15 μm, suboperculate, long-cylindric, gradually narrowing basally, curved at base, inamyloid, 8-spored but sometimes only 6–7 spores mature; paraphyses approximately 3 μm wide in lower part, 3–5 μm wide at the tips, closely septate, often slightly constricted at the septa, branched, anastomosing, embedded in a dark green, amorphous material; hy-

Urnula padeniana

Urnula

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phae of the tomentum irregular, up to 6 μm wide, dark green or brown, often roughened with dark brown granules. OCCURRENCE: solitary to gregarious or clustered on rott ing wood or duff under conifers; late winter, spring, early summer, most abundant just after snowmelt; Pacific Northwest; common. EDIBILIT Y: tasteless and mostly water. COMMENTS: What has long been known as Sarcosoma mexicanum (Ellis and Holway) Paden and Tylutki (as Sarcosoma mexicana) is a complex of two species: a spring species from the Pacific Northwest and a fall species found in the west from Mexico north at least through California. DNA evidence indicates that the genus Sarcosoma is limited to Sarcosoma globosum (p. 249) and that the two western species cluster with Urnula. Urnula padeniana is the Pacific Northwest snow bank species and Urnula mexicana (Ellis and Holway) M. Carbone, Agnello, A. D. Parker, and P. Alvarado is the

more widespread, later-fruiting species originally named Bulgaria mexicana Ellis and Holway 1897 (Carbone, Agnello, and Parker 2013). Pseudosarcosoma latahense (Paden and Tylutki) M. Carbone, Agnello, and P. Alvarado (synonyms Sarcosoma latahense Paden and Tylutki and Plectania latahensis [Paden and Tylutki] M. Carbone), an uncommon Pacific Northwest species known as the black rubber cup, is similar, much less common, smaller (typically 2–4 cm across), and purple-brown before turning black. Its spores measure 24–38 × 9–12 μm and contain a few small oil drops when young but lack oil droplets at maturity, and the tomentum hyphae are usually light in color and smooth except for occasional dark granules (Paden and Tylutki 1969). DNA evidence places Pseudosarcosoma latahense in the Chorioactidaceae (with Neournula and Wolfina) and not in the Sarcosomataceae (Carbone, Agnello, and Alvarado 2013).

Pezizom ycetes

266

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Verpa bohemica var. bohemica (Krombholz) J. Schröter Ea r ly Mor el, W r ink led Thimble Ca p

fruitbody with a cap and stalk; cap up to 5 cm high and wide, broadly conic to bell-shaped or broadly oval, draping from its point of attachment at the top of the stalk, coarsely wrinkled by vertical folds that sometimes branch to form pits, pale yellowish brown to dark yellowish brown; flesh thin, fragile; stalk 5–15 cm high × 1–5 cm thick, nearly equal or tapered in either direction, typically roughened with fi ne granules, whitish to brownish yellow, hollow or stuffed with soft , cottony tissue. MICROSCOPIC FEATURES: spores 55–85 × (15–)17– 22 μm, elliptic-elongated, sometimes slightly curved, smooth, egutt ulate, hyaline; asci 250–350 × 18–25 μm, clavate, inamyloid, 2-spored; paraphyses fi liform, septate, with slightly clavate tips. OCCURRENCE: scattered or in groups on the ground under several types of hardwoods; spring, also winter in parts of California; northeastern North MACROSCOPIC FEATURES:

America west to the Pacific Northwest, south into California and the Rocky Mountains; locally common. EDIBILIT Y: edible for some but poisonous to many others, causing variable reactions, including severe gastrointestinal upset and temporary loss of coordination; never taste raw. COMMENTS: Ptychoverpa bohemica (Krombholz) Boudier is a synonym according to Species fungorum while MycoBank lists Ptychoverpa bohemica as the preferred name. The massive spore size and 2-spored asci are a most unusual combination. Verpa conica (p. 268) is similar, but has a nearly smooth to fi nely wrinkled cap and much smaller spores. Morchella punctipes (p. 189) and Morchella populiphila M. Kuo, M. C. Carter, and J. D. Moore are also similar but the lower edge of the cap is attached to the stalk with a deep sinus.

Verpa bohemica var. bohemica

V e r pa

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Verpa conica (O. F. Müller) Swartz

Smooth Thimble-Ca p

fruitbody with a cap and stalk; cap 1–3 cm wide, 1.5–3 cm high, thimble-shaped to bell-shaped, broadly rounded to flattened over the center, att ached at the top of the stalk, with skirt-like sides that sometimes flare in age, nearly smooth to fi nely wrinkled, flesh thin and very britt le; underside of cap whitish to tan and smooth; stalk 4–11.5 cm long, 5–15 mm thick, nearly equal or tapered toward the apex, hollow or stuffed with white cottony hyphae; surface whitish, smooth to slightly granular, sometimes ribbed. MICROSCOPIC FEATURES: spores 17–26(–34) × 11– 16(–19) μm, elliptic to oval, smooth, uniseriate, hyaline; asci 250–350 × 18–23 μm, cylindric-clavate, inamyloid, 8-spored; paraphyses fi liform, septate, branched toward the bases, with slightly clavate tips. OCCURRENCE: solitary, scattered, or in groups on the ground in hardwoods, mixed woods, and especially old apple orchards; spring, also winter

in parts of its western range; northeastern North America west to the Pacific Northwest, south along the Rocky Mountains and California; occasional to locally common. EDIBILIT Y: listed as edible in some guides, but more recent reports suggest otherwise, and therefore it is not recommended. COMMENTS: Th is is most likely a species complex with at least two species in North America and is in need of further study. Verpa bohemica (p. 267) is similar. It has a much larger fruitbody, its cap is conspicuously wrinkled by vertical folds and is attached only at the top of the stalk, and it has extremely large spores that measure 55–85 × 15– 22 μm. Morchella punctipes (p. 189) and Morchella populiphila M. Kuo, M. C. Carter, and J. D. Moore are also similar, but have conspicuously wrinkled and pitted cap surfaces, and the lower edge of the cap is attached to the stalk with a deep sinus.

MACROSCOPIC FEATURES:

Verpa conica

Pezizom ycetes

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Wolfina aurantiopsis (Ellis) Seaver fruitbody up to 7 cm wide, a stalkless shallow cup; inner surface pale yellow to ochraceous or somewhat orange, smooth; outer surface with projecting folds, minutely roughened, brownish to black; flesh tough, becoming corky on drying, whitish. MICROSCOPIC FEATURES: spores 27–33 × 16–18 μm, broadly elliptic, granular within, smooth, uniseriate or slightly overlapping, hyaline to slightly yellowish; asci up to 300 × 17–18 μm, cylindrical; paraphyses fi liform, slightly enlarged at the tips. OCCURRENCE: solitary or in small groups or clusters on decaying wood in mixed woods, especially with pine and hemlock, on woodchips used MACROSCOPIC FEATURES:

for landscaping, and also reported to grow on soil; summer, fall, and early winter in the southern parts of its range; widely distributed in eastern North America, also reported from Colorado; uncommon. COMMENTS: The epithet aurantiopsis means “appearing orange-colored.” Until its fi rm, corky texture is noted, this uncommon cup fungus is likely to be mistaken for a pale form of Galiella rufa (p. 142). Galiella rufa is similar but the inner surface of its fruitbody is pale orange to dull orange or reddish orange, the outer surface has a dense layer of matted wooly hairs, its flesh lacks the corky texture, and it has smaller spores.

Wolfina aurantiopsis

Wolfina

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Wynnea americana Thaxter

Moose A ntlers, R a bbit Ea rs

fruitbody 2.5–14 cm wide, 6–13 cm high, composed of several (up to 24) elongated half-cups resembling rabbit ears on a short stalk; cup inner surface smooth, pinkish orange to pinkish red or orange to brownish orange; outer surface covered with small rounded verrucae, sometimes wrinkled at maturity, blackish brown to reddish brown; flesh fi rm, somewhat tough, brown; stalk up to 2 cm long, tough, solid, dark brown on the outer surface, whitish within. MICROSCOPIC FEATURES: spores 32–40 × 15–16 μm, elliptic, extremities apiculate, striate with several alternately light and dark bands extending MACROSCOPIC FEATURES:

the length of the spores, multigutt ulate; asci 500– 540 × 18 μm, subcymbiform; paraphyses septate, with enlarged tips, pale brown. OCCURRENCE: solitary or scattered on soil under hardwoods, arising from a tough, underground, brown sclerotium; summer and fall; widely distributed in eastern North America; uncommon to locally common. COMMENTS: Wynnea sparassoides (p. 271) is similar but its fruitbody is composed of numerous apothecia united into a beige to yellow-brown, cauliflower- to brain-like mass on a long, unbranched stalk, and it lacks a sclerotium.

Wynnea americana

Pezizom ycetes

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Wynnea sparassoides Pfister

Sta lk ed Cau liflow er Fu ngus

fruitbody consisting of a head and a long stalk; head 6–8 cm high and wide, more or less rounded, composed of numerous apothecia united into a compound cauliflowerto brain-like mass, beige to yellow-brown; stalk 20–30 cm long × 2–3 cm thick, unbranched, often grooved, solid, cartilaginous, brown on the outer surface, beige to whitish within, deeply buried in the ground. MICROSCOPIC FEATURES: spores 32–36 × 12–15 μm, bilaterally symmetrical, elliptic to naviculate, with prominent longitudinal ridges and 3 oil drops, hyaMACROSCOPIC FEATURES:

line; asci 350–375 × 16–20 μm; paraphyses fi liform, septate, with enlarged tips. OCCURRENCE: solitary on soil under leaf litter; summer and fall; New England, also known from central Mexico; rare. EDIBILIT Y: unknown. COMMENTS: Wynnea americana (p. 270) is similar but its fruitbody is composed of several rabbit-ear-like cups with an orange inner surface, a short stalk, and a tough, underground brown sclerotium.

Wynnea sparassoides

Wynnea

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Wynnella silvicola (Beck) Nannfeldt MACROSCOPIC FEATURES: fruitbody 1–5 cm broad × 1–8 cm tall, rabbit-ear-shaped or, rarely, irregularly cup-shaped, with one enlarged lobe and a short stalk; margin inrolled at fi rst, expanding when old; inner surface smooth or, rarely, wavy-wrinkled when old, dark purplish red-brown to dark brown when fresh, blackish brown when dry; undersurface bald, red-brown or pale red-brown to white near base; stalk 0.2–1.0 cm long × 0.2–2 cm thick, equal or narrowing downward, shallowly fluted or ribbed, solid or with a central chamber, yellowbrown to pale yellow overall, or white at the base; odor of the fruitbody unique and strong when dried, but not detected when fresh. MICROSCOPIC FEATURES: spores 17–25 × 11.5–17 μm, broadly elliptic, smooth, with 1 large oil drop, hy-

aline; asci 200 × 15–19; paraphyses clavate, 5– 6 μm wide at apex, brown, contents fi nely granular. OCCURRENCE: solitary to gregarious in duff, rarely in burned areas, under conifers; summer and fall; North America, Europe, and Asia; uncommon. COMMENTS: Wynnea atrofusca R. Heim, Helvella silvicola (Beck ex Saccardo) Harmaja, and Otidea silvicola Beck are synonyms. As of early 2013, Index fungorum gives Wynnella silvicola (Beck) Nannfeldt as the preferred name, while MycoBank gives Helvella silvicola (Beck) Harmaja as the preferred name. Otidea smithii (p. 202) is very similar macroscopically, but has smaller, narrowly elliptic spores (10–12[–14] × 6–7 μm) and larger hooked or bent paraphyses that are sometimes ornamented with small, irregular protuberances.

Wynnella silvicola

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Chapter 5 Sordariomycetes

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he Sor da r iom ycetes is a cl ass of Ascomycota that as of this writing is known to comprise 15 orders, 64 families, many more than 1,000 genera, and far more than 10,000 species. The Sordariomycetes are informally referred to as “pyrenomycetes.” The word is derived from Greek meaning “stone fungi,” in reference either to the pebble-like appearance of the ascomata or to their often quite hard consistency. Sordariomycetes usually produce at least two kinds of spores in their life cycles: sexual and asexual. At least in temperate regions, asexual spores, known as conidia, begin to form in the spring and continue to shed throughout the growing season. The conidia germinate and the mycelium produced from them grows in or on soil, litter, or wood, or on fungi, insects, plants, and in some cases humans. When sexually compatible mycelia meet, their meeting initiates the sexual process, which leads ultimately to the formation of the pyrenomycete fruit body, or perithecium. In temperate regions, the formation of the perithecia occurs at the end of summer, and mature perithecia are found in late summer and autumn. The perithecia of most species are found on woody substrata as well as on herbaceous material, other fungi, insects, or even bones. Many pyrenomycetes appear to arise directly on soil, but often in those cases the true substratum is an insect that is buried in the soil. The vast majority of perithecia are far too small to catch the eye of even serious mushroom hunters, but a number of species are so spectacular in form or lifestyle that they have caught our eye and may fascinate you as well. Among the pyrenomycetes that are the most conspicuous because of the large size and/or bright coloration of their stromata are the Hypocreales, the Xylariales, and a few of the Sordariales.

In the past, most of the brightly colored pyrenomycetes have been classified in either the Hypo-

creales, which are usually plant- (but not grass-) or fungus-associated, or the Clavicipitales, which are usually thought of as insect or grass parasites but also include some mycoparasites. Prior to 1970, the Clavicipitales were recognized by some as a distinct order and by others as a family of the Hypocreales (Clavicipitaceae). In 1970 (Rogerson 1970), the orders were separated. Until recently (Rossman et al. 1999), each order included only a single family, the Hypocreaceae and the Clavicipitaceae, respectively. However, recent evidence from DNA sequences has supported the merger of these two orders into one, the Hypocreales. The distinguishing features of members of the Hypocreales order are summarized as follows. The perithecia are free, often in more or less conspicuous, compacted mycelium (subiculum), or are seated on or enclosed within a stroma. They are usually brightly colored shades of red and yellow, or sometimes brown or black. When they are red or red-orange, they sometimes become darker red in 3 percent KOH and yellow in lactic acid. These reactions are reversible. Perithecia are typically soft and easily crushed. There are no paraphyses among the asci or, if present, they are rudimentary. The asci are unitunicate and usually have an apical discharge mechanism. In the Clavicipitaceae, the asci have an apical ring that appears as a thick cap pierced by a pore. Ascospores are hyaline or lightly pigmented brownish, rarely green. They are variable in septation from aseptate to phragmosporous, although rarely aseptate. Conidia arise from phialides on free conidiophores, or on conidiophores arising from stromata or pycnidia. Today the order Hypocreales includes the families Hypocreaceae, Nectriaceae, Bionectriaceae, Niessliaceae, Clavicipitaceae, Cordycipitaceae, and Ophiocordycipitaceae. The latter three make up the old Clavicipitales order. Some members of the order Hypocreales are subjects of intense research because of their usefulness in biological control, medicine, and energy production. The Clavicipitaceae, Cordycipitaceae, and

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T

Hypocreales

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Ophiocordycipitaceae are distinguished from the other families in the Hypocreales order by having mostly multiseptate spores that are fi liform and asci with a distinct apical crown. Most Cordycipitaceae and Ophiocordycipitaceae parasitize insects, less frequently subterranean fungi (Elaphomyces; Th iers and Largent 1973), and are more or less conspicuous for the formation of brightly colored (often yellow or red) stromata that arise from the cadavers of their hosts (Mains 1934, 1939, 1940a, 1950a, 1950b, 1954b, 1955, 1957, 1958, 1959). An overview of these fungi is found in a publication by Sung et al. (2007). In North America, the greatest diversity of Cordycipitaceae and Ophiocordycipitaceae occurs in the Appalachians. Many of the Appalachian species are also found in disjunct populations in Asia, and it is anticipated that Asian species like Cordyceps pseudomilitaris Hywel-Jones and Sivichai may also be discovered as adjuncts in the Appalachians. A paper on Ophiocordyceps unilateralis by Evans, Elliot, and Hughes (2011) sheds some light on the variability that may be encountered in just this one species (actually a species complex) that grows on formicine ants. They note that it is stunning to begin to understand that “for most of the insectmicrobial interactions, the ecology and, more pragmatically, the pharmacological properties of the microbes involved remain unknown” (Evans, Elliot, and Hughes 2011, p. 598). In studying Ophiocordyceps variabilis on its carpenter ant hosts, Evans, Elliot, and Hughes (2011, p. 599) discovered a complex scenario, “one that will necessitate a re-evaluation of the diversity of species within Ophiocordyceps unilateralis s.l., as well as in the genus Ophiocordyceps and related genera of Hypocreales—and eventually, within the kingdom of fungi.” The biological scenario is fascinating. The fungal pathogen must att ach securely to the host and penetrate the exoskeleton while avoiding or suppressing host defenses. Then it must control the behavior of the host before killing it. Finally it must protect the cadaver from micro-

bial and scavenger att ack. “What is not generally realized—certainly not by drug companies—is that such coevolved fungi are pleomorphic, with well-defi ned parasitic, necrotrophic and saprotrophic phases: each one morphologically, genetically and physiologically distinct” (Evans, Elliot, and Hughes 2011, p. 599). The Chinese people have long recognized the medicinal potential of Ophiocordyceps sinensis (Berkeley) G. H. Sung, J. M. Sung, Hywel-Jones and Spatafora, and numerous publications in recent years rank the medicinal properties of Cordyceps militaris with that of Ophiocordyceps sinensis (Yu et al. 2004; Mao and Zhong 2006). Elaphocordyceps ophioglossoides has been used traditionally and has shown some interesting medicinal properties in animal testing. While we include numerous representatives of the Cordycipitaceae and the Ophiocordycipitaceae, we include just one species from the Clavicipitaceae, Claviceps purpurea. Members of the Clavicipitaceae are parasites of grasses, less frequently sedges. Most often, they form black, clublike stromata in place of the seeds of their hosts. Claviceps purpurea is notable for the formation of distinctive, dark-colored, horn-like sclerotia, known as ergots, which grow in the ovary of the flowers of their hosts. Ingestion of the sclerotia was often the lot of the poor in times of famine, when they could only afford to use rye flour. Unlike flour made from wheat, which is rarely infected by ergot, rye flour often was infected with ergot, and its ingestion resulted in an appearance of being possessed, hot burning sensations in limbs followed by a gangrenous condition, and often loss of limbs. In Medieval Europe, the disease became known as St. Anthony’s fi re (Hudler, Jensen-Tracy, and Banik 1998). Hudler, Jensen-Tracy, and Banik (1998) note that from 1560 to 1689 in southwest Germany and from 1650 to 1700 in Russia, years with cold, wet spring weather were correlated with a marked rise in trials for witchcraft the following fall and winter. The trial victims were typically peasants who lived

Overview

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The Hypocreaceae family is represented by 5 genera that are most easily distinguished from each other by DNA. We include 28 Hypocrea species, 20 Hypomyces species, 2 Protocrea species, 1 Podostroma species, and 1 Sporophagomyces species. Species of the genus Hypocrea grow on a variety of substrates, with stromata varying from spongy and loose to fi rm with a distinct rind. They have 2-celled spores that break apart at maturity into part-spores, causing the asci to appear to have 16 spores per ascus. Most Hypocrea species form light-colored but inconspicuous stromata on woody substrata. However, there are outstanding exceptions, such as Hypocrea sulphurea, which forms extensive bright yellow stromata that overgrow fruitbodies of the jelly fungus Exidia nucleata. Hypocrea avellanea transforms gills of Marasmius subnudus (Carey and Rogerson 1976). Hypocrea latizonata spreads over basidiomes of the

bird’s nest fungus Cyathus striatus, and club-like yellow-brown stromata of Hypocrea alutacea arise from wood (Chamberlain et al. 2004). Asexual stages of Hypocrea are known as Trichoderma. It is as the powdery, green-spored Trichoderma stage on logs or in Petri plates that these fungi are most commonly seen. A vast amount of published experimental work has been done on Trichoderma in areas as diverse as cellulases and biofuels production, opportunistic mycoses and allergic reactions, bioremediation, biological control of plant diseases, and “green mold” of commercial mushrooms. Many of the biological activities of the combined genus Hypocrea/Trichoderma are summarized in a two-volume work that was published in 1998 (Harman and Kubicek 1998; Kubicek, Harman, and Ondik 1998). Hypomyces stromata develop on fungi or on a thin subiculum, and have densely verrucose and somewhat fusoid 2-celled ascospores that do not break into part-spores at maturity. The spores are greater than 15 μm long, often with distinct apiculi. Hypomyces species are among the most conspicuous members of the Sordariales and are found on boletes, polypores, agarics, other ascomycetes, and soil (Rogerson and Simms 1971; Rogerson and Samuels 1985, 1989, 1993, 1994; Pennycook 2009). Hypomyces lactifluorum, the “lobster fungus,” completely parasitizes its large host (Lactarius or Russula species, often Russula brevipes), transforming it into a brilliant orange to reddish purple launching platform for its ascospores. Different species of Hypomyces parasitize other members of the Russulaceae family and, less frequently, members of the genus Amanita. Still others turn boletes and bolete relatives (such as Scleroderma) into slimy piles of goo, while yet another group of Hypomyces species transforms the undersurface of polypores and Stereum species. Species of Hypomyces that occur on boletes often have bright yellow (Sepedonium) conidial stages that are more common than the corresponding perithecial forms. Hypomyces cervi-

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in river valleys where rye was a staple food and the weather was the coolest and wettest, thus most favorable to the growth of Claviceps purpurea. From 1690 to 1692 it was unusually cool and wet in Salem, Massachusett s. The women who were placed on trial for witchcraft all reported symptoms consistent with ergotism, including pricking sensations like ants crawling on their skin, dementia, hallucinations, convulsive seizures—the same symptoms reported by the young adolescents who were supposedly “bewitched.” Recent research has shown that adolescents are more susceptible to the toxic alkaloids. Animals in the Massachusett s colony were also affected, and some died. Milk production by cows dropped dramatically. Some domestic animals lost limbs due to dry gangrene. The fact that animals as well as people were affected leads many scientists to suspect that the root cause of the problems in Salem was not simply hysteria, the common explanation for the Salem witch trials. Hypocreaceae

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nigenus (on Helvella) and Hypomyces ochraceus (on Lactarius and Russula) are usually seen in their asexual stage. The vast majority of species in the Hypocreales order have two spore forms, each with independent names. Despite the fact that a single species can have two (or even more) genus names, those names taken together include but one species. The teleomorph and anamorph stages may be separated from each other geographically or in time (e.g., asexual spores form during the summer, sexual spores appear only late in the year). Some species may have lost the ability to undergo sexual reproduction entirely, while others may have lost the ability to undergo asexual reproduction. For example, no anamorph has been discovered for Hypomyces lactifluorum. The teleomorph genus Cordyceps is genetically linked to at least five less well-known anamorph genera (Beauveria, Lecanicillium, Isaria, Microhilum, and Simplicillium). The different anamorph names refer to different specific morphologies of the asexual stage. When one reads Beauveria, the names Cordyceps or Isaria may, for example, be substituted from a nomenclatural point of view. Biologically, the anamorph names may not be completely interchangeable even though nomenclaturally they are. Members of the Niessliaceae family form tiny black perithecia, are inconspicuous and uncommon, and so are not included. Members of the Nectriaceae family typically form red, orange, or pale perithecia on woody plants. They are common in the Northern Hemisphere. The former genus, Nectria, has been broken up into smaller genera (Hirooka, Rossman, and Chaverri 2011; Hirooka et al. 2012). The current, revised “true” Nectria is mainly a tropical genus and has perithecia that are red. However, it also includes Nectria cinnabarina (Tode) Fries, the “coral spot” fungus that forms pinkish stromata with masses of conidia on many shrubs in North America and Europe. Nectria cinnabarina has 1-septate

spores in perithecia that form in dense clusters on Berberis or Ribes, or in conspicuous red perithecia on hardwood bark. Neonectria coccinea var. faginata (Persoon) Rossman and Samuels is the dominant species found on scale-infested beech trees in eastern North America. Species of the old genus Nectria that have brown, orange, yellow, or white (and in one case purple) perithecia and do not change color in 3 percent KOH or lactic acid are now classified in several genera of the Bionectriaceae. Bionectria species form orange perithecia on hardwood trees and are especially common in warmer parts of North America. Hydropisphaera is distinguished by a relatively wide ascomatal wall, a greater than 25 μm width, being composed of thin-walled cells that often collapse on drying to form cupulate perithecia. There are 23 recognized species and we include the type for the genus Hydropisphaera peziza (Tode: Fries) Dumont. The sporangium of the “dog vomit” myxomycete (Fuligo septica) turns purple when infected by Nectriopsis violacea. Nectriopsis species grow on myxomycetes and fungi, and have 2-celled spores that do not break into part-spores. Spores are more or less fusoid, less than 15 μm long, and never apiculate.

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Sordariales The Sordariales includes five families, two of which are represented here: Helminthosphaeriaceae, which contains 9 genera, and Lasiosphaeriaceae, which contains 40 genera. Helminthosphaeria clavarium, the one included species in the Helminthosphaeriaceae, parasitizes certain Clavaria species, slightly deforming the coral fungus and covering the surface with minute gray to black perithecia. The Lasiosphaeriaceae are represented by three species of Lasiosphaeria, a genus characterized by small (< 1 mm × 1 mm), hairy perithecia found on wood, with very long wormlike spores.

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The Xylariales order is composed of seven to nine families with anywhere from around 170 to more than 200 genera. A limited amount of work has been done with this class, and revisions are actively under way. Two families are included in our work: the Diatrypaceae and the Xylariaceae. The Diatrypaceae, once in the Diatrypales order (Kirk et al. 2008), are currently considered by MycoBank to be part of the Xylariales. Diatrypaceae species have perithecia with short necks that separately reach the surface of welldeveloped stroma composed mainly of fungal tissue. We include one representative from each of two genera, Diatrype (with eight spores per ascus) and Diatrypella (with more than eight spores per ascus) (Vasilyeva and Stephenson 2004). Unlike many Sordariomycetes, anamorph morphology for the Diatrypaceae is virtually useless at both the genus and the species level since the Diatrypaceae anamorphs look indistinguishable. Xylariaceae species are placed in up to 73 genera, ten of which produce fruitbodies likely to attract mushroom hunters and naturalists. We include Annulohypoxylon, Biscogniauxia, Daldinia, Entonaema, Hypoxylon, Kretzschmaria, Rosellinia, Thuemenella, and Xylaria. Biscogniauxia species have a broadly att ached applanate, eff used-pulvinate, or raised discoid stromata. They are uniquely two-layered with a dehiscent outer layer that often adheres to overlying bark and an inner layer that initially bears the anamorph or papyraceous remains of the anamorph. The mature surface is white, dark brown, or black with carbonaceous material immediately below it, with or without carbonaceous material surrounding the perithecia. The interior is essentially homogenous, not zoned, with ostioles at the same level as the stromatal surface or lower. They often serve more than one perithecium. The asci are

8-spored, typically with an amyloid apical ring. The spores are pale brown to blackish brown, and unicellular. Sometimes they have a straight to sigmoid germ slit that extends part way or along their entire length (Vasilyeva, Rogers, and Miller 2007). Daldinia species differ from Biscogniauxia species primarily in having just one stromatal layer, a zoned interior, and a constricted or a stalked attachment to the substrate (Ju, Rogers, and San Martín 1997; Ju et al. 1998; Rogers et al. 1999). Hypoxylon species are like Daldinia species except that they are not zoned internally. Kretzschmaria species also have just one stromatal layer, are usually stalked, or if sessile are constricted at the point of attachment, and form a crust (Rogers and Ju 1998). Rosellinia species are distinguished by having a subiculum and being composed of a single perithecium. Xylaria species are also single-layered and unzoned (Rogers 1983, 1984a, 1984b, 1986; Rogers and Callan 1986a, 1986b; San Martín, Lavin, and Rogers 2001). Entonaema species are fi lled with liquid when fresh and have orange granules surrounding the perithecia (Rogers 1981). Thuemenella species are distinctive for their fully sessile, broad attachment to the substrate, the lack of apical rings on the asci, and having cubical, pleated spores lacking a germ slit (Parker 1990). A synoptic key to 23 North American Xylaria species was produced by Callan and Rogers (1993). Ju and Rogers (1996) recognize two sections of the genus Hypoxylon: section Annulata (which we treat as Annulohypoxylon) and section Hypoxylon. Carbonaceous material surrounds the perithecia of Annulohypoxylon, and the ostiolar disks are annulate and always above the surrounding stromatal surface (Vasilyeva, Rogers, and Miller 2007). Species of the section Hypoxylon lack carbonaceous material surrounding the perithecia and the ostioles, which are rarely higher than the surrounding stromal surface and never have an annulate disk (Hsieh, Ju, and Rogers 2005).

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Xylariales

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Akanthomyces aculeatus Lebert fruitbody cream to yellowish, with variable 1–8 × 0.1– 0.5 mm branchlike structures arising from various parts of adult moths, often on the wings, tracing patterns of the moth’s imbedded wing veins; mycelium partially covering the insects, fi xing them to the substratum. MICROSCOPIC FEATURES: conidia 3– 6 × 2–3 μm, broadly elliptic or obovoid, often acute at the lower end, catenulate, smooth, hyaline; phialides 6–16 × 2.5–4 μm, smooth, densely compacted into a layer covering the synnemata, arising from lateral cells or terminating the hyphae, narrowed above to an MACROSCOPIC FEATURES:

acute apex terminated by short sterigmata up to 4 μm long. OCCURRENCE: covering dead adult moths that are often adhered by the fungus to a branch, leaf, fern frond, or other such substrate; summer; known from eastern and western North America; rare. COMMENTS: E. B. Mains (1950a) wrote that Akanthomyces aculeatus is highly variable in its development of the synnemata and in the shape and size of the conidia. It was once believed to be the anamorph of Cordyceps tuberculata (Lebert) Maire. Cordyceps tuberculata itself is a rather spectacular species in which several stromata arise from white

Akanthomyces aculeatus

A k a nthom yces

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mycelium on the host moth. The stromata are quite variable, measuring 2–11 mm × 1.5–2 mm, and are yellowish white or gray branches arising from a central core on a brown stalk. The perithecia, 400– 900 × 180–370 μm, are dark brown and partially embedded. The ascospores are fi liform and multiseptate, breaking into 2– 6 × 0.5–1 μm part-spores. The perithecia of Akanthomyces aculeatus are not embedded. Isaria sphingum Schweinitz, Hymenostilbe sphingum (Schweinitz) Petch, and Akanthomyces sphingum (Schweinitz) Petch are all synonyms of the anamorph of Cordyceps sphingum (Schweinitz) Berkeley and M. A. Curtis (not illustrated). They were also once thought to be the same as Akanthomyces aculeatus because of their marked similarities. The conidia of Akanthomyces are catenulate and the conidia of Hymenostilbe are solitary on the phialide. Other Akanthomyces species are found on Lepidoptera larvae and on spiders. Akanthomyces aranearum (Petch) Mains (not illustrated) grows on spiders and produces white, powdery, branch-like structures covered with chains of narrowly obclavate hyaline conidia mea-

suring 8–14 × 1.5–3 μm. The teleomorph is Cordyceps thaxteri Mains (not illustrated). It is characterized by small, villose, brownish, cylindric growths measuring 1.5–2.5 × 0.15– 0.2 mm and projecting from the spider. The fi liform ascospores of Cordyceps thaxteri measure 6–10 × 1–1.5 μm, are multiseptate, and tardily break into one-celled fragments. Isaria farinosa (Holmskjold) Fries is a striking fungus that forms beautiful coremia on a number of different insects. In culture, the spore-producing structures are highly variable, ranging from single detached phialides to penicillate heads with several stages of branching. The colors range from white to yellow to orange. The variability has resulted in authors assigning many different names to this species, including Coremium gracile Petch, Paecilomyces farinosus (Holmskjold) A. H. S. Brown and G. Smith, and Penicillium alboaurantium G. Smith. The coremia growing from the affected insect are typically about 1–2 mm high by 0.3–0.7 mm broad with irregularly ovoid heads. The conidia are hyaline, oval to subglobose, 1–2.5 × 1–1.5 μm. The teleomorph is unknown.

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Annulohypoxylon cohaerens (Persoon) Y. M. Ju, J. D. Rogers, and H. M. Hsieh fruitbody 1.5–8 mm wide, nearly globose to pulvinate, dry, hard, surface inconspicuously roughened, blackish; interior dark brown to blackish with embedded perithecia surrounding a solid center; ostioles papillate, lacking an ostiolar disc. MICROSCOPIC FEATURES: spores 9–12 × 4–5 μm, elliptic-inequilateral with narrowly rounded ends, smooth, brown to dark brown; asci 120–160 × 6– 7 μm. OCCURRENCE: scattered, in groups, or in clusters typically on decaying beech logs, stumps, and branches; summer and fall; widely distributed in eastern North America; fairly common. COMMENTS: Hypoxylon cohaerens (Persoon) Fries and Hypoxylon cohaerens var. microsporum J. D. Rogers and Candoussau are synonyms. Annulohypoxylon annulatum (Schweinitz) Y. M. Ju, J. D. MACROSCOPIC FEATURES:

Rogers, and H. M. Hsieh (not illustrated) is very similar but typically grows on oak, has coarsely conical-papillate ostioles that are encircled by convex discs, and has spores that measure 7.5–10.5(– 11) × 3.5–5(– 6) μm. Annulohypoxylon truncatum (Schweinitz) Y. M. Ju, J. D. Rogers, and H. M. Hsieh (not illustrated) is also similar but typically grows on oak. Its fruitbody is dark reddish brown, has papillate ostioles that are encircled by convex discs, and has spores that measure 8–10.5 × 4–5 μm. The birch hypoxylon, Annulohypoxylon multiforme var. multiforme (Fries) Y. M. Ju, J. D. Rogers, and H. M. Hsieh, forms hemispherical and irregularly shaped, roughened, spreading crusts over the bark of birch logs and branches. The fruit bodies are redbrown when young, become blackish brown to black in age, and produce spores that measure 8.5– 12 × 3.5–5 μm.

Annulohypoxylon cohaerens

A n n u loh y pox y lon

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Annulohypoxylon thouarsianum var. thouarsianum (Léveillé) Y. M. Ju, J. D. Rogers, and H. M. Hsieh MACROSCOPIC FEATURES:

fruitbody hemispheric to spherical, lacking a stalk, 0.8–5 cm across × 0.4– 3 cm thick; surface carbonous, blackish brown, smooth or roughened with projecting perithecia; perithecia peripheral, spherical to compressedspherical, with inconspicuous perithecial mounds; granules immediately beneath the surface, blackish with dull green KOH-extractable pigments; tissue below the perithecial layer massive, woody, dark brown; ostioles 0.2– 0.5 mm in diameter, papillate, encircled with a sunken but convex Hypoxylon truncatum–type disc that flakes off gradually from the ostiole outward. MICROSCOPIC FEATURES: spores 14–24 × 4– 6 μm, elliptic-inequilateral, shriveled, with narrowly rounded ends, unicellular, with a straight germ slit spore-length on the flattened side, overlapping uniseriate, light brown to brown; perispore indehiscent in 10 percent KOH, smooth; epispore smooth; asci 130–140 × 6–10 μm with a stalk 30–40 μm long, cylindric, very evanescent; perithecia 300–700 μm in diameter, 500–1,300 μm high, obovoid; paraphyses soon deliquescing. OCCURRENCE: on wood, usually limbs with bark, chiefly oak; fruits summer, woody and persistent; North America, South America, and Africa; common. COMMENTS: Synonyms include Daldinia malleola (Berkeley and Ravenel) Kauff man, Hypoxylon oc-

cidentale Ellis and Everhart, Hypoxylon thouarsianum (Léveillé) Lloyd var. gilletianum (Saccardo) J. H. Miller, and Hypoxylon amaniense Hennings. Annulohypoxylon thouarsianum var. macrosporum (F. San Martín, Y. M. Ju, and J. D. Rogers) Y. M. Ju, J. D. Rogers, and H.M Hsieh differs in having larger spores measuring 24–30 × 6–7 μm. Annulohypoxylon truncatum (Schweinitz) Y. M. Ju, J. D. Rogers, and H. M. Hsieh (not illustrated) has a smaller and much thinner fruitbody, 0.2–1 cm wide × 0.6–2 mm tall, with a dark reddish brown surface, much smaller spores (8–10.5 × 4–5 μm), and perispores that are dehiscent in 10 percent KOH. Annulohypoxylon annulatum (Schweinitz) Y. M Ju, J. D. Rogers, and H. M. Hsieh = Hypoxylon annulatum (Schweinitz) Montagne (not illustrated) measures 0.2–2 cm × 2–4.5 mm and is intermediate in size between A. truncatum and A. thouarsianum. It has a blackish surface with olivaceous tones, spores 7.5–11 × 3.5– 6 μm, and perispores that are dehiscent in KOH. All three species are found mainly on oaks, and all have dull green KOH-extractable pigments. Daldinia Cesati and De Notarius species may be outwardly similar but when cut in half reveal alternating light and dark bands, a feature not observed with Hypoxylon or Annulohypoxylon.

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Annulohypoxylon thouarsianum var. thouarsianum

A n n u loh y pox y lon

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Beauveria bassiana (Balsamo-Crivelli) Vuillemin fruitbody a white mycelium bearing masses of powdery white spores appearing on body parts of infected insects. MICROSCOPIC FEATURES: conidia 2–3 × 2 μm, produced sympodially in acropetal succession on the end of delicate conidiogenous cells. OCCURRENCE: appears as a powdery white growth on larvae and adult arthropods, present in soils; observed in all seasons when insect larvae or adults are found, but principally summer; worldwide; common. COMMENTS: Synonyms include Isaria shiotae Kuru and Tritarchium shiotae (Kuru) Langeron. Th is species is the anamorph of Cordyceps bassiana Z. Z. Li, C. R. Li, B. Huang, and M. Z. Fan, which to date is MACROSCOPIC FEATURES:

known only from eastern Asia. There are many distinct lineages that may someday be recognized as distinct phylogenetic species. It is widely used as a biological control agent to control termites, thrips, whitefl ies, aphids, and various beetles. When the spores contact the body of an insect host, they germinate, penetrate the cuticle, and grow inside the host. Th is leads to the death of the insect in a few days, after which the white powdery growth appears on the cadaver and produces new spores. The composite image shows a cicada cadaver on the left , uncommon synnemata on a hawk moth shown in the upper right, and a photomicrograph at lower right showing a hyphal strand, two conidiophores, and two clusters of conidia.

Beauveria bassiana

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Biscogniauxia atropunctata (Schweinitz: Fries) Pouzar fruitbody an applanate stroma 2–50+ cm long × 2–25+ cm broad × 0.4– 0.7 mm thick; outer dehiscing layer thin, white, becoming blackish; at maturity white punctuated by black ostioles, eventually almost entirely black; carbonaceous beneath the surface; perithecia obovoid, 0.2– 0.3 mm wide × 0.3– 0.5 mm high; ostioles slightly higher than the stromatal surface with papillate openings. MICROSCOPIC FEATURES: spores (20–)23–30 × 11.5– 14.5 μm, elliptical with rounded ends, smooth with a straight spore-length germ slit, brown to dark MACROSCOPIC FEATURES:

brown; asci 150–170 × 16–18 μm, spore-bearing part 140–155 μm with an amyloid apex. OCCURRENCE: primarily on oak, possibly other hardwoods; common in southeastern North America, found as far north as Indiana and Pennsylvania, Europe. COMMENTS: Hypoxylon atropunctatum (Schweinitz) Cooke, Diatrype atropunctata (Schweinitz: Fries) Berkeley, and Nummularia cinerea Rehm are some of the synonyms of this distinctive species.

Biscogniauxia atropunctata

Biscogni au x i a

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Biscogniauxia mediterranea (De Notaris) Kuntze fruitbody erumpent through bark or wood, irregularly rounded to crust-like; surfaces several centimeters square, dotted with ostioles of the perithecia; perithecia 4– 5 mm wide, black, evenly embedded in a single layer in the black, coal-like stroma, with a dehiscing dark brown outer layer. MICROSCOPIC FEATURES: spores 17–24 × 7–10 μm, narrowly elliptic, smooth, with 1 oil droplet and an indistinct germ slit, uniseriate, dark brown; asci 150–185 × 7–12.5 μm, 8-spored, spore-bearing upper part 120–150 μm, with an amyloid apical ring, discoid, 2–3 μm high × 4–5 μm wide; paraphyses cylindric, indistinctly visible, numerous. OCCURRENCE: on hardwood, chiefl y oak, usually following fi re or drought injury; year-round; worldwide distribution. MACROSCOPIC FEATURES:

Biscogniauxia species, which are only found on deciduous trees, differ from Hypoxylon and Annulohypoxylon in having an inner material composed of host and fungus tissue versus the black to red to dark brown purely fungal tissue of Hypoxylon and Annulohypoxylon. Biscogniauxia marginata (Fries) Pouzar has strongly cupulate stromata, is black at maturity, and is dotted with light-colored umbilicate ostioles; spores are globose to subglobose, 11–14 × 9–12 μm, with a spiral germ slit. Biscogniauxia repanda (Fries) Kuntze is a boreal species found mostly on Sorbus species and characterized by cupulate black stromata with a raised margin, papillate ostioles, and elliptic spores measuring 10–12 × 4.5–5 μm with a straight germ slit.

COMMENTS:

Biscogniauxia mediterranea

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Camarops petersii (Berkeley and Curtis) Nannfeldt Dog’s Nose Fu ngus, Split Sk in Ca r bon Cushion

fruitbody 2–7 cm wide, cushion- to top-shaped, oval or irregular, initially covered by a white to dull brownish felt-like veil; stalk absent or rudimentary; outer surface grayish brown to yellowish brown, splitt ing open to expose a shiny, black, tar-like, slightly roughened inner surface and leaving a sheath around the sides of the fruitbody; margin typically ragged; flesh reddish brown on the upper portion, blackish below, not concentrically zoned when cut vertically, fi rm, rubbery, containing numerous oval, seed-like perithecia arranged in 5– 6 layers, odor disagreeable, creosote-like. MICROSCOPIC FEATURES: spores 6–8.5 × 3–4.5 μm, broadly elliptic at one end, broadly fusiform with a minute apical germ pore at the other end, smooth, usually bigutt ulate, dark brown; asci 8-spored, 50– 55 × 5– 6 μm, clavate. MACROSCOPIC FEATURES:

solitary, in groups, or in clusters on decaying hardwoods, especially elm and oak; summer and fall; eastern and central North America; rare to uncommon. COMMENTS: Peridoxylon petersii (Berkeley and Curtis) Shear, Hypoxylon petersii Berkeley and Curtis, and Numariola petersii (Berkeley and Curtis) Shear are synonyms. Kathie Hodge nicknamed this species “dog’s nose fungus” since it was roughly the size, surface texture, and color of her dog’s nose. Daldinia concentrica (p. 293) is similar and has a reddish brown to black, cushion-shaped to nearly round or irregular fruitbody, but its outer surface does not split to expose a shiny black inner surface, it lacks a ragged margin, and it is concentrically zoned when cut vertically. OCCURRENCE:

Camarops petersii

Ca m a rops

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Claviceps purpurea Fries

Ergot

fruitbody consisting of two stages, a sclerotium infecting various grasses and a stroma that develops on damp fallen sclerotia that were formed during the previous year; sclerotium 1.5–3.5 cm long, more or less cylindric with tapered rounded ends, typically curved, with shallow longitudinal grooves, hard, purplish black to brownish black; interior hard, white; stroma divided into a head and stalk; head 1.5–3 mm in diameter, rounded to somewhat flattened, orangeyellow to ochre or pale purple, darkly punctate from the protruding ostioles of embedded perithecia; stalk up to 3 cm long, 1–1.5 mm thick, cylindrical, smooth, usually twisted, colored like the head, attached to the sclerotium by a dense, white mycelial mat. MICROSCOPIC FEATURES: spores 65–120 × 0.5–1 μm, fi liform, smooth, multiseptate when mature, hyaMACROSCOPIC FEATURES:

line; asci up to 160 × 4.5–5 μm, elongated-cylindric, thick-walled, 8-spored; paraphyses lacking. OCCURRENCE: solitary to clustered on the inflorescences of many species of grasses, especially rye, wheat, and barley; sclerotia occur from late summer into early spring, stromata occur from the early summer into fall; widespread; common. EDIBILIT Y: poisonous, causing ergotism or St. Anthony’s Fire, which can be fatal. COMMENTS: Th is fungus is believed by some authors to be the cause of the abnormal behavior exhibited by individuals accused of being witches in Salem, Massachusett s, during the 1600s. The extremely potent hallucinogen LSD as well as ergotamine and other alkaloids are derived from this fungus.

Claviceps purpurea

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Cordyceps hesleri Mains fruitbody a club-shaped clava up to 4.5 cm long growing from the head of a cicada nymph; fertile upper portion about 7 mm broad, black; stalk 2 cm long × 0.5 cm thick, gray tinged with olive, and yellowish at the base. MICROSCOPIC FEATURES: spores fi liform, nearly as long as the asci, and breaking into part-spores 3– 4 × 1.5–2 μm; asci 300–360 × 6–7 μm, cylindric; perithecia 770– 960 × 360–420, fusoid-elliptical, entirely embedded. OCCURRENCE: parasitic on cicada nymphs; fall; southeastern North America in the Great Smoky Mountains; not common. COMMENTS: Ophiocordyceps sobolifera (Hill ex Watson) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora (not illustrated), which also grows from the head of a cicada nymph, is distinctly different, with a commonly solitary pale ocher, clavate to cylindrical, 20–80 × 2– 6 mm stroma that is internally white, and part-spores hyaline and truncate, measuring 6–13 × 1–1.5 μm. Ophiocordyceps sobolifera is known from Cuba, Mexico, and Asia. MACROSCOPIC FEATURES:

Cordyceps hesleri

Cor dyceps

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Cordyceps militaris (Linnaeus) Link fruitbody consisting of an orange-buff to orange stroma giving rise to a head and stalk, 6–10 mm wide, up to 7.5 mm tall, cylindric to clavate or fusiform, fi nely roughened on the head from the apices of the partially embedded perithecia, smooth and tapered downward on the lower stalk. MICROSCOPIC FEATURES: spores 250–350 × 1– 1.5 μm, fi liform, multiseptate, breaking into smaller barrel-shaped spores that measure 2–5(7) × 1– 1.5 μm, smooth, multiseriate, hyaline; asci 300– 510 × 3– 6 μm, 8-spored, narrowly cylindrical with a thick apical cap; paraphyses lacking. OCCURRENCE: solitary or in groups on the pupae, or less commonly the larvae, of partially buried Lepidoptera; summer and fall; widely distributed in North America; fairly common. COMMENTS: Cordyceps militaris is often collected without its host, which is shallowly buried in soil or under mosses. Cordyceps cardinalis G. H. Sung and Spatafora (not illustrated), reported from the southern Appalachian Mountains of the eastern United States, is very similar but has a small reddish stroma that emerges from the larval stage rather than the pupal stage of Lepidoptera. Its stalk is ochraceous orange to red and its head is reddish orange to reddish. Microscopically it has asci that measure 175–330 × 3–5 μm with a thick apical cap, and fi liform, irregularly multiseptate, nonfragmenting spores that measure 160–320 × 1 μm. Cordyceps olivascens Mains is distinguished by a very light green to olive-buff stroma that is minutely bumpy due to slightly projecting ostioles of the perithecia. Its flesh is thin, whitish, and fi rm, and the stalk is colored like the head or slightly darker olive, usually with a whitish base. Cordyceps olivascens, a rare species known from Alabama and Mississippi, produces stromata on the remains of buried or partially buried unidentified insects

Trooping Cor dyceps

MACROSCOPIC FEATURES:

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Cordyceps militaris

in rotten wood or humus and has smooth, hyaline, multispetate, fi liform spores that soon break into oblong fragments 3– 6 × 1–1.5 μm. Cordyceps washingtonensis Mains, known from Washington State, is closely related to Cordyceps militaris, grows on pupae and larvae of Lepidoptera, and differs in being creamy white to yellow. The perithecia are embedded in the early developmental stages, and once fully mature they can protrude as much as half of their length out of the stroma and are bright yellow; stalk whitish. Cordyceps washingtonensis spores are 80–110 × 1–1.5 μm, multiseptate, not breaking into part-spores (unlike Cordyceps militaris), narrowly cylindric-clavate, and narrowed in the lower part.

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Daldinia childiae J. D. Rogers & Y. M. Ju Cr a mp Ba lls, K ing A lfr ed’s Ca k es

fruitbody spherical to turbinate, sessile or with a short, stout stalk; 0.5– 5 cm × 0.8–4 cm; outer surface smooth or with inconspicuous to conspicuous perithecial mounds, chestnut to grayish sepia, darkened and dull when old; dull orange-brown or dull reddish brown granules immediately below the surface, with hazel to cinnamon KOH-extractable pigments; flesh pithy to woody with alternating zones, the darker zones 0.02– 0.06 cm thick and dark brown, the lighter zones 0.06– 0.1 cm thick and brown; perithecia 0.3– 0.5 mm across and 0.7–1.5 mm high.

MACROSCOPIC FEATURES:

MICROSCOPIC FEATURES: spores 12–16 × 5.5– 7.5 μm, elliptic-inequilateral, with narrowly rounded ends, unicellular, with straight germ slit spore-length on convex side, perispore dehiscent in 10 percent KOH, epispore smooth, brown to dark brown; asci at least 180 μm long × 8–12 μm wide, the sporebearing part about 90 μm long, the stalk at least 90 μm long, with amyloid apical ring; conidia 7– 9(–10.5) × 4.5–5.5 μm if present. OCCURRENCE: on hardwood, solitary to infrequently aggregated; year-round; widespread in North America and Europe; common.

Daldinia childiae

Da l di n i a

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COMMENTS:

The common name “cramp balls” is a reference to the folk belief that this fungus can prevent or cure menstrual cramps. Daldinia childiae has been known as three different forms or varieties of Daldinia concentrica f. confluens C. G. Lloyd, D. f. intermedia C. G. Lloyd, and D. var. minuta Waraitch. Daldinia concentrica (p. 293) differs from D. childiae in that it has stromatal surface scales and purple KOH extracts of the stromatal tissue. Daldinia lloydii Y. M. Ju, J. D. Rogers, and F. San Martín (not illustrated) has surface scales like D. concentrica, but lacks the KOH-extractable pigments. Daldinia clavata Hennings (not illustrated), Daldinia grandis (p. 294), Daldinia loculata (Léveillé) Saccardo (not illustrated), and Daldinia

gelatinosa Y. M. Ju, J. D. Rogers, and F. San Martín (not illustrated) all lack surface scales on the fruitbody. Daldinia gelatinosa is distinguished by a turbinate fruitbody shape, unique gelatinous disintegrating flesh, and whitish to grayish zone colors. Daldinia clavata also has whitish to grayish zone colors and has a distinctive clavate to cylindrical shape plus spores that are both shorter and narrower, 8–11.5 × (3.5–)4–5.5 μm. Daldinia grandis has longer and wider spores than all of the other species, 14–22(–22.5) 7–11 μm. With the exception of Daldinia clavata and D. grandis, the species in this group are microscopically highly similar and differ primarily in the features explicitly mentioned above.

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Daldinia concentrica (Bolton: Fries) Cesati and de Notaris Ca r bon Ba lls, Coa l Fu ngus, Cr a mp Ba lls, K ing A lfr ed’s Ca k es

fruitbody 2–5 cm wide, cushion-shaped to nearly round or irregularly shaped; surface uneven and furrowed, reddish brown at fi rst, becoming somewhat shiny black, fi nely roughened, and often with minute pores; flesh concentrically zoned when cut vertically, fibrous to powdery and carbon-like, dark purplish brown alternating with darker or sometimes whitish zones; perithecia embedded in a single layer near the surface. MICROSCOPIC FEATURES: spores 12–17 × 6– 7.5 μm, irregularly elliptic with one side flattened, smooth, unigutt ulate, with a barely visible germination cleft , uniseriate, dark brown; asci 210–250 × 8–11 μm, amyloid tips, 8-spored; paraphyses fi liform, septate. OCCURRENCE: solitary or in clusters on beech (Fagus) in beech-birch-maple forests; year-round; northern North America; common. MACROSCOPIC FEATURES:

According to legend, during wartime King Alfred was hiding in a country home. Unaware of his identity, the mistress of the house put him in charge of watching the baking and taking the cakes out of the oven when they were done. King Alfred fell asleep and the cakes burned. Daldinia concentrica supposedly resembles those burnt cakes. Daldinia tuberosa (Scopoli) J. Schröter, Hemisphaeria concentrica (Bolton) Klotzsch, and Hemisphaeria tuberosa (Scopoli) Kuntze are synonyms. Daldinia grandis (p. 294) is very similar and grows on decaying hardwood in western North America, but its fruitbody is dark brown when young, black when mature, and it has larger spores that measure 14–22(–25.5) × 7–11 μm. COMMENTS:

Daldinia concentrica

Da l di n i a

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Daldinia grandis Child fruitbody depressed spherical to hemispheric, sessile or nearly so, 2.5– 8 cm wide × 1.5–5.5 cm tall; surface smooth with inconspicuous to conspicuous perithecial mounds, brown-vinaceous, varnished black in age; dull reddish brown granules immediately beneath the surface, with or without livid purple to dark livid KOH-extractable pigments; interior tissue pithy to woody, with 0.3– 0.8 mm thick dark brown zones alternating with 0.8–2 mm thick gray to grayish brown zones that are gelatinous when fresh but very fi rm and hard when dry. MICROSCOPIC FEATURES: spores 14–22(–25.5) × 7– 11 μm, with straight germ slit spore-length on the more convex side, perispore indehiscent in 10 percent KOH, epispore smooth, dark brown; asci 220–250 × 10–13 μm, spore-bearing part 90–120 μm long, the stalk 100–150 μm, apical ring bluing in Melzer’s. MACROSCOPIC FEATURES:

solitary or sometimes aggregated on hardwood; year-round; known from western North America, and South America; regionally common. COMMENTS: Daldinia concentrica (Bolton: Fries) Cesati and De Notaris f. californica C. G. Lloyd is a synonym. The species is readily distinguished by larger spores than the other Daldinia species. Daldinia gelatinosa Y. M. Ju, J. D. Rogers, and F. San Martín (not illustrated) has white gelatinous zones alternating with darker zones and disintegrates rather than becoming hard on drying. Daldinia gelatinosa spores are 12.5–15.5 × 6–7.5 μm (similar to D. childiae). See Daldinia childiae (p. 291) for additional comments. An outstanding website, Mycology (mycology.sinica.edu.tw), contains extensive information on all members of the Xylariaceae if more information is desired. OCCURRENCE:

Daldinia grandis

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Diatrype stigma (Hoff mann) Fries fruitbody forms a widespread crust, highly variable in shape and size, up to 23 cm or more long × 1–1.5 mm thick; surface shiny, fi nely roughened to nearly smooth, sometimes minutely cracked, purplish to reddish brown when young, soon blackish brown to black; flesh whitish, fibrous-tough; perithecia embedded just beneath the surface in a single layer. MICROSCOPIC FEATURES: spores 6–12 × 1.5–2 μm, allantoid, smooth, usually with 1–2 oil droplets at each end, irregularly biseriate, hyaline; asci 35– 40 × 5– 6 μm, clavate, 8-spored; paraphyses lacking. OCCURRENCE: on decaying hardwood branches, especially beech, often surrounding them; yearround; eastern North America; common. MACROSCOPIC FEATURES:

The nearly smooth, shiny surface and hyaline spores are characteristic of this species. Other similar black, spreading crusts have more coarsely roughened surfaces or dark brown spores. Diatrype undulata (Persoon) Fries and Sphaeria stigma Hoff mann are synonyms. Diatrypella favacea (Fries) Cesati and De Notaris = Diatrypella verruciformis (Ehrhart) Nitschke is a widespread parasite that erupts through the bark of hardwoods. It is brown-black and whitish within, and has allantoid spores that measure 6–8 × 1.5 μm, with many spores per ascus.

COMMENTS:

Diatrype stigma

Di atry pe

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Elaphocordyceps capitata (Holmskjold) G. H. Sung, J. M. Sung, and Spatafora Hea d-Lik e El a phocor dyceps, Rou nd-Hea ded El a phocor dyceps

MACROSCOPIC FEATURES:

fruitbody consisting of a head and stalk attached to its host; head 6–20 mm wide and high, irregularly rounded, fi nely bumpy, dark reddish brown to olive-black; flesh white, thick, fi rm, odor and taste not distinctive; stalk 2–8 cm long, 5–16 mm thick, nearly equal overall, smooth to slightly ridged, fibrous, yellow to dull yellow, becoming olive-brown in age. MICROSCOPIC FEATURES: spores cylindrical, multiseptate, hyaline, disarticulating into part-spores 8–25(–32) × 1.5–3 μm; asci 350–540 × 10–12 μm, cylindrical with a thickened apex, inamyloid; paraphyses lacking; perithecia ovoid, 650– 950 × 250– 420 μm, mostly immersed in stromata. OCCURRENCE: solitary or in groups in woods, attached to buried, walnut-shaped, reddish brown fruitbodies of Elaphomyces species; late summer, fall, and early winter; widely distributed in North America from eastern Canada to Florida, Europe and Asia; uncommon to locally common. COMMENTS: Cordyceps capitata (Holmskjold: Fries) Link, Cordyceps canadensis Ellis and Everhart, and Torrubia capitata (Holmskjold: Fries) Tulasne and C. Tulasne are synonyms. Elaphocordyceps longisegmentis (Ginns) G. H. Sung, J. M. Sung, and Spatafora, known from North America and Japan, is macroscopically identical but is easily identified by microscopic examination (Ginns 1988). Its smooth, nonamyloid, multiseriate, hyaline spores when mature are segmented into long truncated part-spores that measure 40– 65 × 4– 5 μm (although terminal segments can be as small as 12 × 3 μm). Asci are up to 440 × 10–15 μm, cylindrical to narrowly elliptic, and nonamyloid, with walls fragile at spore maturity and an apex with a thick apical cap with a narrow pore. Perithecia are

about 500 × 300 μm, imbedded, elliptic, and hyaline. Elaphocordyceps fracta Mains (not illustrated) has a purplish black head measuring 0.25 cm diameter that is distinctly separated from the slender, yellow-green to olivaceous stalk, has part-spores measuring 2–5 × 1.5–2 μm, and was fi rst collected on Elaphomyces appalachiensis Linder. Elaphocordyceps japonica (Lloyd) G. H. Sung, J. M. Sung, and Spatafora (not illustrated), known from Australia, Austria, Japan, and North America, has a deep olive-gray head not differentiated from the stalk, and fi liform spores that disarticulate into partspores and measure 10–18 × 2.5–4 μm. Th is overlaps in size with Elaphomyces capitata. Both species grow on members of the Elaphomyces granulatus group and some other Elaphomyces species. Elaphocordyceps valliformis (Mains) G. H. Sung, J. M. Sung, and Spatafora (not illustrated) has a distinctly separated dark brown head measuring 0.3– 1.5 × 0.3–1.2 cm, on a smooth or scurfy dark brown stalk, and part-spores 3–8 × 2 μm. Elaphocordyceps tenuispora (Mains) G. H. Sung, J. M. Sung, and Spatafora (not illustrated) has a dark brown obovoid fertile area measuring 1–1.5 × 0.8–1 cm that is not distinctly separated from the yellowish brown stalk and has part-spore segments of 6–8 × 1– 1.5 μm. There are also several other Elaphocordyceps species that have a clavate rather than a capitate head. See Elaphocordyceps ophioglossoides (p. 298). Note: All Elaphocordyceps species grow on species of the truffle Elaphomyces. Several Ophiocordyceps species—see O. dipterigena (p. 337) and O. myrmecophila (p. 340)—also have capitate heads, but if you dig into the soil, you will fi nd that they are attached to a mummified insect larva or pupa and not to a truffle.

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Elaphocordyceps capitata

El a phocor dyceps

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Elaphocordyceps ophioglossoides (Ehrhart) G. H. Sung, J. M. Sung, and Spatafora Anamorph: Verticillium-like A dder’s Tongue, Golden-Thr ea d El a phocor dyceps

stromata consisting of a head and stalk attached to its host; head 1–1.6 cm wide, 2–2.5 cm high, fusiform to oval or clavate, fi nely bumpy, yellowish brown to dark reddish brown, becoming olive-black in age; flesh whitish, thick, fi rm; stalk 2.5–14 cm long × 1.5–10 mm thick, nearly equal overall, smooth, yellow, with conspicuous golden yellow basal rhizomorphs. MICROSCOPIC FEATURES: spores cylindric, multiseptate, disarticulating into truncated part-spores 2–5 × 1.5–2 μm, smooth, hyaline; asci 280–450 × 5–7 μm, cylindrical, 8-spored; paraphyses lacking; perithecia 600–700 × 260–370 μm, ovoid, immersed. MACROSCOPIC FEATURES:

solitary or in groups in woods, on buried walnut-shaped reddish brown fruitbodies of Elaphomyces species, known to grow on Elaphomyces granulatus and Elaphomyces variegatus; summer and fall; widely distributed in North America; uncommon to locally common. COMMENTS: Cordyceps ophioglossoides (Ehrhart: Fries) Link and Torrubia ophioglossoides (Ehrhart: Fries) Tulasne are synonyms. Elaphocordyceps paradoxa Kobayasi, reported from Japan, is very similar in its macroscopic and microscopic features but is a pathogen of cicadae. For species with a capitate stromata and lacking golden yellow rhizomorphs, see Elaphocordyceps capitata (p. 296). OCCURRENCE:

Elaphocordyceps ophioglossoides

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Elaphocordyceps subsessilis (Petch) G. H. Sung, J. M. Sung, and Spatafora Anamorph: Tolypocladium inflatum W. Gams solitary stromata arising from Coleopteran larvae buried in wood; stromata 4–15 cm tall × 1–5 mm thick, irregular shape, whitish; stalk present or absent, often bifurcating if present and composed of loosely woven hyaline hyphae with pads of perithecia that appear at the surface of rotten wood. MICROSCOPIC FEATURES: spores fi liform, hyaline, disarticulating into 3– 6 × 0.5–1 μm truncate partspores; asci cylindrical, 430– 600 × 3–4 μm, apical cap 2 μm thick; perithecia 800–1100 × 320–450 μm, narrowly ovoid to conoid, pseudoimmersed, produced on a lateral pad, yellow to brown. MACROSCOPIC FEATURES:

on larvae of Scarabaeidae (Coleoptera) buried in wood and possibly also on larvae in soil and dung; presumably fall; known from Tennessee, North Carolina, Michigan, New York, and Washington as well as Japan; common. COMMENTS: The teleomorph-anamorph connection was established by Hodge, Krasnoff, and Humber (1996). Cordyceps facis Kobayasi and Shimizu from Japan is treated as a synonym but more research is needed to clarify host affi liation among numerous families of Coleoptera and habitat requirements (e.g., rotten wood versus soil or dung). OCCURRENCE:

Elaphocordyceps subsessilis

El a phocor dyceps

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Entonaema liquescens Möller Anamorph: Nodulisporium-like stromata 1– 6 cm high, 1–13 cm wide, sessile, somewhat globose or pulvinate to convoluted and cerebriform, hollow; surface bright sulphur-yellow to dull yellow or oliveyellow, occasionally with orange tones, sometimes staining greenish when handled, easily rubbed off and exposing a darker stroma below, conspicuously or inconspicuously dotted with black, punctate to barely papillate ostioles; flesh surrounding the perithecia and bordering cavities 2–5 mm thick and gelatinous when wet, but when dry less than 1 mm thick, hard, reddish to blackish; stromal cavity fi lled with watery liquid that escapes and causes the stroma to collapse when punctured. MICROSCOPIC FEATURES: spores (8–)9.5–11.7(–13) × (4–)5– 6(–7.5) μm, elliptic-inequilateral to rectangular with blunt ends, with or without a straight germ slit that runs less than the full length of the spore, usually conspicuously bigutt ulate, pale to dark brown; asci 120–160 × 6–10 μm, 8-spored. OCCURRENCE: scattered or in clusters on decaying wood; summer, fall, and early winter; most commonly collected in the southeastern United States, but also reported from Illinois and Kansas; occasional to locally fairly common. COMMENTS: Although several different species of Entonaema have been reported from many other countries, Entonaema liquescens is the only Entonaema described from North America, to date. The epithet liquescens refers to the watery liquid contained in the stroma of fresh specimens. MACROSCOPIC FEATURES:

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Helminthosphaeria clavariarum (Desmazières) Fuckel Anamorph: Spadicoides clavariarum (Desmazières) S. Hughes fruitbody black to gray-black, consisting of minute perithecia on the branches of Clavaria coralloides (Linnaeus) J. Schroeter (synonym Clavulina cristata [Fries] J. Schröter) and Clavulina cinerea (Fries) J. Schröter; perithecia 0.3 mm across, lacking a distinct papilla, scattered over a web of dark brown hyphae, superficial with conspicuous black setae on the perithecia. MICROSCOPIC FEATURES: spores 10–14 × 6– 7 μm, broadly fusiform or one-sided, nonseptate, dark brown; asci up to 90 × 8 μm, cylindric, stalked, MACROSCOPIC FEATURES:

8-spored; ascospores uniseriate, paraphyses numerous; some hairs tapering to a point, others cylindric and bearing elliptic, 1-septate, brown conidia. OCCURRENCE: known only from fruitbodies of Clavulina coralloides and C. cinerea; fall; in temperate regions worldwide; common. COMMENTS: The presence of Helminthosphaeria clavariarum seems to reduce the division of the branch tips of its host fungus. The conidial state darkens the base of the host Clavulina.

Helminthosphaeria clavariarum

H elminthosph a er i a

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Hypocrea alutacea (Persoon) Cesati and De Notaris Anamorph: Trichoderma alutaceum Jaklitsch fruitbody up to 5 cm high, consisting of a head and stalk; head 5–10 mm wide × 0.5–2 cm high, cylindric to clavate; surface fi nely pimpled due to the partially embedded perithecia, whitish to pale yellow at fi rst, becoming pale yellow-orange and darkening to orangebrown or brown at maturity; ostioles appear as tiny brownish dots; stalk 1.5–3 cm high, smooth, whitish to pale yellow, darkening to brownish in age; flesh white, fibrous-tough, odorless. MICROSCOPIC FEATURES: Part-spores hyaline, fi nely verrucose, uniseriate, dimorphic; upper part globose to subglobose, 2.7–3.7 × 2.5–3.5 μm; lower part subglobose to conical, 3–4 μm; asci cylindriMACROSCOPIC FEATURES:

cal, 65– 90 × 2.5–4.5 μm, 8-spored, apex slightly thickened. OCCURRENCE: solitary or in small groups on decaying wood; summer and fall; widely distributed in North America and Europe; rare. COMMENTS: Cordyceps alutacea (Persoon) Quélet and Podostroma alutaceum (Persoon) Atkinson are synonyms. Hypocrea alutacea is similar to Hypocrea leucopus (P. Karsten) H. L. Chamberlain, which grows on the ground among leaves and needle litter. The fertile head of Hypocrea leucopus is sharply delimited from the stalk while the fertile head of Hypocrea alutacea is not sharply delimited from the stalk.

Hypocrea alutacea

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Hypocrea latizonata Peck fruitbody a dirty white subiculum forming a 0.5– 0.8 cm wide band around the outside of cups of Cyathus striatus Hoff man, punctuated by the orange to dark brown, slightly prominent ostioles. MICROSCOPIC FEATURES: spores 2-celled, dividing into part-spores at maturity such that the asci apMACROSCOPIC FEATURES:

pear to contain 16 spores, hyaline; asci cylindrical, 75–80 × 3.5–4 μm. OCCURRENCE: on Cyathus striatus year-round; widespread in all temperate regions; common. COMMENTS: Because this grows on the birds’s nest fungus, Cyathus striatus, it is unique and distinctive.

Hypocrea latizonata

Hypocr ea

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Hypocrea pachybasioides Yoshimura Doi Anamorph: Trichoderma polysporum (link) Rifai stromata solitary to gregarious, occasionally cespitose, 0.3– 6+ cm broad × 0.2–4.5 mm thick, variable in outline, typically brown to reddish brown, sometimes yellow to light brown when young; margins often white, usually fully attached to the substratum; surface variously wrinkled or creased; ostiolar openings mostly visible as purple-brown to blackish brown slightly raised dots. MICROSCOPIC FEATURES: ascospores dimorphic part-spores; distal part-spores globose to conical, 2.2– 6 × 2–5.7 μm, proximal part-spores mostly subMACROSCOPIC FEATURES:

globose to oblong or wedge-shaped, 2.5–7.2 × 1.7– 4.5 μm, fi nely spinulose, uniseriate, hyaline; asci 50–130 × 3.5–8.5 μm, tip thickened with a pore; perithecia immersed in the stroma, globose to subglobose, 75–350 μm tall × 55–230 μm wide. OCCURRENCE: on wood; found in much of North America, Europe, Australia, and Asia; not frequently noticed. COMMENTS: Hypocrea pachybasioides is quite distinctive. If the stromata is blackish, see Hypocrea schweinitzii (p. 308).

Hypocrea pachybasioides

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Hypocrea peltata (Junghuhn) Berkeley fruitbody a cerebriform to shield-shaped stroma attached at a central point; 0.7–8 cm × 3 cm; fleshy or leathery, often convoluted, light tan; underside with somewhat conspicuous ridges radiating from center to margin as if it was a parasitized gilled mushroom, smooth, lighter in color than the upper surface, often with cottony white hyphae at the surface; perithecia numerous, crowded, immersed, and inconspicuous. MICROSCOPIC FEATURES: spores four large 2-celled spores 2.5–6 × 2–5 in distal part and 2–6.5 × 2–4.5 in proximal part, four small ascospores 2–4.5 × 2– 4 μm in distal part and 2–4.5 × 2–5 μm in proximal part, disarticulating at the septum, subglobose, fi nely spinulose, hyaline; asci 60–130 × 4–8 μm, MACROSCOPIC FEATURES:

narrowly clavate, sessile, apex slightly thickened with an obscure pore, 8-spored; perithecia 310– 385 × 170–240 μm, elliptic, immersed in a single row below the stroma surface; ostiolar canal 60– 120 μm long, wall 30 μm wide. OCCURRENCE: usually on hardwood, rarely on pines; presumably year-round; subtropical areas globally, extending as far north as North Carolina; common. COMMENTS: The large fruitbodies (stromata) with ridges on the underside and a central point of attachment give the appearance of a Hypomyces or a Hypocrea parasitizing an agaric, but there is no evidence of a parasitized fungus being present (Samuels and Ismaiel 2011). Th is is a highly un-

Hypocrea peltata

Hypocr ea

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usual Hypocrea species in that it does not produce a Trichoderma in culture. Trichoderma characteristically produces masses of green conidia. Hypocrea peltata causes a great deal of damage to shiitake production in Japan, and the destruction has been traced to the production of hypecelin A and B, which have been shown to have high antibiotic/ antifungal activity against a range of human pathogens (Samuels and Ismaiel 2011). Hypocrea peltata was once isolated from a human lung in Washington State. Th is unusual fi nding is att ributed to the fact that this species grows well at human body

temperature while almost all other Hypocrea species cease growth by 35 ºC. Gary Samuels (personal communication) cautions that “given its ability to infect a human, one should be a bit careful in ‘playing’ with this species (i.e., classroom demo and so on).” A number of Hypocrea species in the Longibrachiatum clade are known to cause invasive mycoses in humans. However, in the Washington State example, there was no evidence that the presence of this species in the lungs was the cause of the observed fibrosis.

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Hypocrea pulvinata Fuckel fruitbody rounded to irregularly oval, cushion-like, solitary or coalescing to form a crust; 3– 60 × 2–20 mm; surface smooth at fi rst, eventually roughened and dotted by tiny brown ostioles of the embedded perithecia, color yellow to grayish orange, sometimes brownish yellow to golden yellow at maturity, stromata generally turning orange or red with KOH but reaction sometimes weak. MICROSCOPIC FEATURES: part-spores slightly dimorphic, distal part 3.7–4.7 × 3.0–3.7 μm, proximal part 3.7–5 × 2.7–3.7 μm, irregularly globose to angular-oval, smooth to minutely spinulose, thinwalled, uniseriate, hyaline; asci 60–70 × 3.5–4 μm, cylindrical, 8-spored, forming 16 part-spores beMACROSCOPIC FEATURES:

fore maturity; lacking paraphyses; perithecia 190– 275 μm long, elliptic; ostioles 40–80 × 30–70 μm. OCCURRENCE: solitary, in groups, or in clusters, on a variety of polypores, including Laetiporus sulphureus, Fomitopsis pinicola, Piptoporus betulinus, and Ganoderma species, often coalescing and spreading on decaying stumps, branches, leaves, and the surrounding soil; late spring, summer, and fall; known from northeastern and northwestern North America; uncommon. COMMENTS: Hypocrea fungicola (P. Karsten) Saccardo and Hypocrea citrina f. fungicola P. Karsten are synonyms. A number of different Hypocrea species with eff used stromata are found in North America; see Hypomyces aurantius (p. 315).

Hypocrea pulvinata

Hypocr ea

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Hypocrea schweinitzii (Fries) Saccardo Anamorph: Trichoderma citrinoviride Bissett MACROSCOPIC FEATURES:

fruitbody a stroma, circular to irregular in outline, 1–4(–12) mm diameter; broadly att ached but margins free; plane to slightly wrinkled, very dark green to black; ostiolar openings appearing as small cracks in the stroma surface; no reaction to KOH, dark green all over with lactic acid; fruity odor in culture. MICROSCOPIC FEATURES: ascospores 2-celled, distal part 3.3–3.8 × 2.7–3.3 μm, proximal part 3.4– 4.0 × 2.7–3.2 μm, spinulose, disarticulating early to give 16 monomorphic part-spores, (1.7–)2.7– 4.2(– 6.0) μm, subglobose, linear in the ascus, hyaline; asci (35–)60–80(–100) × (3–)4– 6(– 9) μm, cylindrical; perithecia 170–185 × 105–135 μm; conidiophores with a conspicuous main axis from which solitary phialides arise for a considerable distance; phialides 5.4–7.2 μm long; conidia 3– 5(–11) × 2–3(–4) μm, narrowly elliptic to oblong, smooth, green. OCCURRENCE: solitary to densely gregarious, individual stromata overlapping each other on bark of hardwoods, anamorph cosmopolitan on a diversity of inorganic and organic substrata, including humans with suppressed immune systems; anamorph widespread in North America, teleomorph known from New York to Louisiana, western Europe; not common. COMMENTS: Th is species, fi rst described from Pennsylvania (teleomorph) and Ohio (anamorph), was believed to be widespread globally, but Sam-

uels, Petrini, and Manguin (1994) demonstrated that the tropical collections were a closely related but distinct species, Hypocrea jecorina Berkeley and Broome, and that the Chinese collections were a third closely related but as yet unnamed species. The anamorph, Trichoderma citrinoviride Bissett (not illustrated), is distinctive, with short, rarely rebranched side-branches and cylindrical to narrowly flask-shaped single phialides arising from the main axis, with smooth-walled elliptic to oblong conidia (Samuels 1998). Th is subclade of Hypocrea is noteworthy both for producing human mycoses and for providing source material for important tools for fighting plant disease. Hypocrea schweinitzii is unusual in being able to grow and sporulate at 40 ºC, making it dangerous for humans with a suppressed immune system. When still immature, members of the Hypocrea lixii Patouillard / Trichoderma hazarianum Rifai species complex are easily confused with Hypocrea schweinitzii. At maturity the stromata of members of the Hypocrea lixii complex are a distinctive dark green to almost black, with litt le visible differentiation in color between the surface of the stromata and the perithecia, and members of the complex have green ascospores (compared to hyaline ascospores in Hypocrea schweinitzii). Th is cosmopolitan species complex is found on wood, bark, and fungi, especially polypores.

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Hypocrea schweinitzii

Hypocr ea

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Hypocrea strictipilosa P. Chaverri and Samuels Anamorph: Trichoderma strictipile Bissett fruitbody a pulvinate or lenticular stroma, centrally att ached, 0.7–4 mm × 0.5–1.5 mm; surface smooth with numerous fi ne, distinct, ostiolar dots, white to pale yellowish when young, yellow to light brown at maturity, sometimes fi nally reddish brown, not changing color in KOH, ostiolar openings at fi rst brownish then turning greenish from the spores; basal part

MACROSCOPIC FEATURES:

pale orange and often like a broad stalk; anamorph mainly in association with young stromata that are eff use, powdery, dark green. MICROSCOPIC FEATURES: ascospores with dimorphic cells, distal part 4– 6.5 × 4–5.5 μm, subglobose or wedge-shaped, proximal part 4.5–8 × 3.5–5 μm, sometimes subglobose, usually oblong or wedgeshaped, often slightly longer in ascus base, ver-

Hypocrea strictipilosa

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ruculose, uniseriate, yellowish green; asci 90–135 × 5–7.5 μm, cylindrical, stalk 6–36 μm long, 8-spored with spores disarticulating to yield 16 part-spores per ascus; perithecia 135–235 × 90–190 μm, flaskshaped or globose; ostioles 50–70 μm long × 25– 60 μm wide at apex, projecting 0–30 μm, wall yellow; conidiophores a central axis with lateral branches terminating in a whorl of phialides; conidia from phialides 2.5–8 × 2.5–5 μm and from conidiophores 3.5–11.5 × 2.5–5.5 μm, elliptic, smooth, green. OCCURRENCE: on Ascomycota, Basidiomycota, and wood of deciduous and coniferous trees; eastern North America, northeastern Europe; whenever fungi are fruiting; common. COMMENTS: Hypocrea aureoviridis f. macrospora Yoshimura is a synonym. Th is is the most common green-spored Hypocrea in north-temperate zones (Chaverri et al. 2003; Chaverri and Samuels 2003). It is one of several Hypocrea species usually identified as Hypocrea gelatinosa (Tode) Fries, a greenspored species known so far only from Europe. The flat, smooth, nonwaxy surface of Hypocrea strictipilosa is less pigmented than the green-spored H. chlorospora Berkeley and M. A. Curtis (distal ascospores 5.0–5.2 × 4.7–5.2 μm, and proximal ascospores 5.0–5.5 × 4.5 4.7 μm). Hypocrea sinuosa P. Chaverri and Samuels and Hypocrea thelephoricola P. Chaverri and Samuels (not illustrated) are two other members of the highly colored H. chlorospora group. Hypocrea thelephoricola P. Chaverri and Samuels is at fi rst white, then dotted with bright yellow projecting perithecia that turn green-

ish as the verrucose olive to dull green spores mature (distal part-spore plump, 3.3–5.2 × 3.0–4.7 μm, proximal cell 3.0– 6.4 × 2.6–4.2 μm). When young, the bright yellow perithecia of Hypocrea thelephoricola resemble the pale yellow perithecia of Hypocrea chlorospora Berkeley and M. A. Curtis; in age it is more similar to Hypocrea strictipilosa, but its growth on Steccherinum ochraceum permits easy identification, which is confi rmed by the peculiar orange reaction of the perithecial wall to KOH while the ostiole remains hyaline. Hypocrea sinuosa P. Chaverri and Samuels is widely distributed and locally common, distinguished when fresh by a citrine color and prominent perithecia (ascospores dull green to olive, distinctly verrucose, distal part 4.0– 6.5 × 3.5–5.8 μm). In dry weather, or when eaten by mites, the stroma can be orange rather than yellow. After repeated drying and rewett ing, Hypocrea sinuosa looks very much like H. strictipilosa. It can also be mistaken for Hypocrea ceracea P. Chaverri and Samuels (not illustrated), another North American look-alike to the European H. gelatinosa. Immature stromata of Hypocrea ceracea are waxy, glassy-translucent, and pale yellowish to orangish, with litt le contrast between the perithecia and the stroma. Hypocrea chromosperma Cooke and Peck grows on decaying wood (1–1.2 mm diameter stromata generally clustered in large numbers), the ostioles are slightly prominent (distal part ascospores 4.2–4.5 × 4.0–4.2 μm, proximal part ascospores 4.2–4.5 × 3.5–3.7 μm, pale yellow to brownish when mature).

Hypocr ea

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Hypocrea sulphurea (Schweinitz) Saccardo fruitbody an eff use stroma almost completely covering the host; from 1 × 1 mm up to 7 × 3 cm, sometimes vivid yellow, usually light yellow to grayish yellow; reaction to KOH very slight, the stroma sometimes becoming light orange. MICROSCOPIC FEATURES: ascospores dimorphic, hyaline, distal part obovate to subelliptic, 4.5–7.5 × 4.5–8 μm, proximal part elliptic to subcylindrical, 4.5–8.5 × 3– 6.5 μm; asci 80–150 × 4.5– 8 μm, tip slightly thickened; perithecia 130–270 × 90–170 μm. MACROSCOPIC FEATURES:

on decorticated wood with Exidia species, sometimes without evidence of any Exidia; North America, Europe, and Japan; common. COMMENTS: The anamorph of this distinctive and common species is in the Trichoderma sect. Hypocreanum (Overton, Stewart, and Geiser 2006). Its closest North American relative is Protocrea (Hypocrea) farinosa (Berkeley and Broome) Petch (see comments under Hypomyces aurantius, p. 315). OCCURRENCE:

Hypocrea sulphurea

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Hypocrea viridescens (Horne and Williamson) Jaklitsch and Samuels Anamorph: Trichoderma viridescens (Horne and Williamson) Jaklitsch and Samuels fruitbody pulvinate, turbinate, or discoid, broadly att ached to substrate, margins free, lacking a stalk, 1–5 mm long × 0.5– 1.5 mm high, sometimes aggregated into lines along wood fibers, downy at fi rst, covered with yellowish to rust-colored hairs, semi-eff used, velutinous, with white mycelial margin that is usually dark at maturity, circular to irregular, smooth to granular, white becoming white with yellow ostiolar open-

MACROSCOPIC FEATURES:

ings, frequently becoming yellowish to pale ochraceous, light brownish, or yellow to orange to rustbrown, mostly dark reddish brown when mature, often with the anamorph in evidence as well. MICROSCOPIC FEATURES: ascospores 3.5– 7.5 × 3– 5 μm, verrucose with warts about 0.5 μm wide and high, dimorphic, distal subglobose to oval, some slightly tapered at upper end, proximal oblong to wedge-shaped, the lower end broadly rounded;

Hypocrea viridescens

Hypocr ea

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asci 55–120 μm including stalk of 5–25 μm, cylindrical, hyaline; perithecia 140–330 × 100–250 μm, flask-shaped, elliptic to globose; ostiolar canal 60–120 μm long × 30– 60 μm wide; conidia 3– 5.5(–8.5) × 2.5–4.5 μm, subglobose to elliptic, grossly warted; chlamydospores present or absent, (2.5–)8–12(–18) μm, subglobose. OCCURRENCE: solitary to gregarious to aggregated on all types of wood, also on polypores and agarics, leaves, apple cores, waste paper, soil, and numerous other substrates; spring through fall; North America, Europe, Asia, the Pacific Rim; common. COMMENTS: The anamorph appears as citrine to sulphur-yellow cottony patches of mycelium with dark green conidia. Trichoderma viride Persoon, the type for the genus Trichoderma, was long thought to be the common and abundant Trichoderma species worldwide, but is actually rather rare. The teleomorph of Trichoderma viride is Hypocrea rufa (Persoon) Fries (not illustrated), the type for the genus Hypocrea (Jaklitsch et al. 2006). Hundreds of collections of purported Hypocrea rufa can be found in herbaria worldwide. However, examination of the collections reveals considerable differences, and it soon becomes clear that many different species have been collectively called Hypocrea rufa. Most material that has been called Trichoderma viride/Hypocrea rufa actually turns out to be Trichoderma viridescens/Hypocrea viridescens. The two species pairs can be distinguished by very careful study of the conidia, which are globose in Trichoderma viride and subglobose to elliptic in T. viridescens. Hypocrea atro-

viridis Dodd, Lieckfeldt & Samuels (anamorph Trichoderma atroviride P. Karsten) (not illustrated), which is known from Europe and North America, is very similar to Hypocrea viridescens, Hypocrea rufa, Hypocrea petersenii Samuels, Dodd, and Schroers (not illustrated), Hypocrea rogersonii Samuels (not illustrated), and other members of the Viride Clade of Trichoderma, but distinguished by its smooth, green conidia and the coconut odor of its colonies. “All members of the Viride Clade, including Hypocrea viridescens, have virtually indistinguishable stromata. They are characterized by being ‘furry’ at fi rst from short hairs arising from the stroma surface. In these species the ostiolar opening is not visible, though white balls of spores can be seen at the openings. In other species of Hypocrea, the ostiolar openings can be seen with a hand lens” (G. Samuels, personal communication). Hypocrea minutispora B. S. Lu, Fallah, and Samuels is another species sometimes mistaken for Hypocrea rufa. Hypocrea minutispora has light brown to yellowish brown to reddish brown stromata 0.7–2 × 0.5–1.5 mm in diameter, the surface is smooth to creased and marked with slightly raised purple-brown ostiolar openings, the asci are cylindrical with a thickened tip, and the spores are dimorphic to monomorphic with the distal part globose to subglobose, 3.2– 6.5 × 2.7–5.5 μm, and the proximal part subglobose, oblong, elliptical to wedge-shaped, and 3.5– 6.5 × 2.2–5.5 μm. The teleomorph has regularly branched conidiophores with short, broad phialides.

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Hypomyces aurantius (Persoon) Fuckel Anamorph: Cladobotryum varium Nees subiculum floccose to matted, often covering its entire host, occasionally spreading over adjacent substrate; variably colored white to buff, most often yellow-orange to rusty red, turning violet to purplish red in KOH; perithecia formed all over or only in some parts of the subiculum, gregarious or in some specimens cespitose, semi-immersed in the subiculum, pale yellow to orange or red, entirely purplish in 3 percent KOH.

MACROSCOPIC FEATURES:

MICROSCOPIC FEATURES: ascospores(13–)20– 25(–27) × (3–)4– 6(–7.5) μm, fusiform to lanceolate, 2-celled, verrucose (warts less than or equal to 0.5 μm high), acutely apiculate (apiculi 2–4.5 μm long), obliquely uniseriate; asci 100–140 × 6–7 μm, cylindric to clavate, apex thickened, with a pore, 8-spored; perithecia 250–575 × 200–375 μm, spherical to ovate or obpyriform, gregarious; papillae typically not well differentiated, 150–200 μm wide × 75–100 μm high; conidia (8.5–)10–16(–21) × 5–

Hypomyces aurantius

H y pom yces

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7.5(–10) μm, at fi rst round or nearly round, becoming elliptic or oboval, (0-)1-septate, with a basal hilum, often with refractive material at the apex, single and end to end in dry chains; chlamydospores 18–48 × 12–18 μm, oblong-oval, 1–4-celled, constricted at septa, pale brown to reddish brown, thick-walled, wall 1–3 μm thick. OCCURRENCE: found on a wide range of polypores and wood-decaying agarics, or spread onto nearby vegetation; spring through fall; common in both north and south temperate regions, rare in the tropics. COMMENTS: In nature, the anamorph is a profuse, pure white, cottony mat, its powdery appearance due to the conidia. Hypomyces subaurantius Heinrichson is a synonym. There is great variation in general appearance and habitat for this widespread species, as well as in the characteristics of the perithecia and ascospores. Typical features are a bright orange subiculum, orange-red perithecia that turn purple in KOH, 2-celled verrucose, apiculate ascospores that are usually less than 25 μm long, and growth on polypores. Hypomyces rosellus (Albertini and Schweinitz) Tulasne and C. Tulasne has a rose-colored subiculum with dark red perithecia that turn purple in 3 percent KOH and has longer ascospores (> 30 μm) that are conspicuously apiculate and verrucose. Protocrea, Hypocrea, and Hypomyces species are very closely related. Protocrea pallida (Ellis and Everhart) Jaklitsch, K. Põldmaa, and Samuels (= Hypocrea pallida Ellis and Ever-

hart) is also found on polypores and is very similar in appearance to Hypocrea aurantius. It also has a strong pink to violet reaction of the subiculum and perithecia to KOH. It is distinguished by 2-celled spores that are minutely warted, hyaline, and more or less dimorphic (distal part 2.7–4.3 × 2.4–4 μm, proximal part 3–5.5 × 2–3.5 μm), with one large and several small oil drops. The ascospores divide into part-spores. Hypocrea americana, which grows on Fomitopsis pinicola and Piptoporus betulinus, has spores that disarticulate into 16 part-spores per ascus at maturity. Each part-spore is subglobose and measures 2.7–3.7 × 3.2–4.2 μm. The KOH reaction is brownish orange to red. Protocrea farinosa (Berkeley and Broome) Petch has a much less well-developed subiculum and the perithecia often emerge directly from the pores of the host. Its reaction to KOH is at most a faint violaceous tint in contrast to the strong reaction of the Protocrea pallida or Hypomyces aurantius subiculum to KOH. Hypocrea delicatula Tulasne and C. Tulasne had been placed into Protocrea by Petch, but DNA confi rms that it belongs in Hypocrea. Hypocrea delicatula grows on hardwood and soft wood debris, rather than polypores, has a white to cream stromata, semi-immersed to immersed perithecia, and dimorphic spores with the upper part turbinate, often with a small apiculus (4–4.5 × 1.5–2.5 μm) and a lower part wedge-shaped and not apiculate (4– 4.5 × 1.5–2.5 μm).

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Hypomyces cervinigenus Rogerson and Simms Anamorph: Mycogone cervina Ditmar fruitbody at fi rst cottony and later a compacted mat covering a parasitized Helvella; white, later becoming pink to brown in places or overall; not changing color with KOH. MICROSCOPIC FEATURES: ascospores rare, 15– 25 × 1.5–5 μm, fusiform to spindle-shaped, sometimes curved, usually acute at the apices but often with lower end obtuse, median or submedian septum, cytoplasm usually vacuolate, septum often obscured, smooth, obliquely uniseriate in ascus, hyaline; ascus 100–140 × 2–4.5 μm, apex thickened, penetrated by a pore, 8-spored; perithecia rare, 165–385 × 130–265 μm, gregarious to scattered, semi-immersed in the subiculum, buff or light yellow, not reactive to KOH; aleuriospores 13.5– 17.5 μm diameter including ornamentation, round, spiny, thick-walled, joined to a secondary, thinMACROSCOPIC FEATURES:

walled, more or less hemispheric cell, in deposit aleuriospores pinkish buff; conidia 14.0–25.5 × 4.0– 5.0 μm, cylindric-elliptic, smooth, thin-walled, contents granular or with one to several oil droplets, some centrally septate. OCCURRENCE: parasitic on Helvella species, including H. elastica, H. ephippium, and H. vespertina, covering cap of the host, or both cap and stalk (host’s fruitbody becomes decayed); spring, summer, or fall—whenever Helvella species are fruiting; western North America, Europe; anamorph common, teleomorph rare. COMMENTS: The two-celled smooth ascospores (less than 25 μm long) with acute, not apiculate ends, the waxy white to pale buff perithecia, and growth on Helvella are all distinctive features.

Hypomyces cervinigenus

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Hypomyces chlorinigenus Rogerson and Samuels Anamorph: Sepedonium chlorinum (Tulasne) Damon the anamorph appears fi rst, is white and cottony, partially or completely covers its host, produces conidia and becomes pale yellow, then yellow to yellow-brown and powdery as darker aleuriospores are produced; the teleomorph appears next and consists of globose to obovoid, buff to brown perithecia containing asci and ascospores, producing a roughened, bumpy texture; unreactive to KOH. MICROSCOPIC FEATURES: ascospores 7.5–12 × 2.5– 5 μm, elliptic-fusiform, 2-celled, smooth to fi nely verrucose, hyaline; asci 70– 90 × 3–5 μm, cylindrical; conidia 10–11 × 4–5 μm, ovoid to ellipticcylindric, 1-celled, hyaline; aleuriospores 35–45 × 15–18 μm, cylindric, longitudinally ridged, yellow to yellow-brown. MACROSCOPIC FEATURES:

on various boletes; summer and fall, also winter in the southern part of its range; eastern North America; fairly common. EDIBILIT Y: inedible; parasitized boletes should not be eaten. COMMENTS: The photograph shows the anamorph parasitizing specimens of Boletus pallidus. Hypomyces boletiphagus Rogerson and Samuels (not illustrated), another eastern species that att acks a wide range of boletes, is very similar in appearance but the 1-septate, smooth to verruculose spores are longer (10–13 × 2–4 μm), and KOH applied to the affected surfaces turns the surface yellow to red to blackish while Hypomyces chlorinigenus shows no color change. OCCURRENCE:

Hypomyces chlorinigenus

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Hypomyces chrysospermus Tulasne Anamorph: Sepedonium chrysospermum (Bulliard) Fries Bolete E ater, Bolete Mold, Golden H y pom yces

fruitbody consisting of two stages: the anamorph appears fi rst, is white and moldy and partially or completely covers its host, produces conidia and becomes yellow to golden yellow and powdery as yellow to yellowbrown aleuriospores are produced; the sexual stage consists of globose to flask-shaped, orange-yellow to red-brown perithecia containing asci and ascospores, producing a roughened, pimpled texture. MICROSCOPIC FEATURES: conidia 10–30 × 5–12 μm, elliptic, 1-celled, smooth, hyaline; aleuriospores 10–25 μm, globose, thick-walled, prominently verrucose, yellow to golden yellow or yellow-brown; ascospores (15–)22–25(–30) × 4–5(– 6) μm, 2-celled, fusiform to lanceolate, hyaline; asci 100–140 × 5– 8(–10) μm, cylindrical. OCCURRENCE: on various boletes and gilled fungi; summer and fall; widely distributed in North America; common. EDIBILIT Y: inedible; parasitized boletes should not be eaten. COMMENTS: The teleomorph is seldom encountered. Hypomyces microspermus Rogerson and Samuels (anamorph Sepedonium microspermum Besl) is MACROSCOPIC FEATURES:

nearly identical, but it has smaller ascospores that measure (8–)10–15 × 2.3–4 μm and 8–15 μm globose aleuriospores. According to Rogerson and Samuels (1989), Hypomyces microspermus is the most frequently collected boleticolous species in North America. Hypomyces melanocarpus Rogerson and Mazzer, with a white to buff subiculum, differs from most other Hypomyces species on boletes by having olivaceous to dark red to black perithecia and ascospores (mostly 30–35 × 7.5–10 μm) that remain 1-celled. Hypomyces chrysospermus has orange-yellow to red-brown perithecia and Hypomyces tulasneanus Plowright (anamorph Sepedonium tulasneanum Saccardo) (not illustrated) has yellow to light brown perithecia. Both Hypomyces chrysospermus and Hypomyces tulasneanus have unequally 2-celled ascospores at maturity. Hypomyces completus (G. Arnold) Rogerson and Samuels (anamorph Sepedonium brunneum Peck) also has dark perithecia (pale to dark brown to black) and 1-celled ascospores (mostly 35–40 × 4– 6 μm) and is further distinguished by a yellow-brown to black subiculum.

Hypomyces chrysospermus

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Hypomyces hyalinus (Schweinitz) Tulasne and C. Tulasne A m a nita Mold

distorted Basidiomycete host 10–30 cm high, up to 7 cm thick, fi rm, solid, columnar to clavate, usually quite phallic, chalky white, yellowish or pinkish to pale orange, roughened where perithecia develop; perithecia pallid to light orange or brownish, immersed in the subiculum except for the papilla, no reaction to KOH; host cap, gills, and other structures rarely discernible. MICROSCOPIC FEATURES: spores 15–20 × 4.5– 6.5 μm, fusiform, 2-celled, septum submedian, prominently verrucose, ends apiculate, hyaline; asci 110–130 × 4– 6 μm, cylindric; perithecia 250– 325 μm tall × 180–210 μm wide, ovate to obpyriform, papilla 125 μm high × 60–100 μm wide at apex, broadly truncate; anamorph not known. OCCURRENCE: on several species of Amanita, especially frequent on Amanita rubescens; early summer and fall; eastern North America, west along the Rocky Mountains, and the Pacific Northwest; common and often abundant. EDIBILIT Y: not recommended since parasitizing an Amanita. COMMENTS: Apiocrea hyalina (Schweinitz) Sydow and P. Sydow and Sphaeria hyalina Schweinitz are synonyms. Although the identification of the host mushroom is sometimes evident, in some cases the host mushroom’s identity can only be presumed; reddish-staining specimens are most likely Amanita rubescens. MACROSCOPIC FEATURES:

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Hypomyces lactifluorum (Schweinitz: Fries) Tulasne Lobster Fu ngus, Lobster Mushroom

fruitbody a fi rm layer of roughened or pimpled tissue covering the entire surface of its distorted host species, the gills of which are reduced to blunt ridges; overall shape often like an inverted pyramid with a depressed top; surface pale yellow-orange to bright orange, aging orange-red to purple-red, occasionally fading to pink; perithecia deep orange to reddish purple, usually darker than the surrounding tissue, immersed except for papilla, turning purple with KOH. MICROSCOPIC FEATURES: spores 30–50 × 4.5–8 μm, fusoid or shaped like caraway seeds, prominently verrucose and apiculate (verrucae 1–1.5 μm high, confluent; apiculi 4.5–7.5 μm long, acute, straight or curved), 1-septate, partially overlapping in the asci, hyaline; asci 200–260 × 5–10 μm, MACROSCOPIC FEATURES:

long-cylindric, apex thickened and with a pore, 8-spored; perithecium 400– 600 × 200–450 μm, papilla averaging 120–220 high × 120–200 μm wide; anamorph unknown. OCCURRENCE: usually found on Russula brevipes, rarely on Lactarius species in the piperatus complex; summer and fall; throughout North America and known only from North America; common. EDIBILIT Y: this is a popular edible fungus, even though the identity of the host species is usually undetermined. COMMENTS: Hypomyces lactifluorum is the type for the genus Hypomyces (Fries) L. R. Tulasne. Hypomyces insignis Berkeley and M. A. Curtis and Hypomyces purpureus Peck are synonyms. Twelve other species of Hypomyces occur on gilled fungi, mostly on members of the Russulaceae but also on Ama-

Hypomyces lactifluorum

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nita, Crepidotus, Entoloma, and Pholiota. Several species are also found on boletes, and some on polypores and thelephores. Hypomyces macrosporus Seaver is distinguished by white perithecia and subiculum that do not react to KOH but is microscopically indistinguishable from Hypomyces lactifluorum. Sometimes both species can occur on a single specimen, giving it an orange and white streaked appearance. Hypomyces macrosporus may be an albino form of Hypomyces lactifluorum (Rogerson and Samuels 1994). Hypomyces banningiae Peck as “banningii” is very similar to Hypomyces

macrosporus Seaver in that the subiculum of both is similarly colored white to buff and is unreactive to KOH. Both have large ascospores. Hypomyces banningiae is an eastern species found on highly decomposed Lactarius species where the host becomes fi rm. Hypomyces banningiae has aseptate spores (28–40 × 4.5–5.5 μm, fusiform or lanceolate, 2–4 oil droplets, smooth to verruculose with bluntly thickened ends). The spores of Hypomyces macrosporus and Hypomyces lactifluorum have a median septum and are prominently verrucose and apiculate with apiculi 3–7 μm in length.

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Hypomyces lateritius (Fries) Tulasne and C. Tulasne Anamorph: Acremonium tulasnei G. R. W. Arnold subiculum a dense white cottony mycelium deforming the gills of Lactarius species; appears restricted to the gill surfaces and makes the host fi rm, initially white to lemonyellow, later buff, yellowish orange or tawny and at full maturity brick-red and brown to reddish black; KOH negative. MICROSCOPIC FEATURES: spores mostly 15–21 × 3.5–5 μm, fusiform, naviculate or lanceolate, 1-celled but with inclusions that appear to be a median septum, verrucose (warts less than 0.5 μm high), acutely apiculate (apiculi mostly 1.5–3 μm), partially overlapping in ascus; asci 90–150 × 4–8 μm, apex thickened and penetrated by a pore, 8-spored; perithecia 300–470 × 170–360 μm, ovate to obpyri form, usually immersed in the subiculum with papilla protruding, dark amber to reddish brown, becoming lighter and yellowish in KOH; papilla 120–150 high × 100–175 μm wide. OCCURRENCE: on gills of Lactarius species; summer and fall or whenever Lactarius species are present; worldwide; common. MACROSCOPIC FEATURES:

Rogers and Samuels (1994) report that Hypomyces lateritius is a variable species that may ultimately be proven to be a species complex. When found on species in the Lactarius camphoratus complex, the ascospores are slightly larger and the aborted gills are yellowish. Hypomyces lithuanicus Heinrichson-Normet (not illustrated) is possibly the same as Hypomyces thiryanus Maire and parasitizes Lactarius torminosus and Lactarius pubescens. Hypomyces lithuanicus is distinguished by a cream-ochre to cinnamon subiculum covering the deformed gills and by perithecia that turn reddish brown in KOH (in the apical portion only). Hypomyces torminosus Durieu (not illustrated) differs from Hypomyces lithuanicus by perithecia that remain very brown in KOH. Hansen and Knudsen (2000) list both Hypomyces lithuanicus and Hypomyces torminosus as synonyms of Hypomyces spadiceus Fries ex Cooke (not illustrated). In contrast to the purple-brown to brown reaction of the perithecia in the Hypomyces spadiceus complex, the perithecia of Hypomyces lateritius become lighter and yellowish in KOH.

COMMENTS:

Hypomyces lateritius

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Hypomyces luteovirens (Fries) Tulasne and C. Tulasne Gr een Lobster, Russu l a Mold

a fi rm layer of roughened or pimpled tissue covering and somewhat distorting the gills and upper stalk of its host species, rarely spreading onto the cap; at fi rst yellowish to bright yellow, becoming yellowish green to dark green, and fi nally blackish green MICROSCOPIC FEATURES: spores 28–35 × 4.5– 5.5 μm, fusiform, aseptate, nearly smooth to prominently verrucose, hyaline; asci 160–200 × 5–8 μm, cylindric; perithecium 380–485 μm tall × 180– 290 μm wide, broadly ovate to obpyriform, immersed except for papilla, papilla 96–120 μm high, 180–230 μm wide at apex, truncate or obtuse, no reaction to KOH; anamorph unknown. OCCURRENCE: on various species of Russula; summer and fall; widely distributed in North America; occasional to fairly common. MACROSCOPIC FEATURES:

Since this parasite rarely att acks the cap surface of the host mushroom, it is usually detected by picking and inspecting the underside of specimens of Russula. No anamorph is known. Byssonectria luteovirens (Fries) Z. Moravec, Hypocrea luteovirens (Fries) Berkeley and Broome, and Peckiella luteovirens (Fries) S. Imai are synonyms. As of 2012, Hypomyces viridis (Albertini and Schweinitz) Berkeley is still in use, though Rogerson and Samuels (1994) treat it as a synonym of Hypomyces lutovirens. Hypomyces tulasneanus Plowright (not illustrated) is a name that has been applied to H. luteovirens, but H. tulasneanus was originally and correctly described as growing on Boletus, not on Russula. COMMENTS:

Hypomyces luteovirens

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Hypomyces ochraceus (Persoon: Fries) Tulasne Anamorph: Cladobotryum verticillatum (Link) S. J. Hughes fruitbody a subiculum on wood, moss, or the ground; up to 5 cm in extent; initially white, then pinkish cream or buff and fi nally apricot, remaining white or lighter at the margin; no reaction to KOH. MICROSCOPIC FEATURES: spores 27–45 × 5–8 μm, fusiform, often curved, nonseptate or septum median, prominently verrucose (warts 0.5–1.2 μm high), acutely apiculate (apiculi 0.5–1.2 μm long); partially overlapping in ascus; asci 170–180 × 6– 8 μm, apex thickened, penetrated by a pore; perithecia 260–360 × 240–250 μm, at fi rst immersed in subiculum, later half-free, yellowish orange or amber to reddish orange; papilla 90–100 × 60–80 μm, prominent; conidia 11–24 μm long, broadly elliptical, unicellular. OCCURRENCE: appears fi rst as the anamorph Cladobotryum verticillatum (Link) S. Hughes on a host agaric (usually members of the Russulaceae, also on boletes) that is so completely destroyed that the species appears to occur on detritus by the time of perithecial formation (in 2–3 weeks); summer and fall; Europe and eastern North America, also Oregon; common. MACROSCOPIC FEATURES:

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Hypomyces ochraceus

Hypomyces armeniacus Tulasne and C. Tulasne is a synonym. Th is fungus is eff used like species of Hypocrea citrina Fries (not illustrated; see Canham [1969] and Overton et al. [2006] for descriptions of H. citrina varieties) and Hypocrea sulphurea (p. 312).

COMMENTS:

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Hypomyces porphyreus Rogerson and Mazzer subiculum sparse, cottony, partly to completely covering the host pileus; brick-red; KOH unreactive or the base or entire perithecium turns purple. MICROSCOPIC FEATURES: spores 20–30 × 3.5– 5.0 μm, fusiform to narrowly elliptic, with or without a median septum, each end with a broad apiculum, verrucose, warts 0.5–1.5 μm high and round, spores overlapping by about one half their length; asci 125–190 × 4–5 μm, long cylindrical, apex thickened and with a pore, 8-spored; perithecia 300– 395 μm tall × 220–275 μm wide, brown with red tints, dark brick-red or chestnut; papilla 40– 60 μm high × 150 μm wide; anamorph not known. OCCURRENCE: on Entoloma strigosissimum (Rea) Noordel = Leptonia strigosissima (Rea) P. D. Orton and on undetermined agarics; summer and fall; Michigan, the Netherlands, and Norway; rare. MACROSCOPIC FEATURES:

Rogerson and Mazzer (1971) suggest an alliance between Hypomyces porphyreus and H. cervinigenus (a smooth-spored species on Helvella; p. 317) and Sporophagomyces chrysostomus (Berkeley and Broome) K. Põldmaa and Samuels = Hypomyces chrysostomus Berkeley and Broome. Sporophagomyces chrysostomus is found in eastern North America on often strongly disintegrated polypores where it forms a brush-like white growth over the pore surface and becomes quite showy as the white mycelial growth catches quantities of the brown basidiospores from the polypore. The smooth, acute to obtuse spores are 10–15 × 3–4 μm with one median septum, and the perithecia are pale yellow, turning olivaceous to reddish.

COMMENTS:

Hypomyces porphyreus

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Hypomyces tremellicola (Ellis and Everhart) Rogerson Anamorph: a Verticillium species subiculum fi rm, white, completely covering the fertile layer and often the entire fruitbody of the host; perithecia of the teleomorph buff or reddish brown; subiculum no reaction to KOH. MICROSCOPIC FEATURES: ascospores 7–13 × 3– 4 μm, elliptic, 1-septate, septum median, spore wall verrucose, ends not apiculate; asci 60–75 × 4–5 μm, apex thickened or not, penetrated by a pore; perithecia 450– 600 × 300–500 μm, cespitose, superficial or immersed in subiculum except for papillae; papillae 220–225 μm high × 200 μm at base, tip obtuse; conidia (4–)5–8(–12) × 2–3(–5) μm, cylindrical to elliptic, basal hilum absent, aseptate, rarely 1-septate. OCCURRENCE: mainly on species of Crepidotus but also on Pleurotus and Polyporus species, sometimes migrating to nearby bark; spring through MACROSCOPIC FEATURES:

fall; known from central and eastern North America, northern South America, and middle Europe; anamorph common, teleomorph common to uncommon. EDIBILIT Y: unknown, not recommended since host unrecognizable. COMMENTS: Nectriopsis tremellicola (Ellis and Everhart) W. Gams is listed by Species fungorum as the currently accepted name. Th is species is intermediate in form between Hypomyces, Hypocrea, and Nectria. Hypocrea avellanea Rogerson & Carey, which grows on Marasmius subnudus Ellis ex Peck in summer to early fall in northeastern North America, differs by having spores that disarticulate at maturity. Hypocrea avellanea subiculum covers and deforms all or most of the gills of the host mushroom, is overall wood-brown to avellaneous, has perithecia partially or completely em-

Hypomyces tremellicola

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bedded in the subiculum, separate or densely compacted, orange, dresden brown at maturity, with apices darker. The part-spores are hyaline, smooth to spinulose, and dimorphic, the upper part typically globose and measuring 2–4 × 2–3.5 μm, lower part usually truncate elliptic and measuring 2.5– 5 × 2–3.5 μm, both parts usually becoming elliptic after discharge. The asci are cylindric-clavate, 50– 100 × 3–5 μm, and 8-spored, the ascospores are uniseriate, with an apex slightly thickened and with an obscure pore. The perithecia are 100–300 × 100–200 μm, globose-ovate to globose-papillate, with apices that turn violet-red in 3 percent KOH. The anamorph is Verticillium-like, with elliptic conidia, measures 3– 9 × 1.5–5 μm, and is hyaline.

Similar Hypocrea species that are eff used over a fungus host were once placed in Protocrea. Protocrea is now known to be distinct from Hypocrea ( Jaklitsch, Põldmaa, and Samuels 2008), but Hypocrea avellanea forms a clade with the European species Hypocrea delicatula Tulasne and C. Tulasne within Hypocrea. Most species of Trichoderma/ Hypocrea are not host specific, but a few species, including Hypocrea avellanea, Hypocrea stromatica J. L. Bezerra, J. C. B. Costa, and C. N. Bastos (not illustrated) (on Moniliophthora perniciosa), and Trichoderma aggressivum Samuels and W. Gams (not illustrated) (on commercial Agaricus), are only found on specific agaric hosts. See also Nectriopsis tubariicola (p. 335).

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Hypoxylon fuscum (Persoon) Fries Anamorph: Virgariella-like stromata soft , fleshyleathery, usually hemispheric to cushion-shaped, sometimes flat or with inconspicuous perithecial mounds; 0.1– 6 cm × 0.1–3 cm and 0.4–2.5 mm thick; surface fairly even, with perithecial elevations usually not prominent, light purplish when very young, brown-vinaceous or dark vinaceous, becoming blood-colored, chestnut, and fuscous in age; buff to dull reddish brown granules immediately beneath the surface and between the peri-

MACROSCOPIC FEATURES:

thecia, with amber to olivaceous KOH-extractable pigments; dark brown to blackish brown tissue below the perithecial layer, inconspicuous to 0.15 cm thick; perithecia black, evenly embedded in a superficially red-brown stroma. MICROSCOPIC FEATURES: ascospores 8–20 × 4– 8 μm, elliptic-inequilateral, with narrowly rounded ends, unicellular, somewhat obliquely uniseriate, with a slightly sigmoid full-length germ slit, perispore dehiscent in 10 percent KOH, smooth or

Hypoxylon fuscum

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with inconspicuous coil-like ornamentation, epispore smooth, brown to dark brown; asci 85–200 × 6– 9(–12) μm, the spore-bearing part 60–115 μm long, stalk 20–100 μm long, with discoid 0.5–2 μm tall × 1.2–3.5 μm wide amyloid apical ring; perithecia 100–300(–400) μm across × 200–500 μm high, spherical to obovoid, ostioles lower than stromatal surface; conidia 4– 6 × 2–2.5 μm, elliptic, smooth, hyaline on Virgariella-like anamorph of hyaline hyphae with conidiogenous cells 20–40 × 1.5–2 μm. OCCURRENCE: gregarious to solitary on dead limbs, bark, and barkless wood of a wide range of hardwoods; year-round; throughout North America, Europe, Asia, and the Philippines; common. COMMENTS: The list of synonyms is vast. Hypoxylon howeanum Peck as “howeianum” is another species found worldwide on a wide range of hardwoods and has an orange or brick-red surface darkening to almost black with inconspicuous to conspicuous perithecial mounds. It has red granules immediately beneath the surface and white granules between perithecia, and tissue below the perithecial layer is satiny black with one or more faint concentric zones, with orange or rust KOH-extractable pigments. Texture is fleshyleathery to woody, and never carbonous. Hypoxylon howeanum ascospores are 6– 9.5 × 3–4.5 μm, elliptic-inequilateral, unicellular, and smooth, with narrowly rounded ends, an indistinct sporelength sigmoid germ slit, and perispore dehiscent in 10 percent KOH. The epispore is smooth and brown to dark brown. The ascus is 80–130 × 4.5– 6.3 μm, the spore-bearing upper part 50–70 μm, with a discoid amyloid apical ring 0.4– 0.8 μm high × 1.2–2 μm broad. Perithecia measure 100–300 ×

200–400 μm, are spherical to obovoid, with an ostiole lower than or at the same level as the stromatal surface. Conidia on the surface of the stroma are 4– 6 μm, round to oblong, and hyaline. Hypoxylon multiforme Fries var. australe Cooke is one of many synonyms. Hypoxylon fragiforme (Pers.) J. Kickx f. = Hypoxylon coccineum Bulliard, commonly known as beech hypoxylon or red cushion hypoxylon, occurs only on beech trees, at least as far west as Michigan, and is the only Hypoxylon species other than H. howeanum with two colors of granules. Hypoxylon fragiforme has inconspicuous perithecial mounds, is 1.5–16 mm wide, dry, hard, conspicuously roughened with ostioles, and grayish white at fi rst, then salmon-pink, becoming cinnamon to brick-red at maturity, and fi nally blackening in age. The interior is a dark brown to black layer of embedded perithecia with a hollow center. Ostioles are lower than or at the same level as the stromatal surface. Spores are (10–)11–15 × 5–8 μm, elliptic with one side flattened, a spore length germ slit, sometimes with 1–3 oil drops, smooth, dark brown, and with a perispore dehiscent in 10 percent KOH. Asci are 155–175 × 6.3–8 μm with a discoid 0.8–1.2 μm long × 2.2–2.8 μm amyloid pore. Perithecia are 0.5– 0.6 mm tall × 0.2 – 0.4 mm wide. The anamorph of Hypoxylon fragiforme and Hypoxylon howeanum is Nodulisporium-like. The genus Annulohypoxylon is distinguished by the presence of black carbonaceous tissue surrounding the perithecia. Daldinia species have alternating light and dark bands observable when cut through the middle.

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Hypoxylon rubiginosum var. rubiginosum (Persoon) Fries Anamorph: Nodulisporium-like fruitbody variable in size and shape but often cushion-shaped; individual fruitbodies usually fusing and forming crustlike patches up to 10 cm or more in diameter; surface uneven, often furrowed, brick-red to reddish brown or purplish brown, in age wearing away and exposing a black interior. MICROSCOPIC FEATURES: spores 9–12 × 4–5.5 μm, irregularly elliptic, inequilateral with one side flattened and narrowly rounded ends, smooth or with inconspicuous coil-like ornamentation, brown to dark brown, straight spore-length germ slit, perispore dehiscent in 10 percent KOH; asci 100–170 × 5.5–8 μm; anamorph (at margins of young stromata) Nodulisporium-like, fi nely roughened, yelMACROSCOPIC FEATURES:

lowish to pale brown, conidiogenous cells hyaline, smooth, 10–25 × 3–4 μm; buff-colored with yellowish conidia 5– 6 × 3–4 μm. OCCURRENCE: in groups or confluent patches on trunks, logs, and stumps of hardwoods; yearround; southeastern United States; occasional to fairly common. COMMENTS: Hypoxylon fragiforme is also brick-red but forms round to cushion-shaped fruitbodies, not crust-like patches. Hypoxylon ferrugineum G. H. Ott h (synonym Hypoxylon rubiginosum var. ferrugineum [G. H. Ott h] J. H. Miller) forms brick-red to hazel crust-like patches on eastern hardwoods, and the spores are larger (13.5–17 × 6.5–8.5 μm).

Hypoxylon rubiginosum var. rubiginosum

H y pox y lon

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Kretzschmaria deusta (Hoff mann) P. M. D. Martin stromata 4.5– 9 cm wide, 0.8–4 mm thick, spreading, crust-like, forming extensive sheets up to 40 cm or more across that are irregular in shape; the anamorph appears fi rst with a grayish white, soft, powdery surface with multiple lobes; as the teleomorph develops the surface becomes hard, black, fi nely roughened, and resembles burnt wood; flesh soft and white at fi rst, becoming black and britt le in age; perithecia embedded just beneath the surface in a single layer. MICROSCOPIC FEATURES: spores (28–)32–34(–36) × 7–10(–12) μm, irregularly elliptic with one side flattened, with a straight germination slit much MACROSCOPIC FEATURES:

Ca r bon Cushion

shorter than the length of the spore, smooth, uniseriate, dark brown; asci 410–480 × 12–15 μm, the spore-bearing part 170–210 μm long, cylindrical, amyloid apical pore, 8-spored; paraphyses fi liform, tips not enlarged; conidia 5– 6.5 × 2–3 μm, smooth, hyaline. OCCURRENCE: on stumps and roots of hardwood trees; year-round; widely distributed in North America; common. COMMENTS: Ustulina deusta (Hoff mann) Lind is a synonym. Mature specimens are easily overlooked. Th is species is essentially a resupinate Xylaria (Rogers 1985).

Kretzschmaria deusta

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Lasiosphaeria spermoides (Hoff mann) Cesati and De Notaris ascomata a dense mass, often several centimeters in diameter, of superficial, subglobose to ovoid perithecia; individual perithecia 0.6–1 mm wide, irregularly rounded to oval or cushion-shaped with slightly raised beaks or expanded ostioles, brownish black to black, hard and carbonaceous; ostioles sometimes tufted with a mass of whitish spores. MICROSCOPIC FEATURES: spores 20–27 × 3.5– 4.5 μm, allantoid, smooth, hyaline; asci cylindric, 8-spored; paraphyses uniformly fi liform. OCCURRENCE: in very dense clusters on decaying wood or bark of conifer or hardwoods, often on stumps; year-round; northeastern North America, distribution range yet to be determined; common. COMMENTS: Ruzenia spermoides (Hoff mann) O. Hilber is a synonym. Lasiosphaeria ovina (Persoon) Cesati and De Notaris is very similar and occurs throughout eastern North America, west to Colorado. When young, its perithecia are clothed MACROSCOPIC FEATURES:

externally with a fi ne white or yellowish tomentum, except for the ostioles, which appear as black dots. At maturity the perithecia are brownish black to blackish, hard, and carbonaceous. Microscopically, its spores are longer and measure 35– 50 × 3–5 μm, are cylindric or vermiform, usually abruptly curved near the lower end, simple or indistinctly septate, and hyaline. Lasiosphaeria lanuginosa (H. Crouan and P. Crouan) A. N. Miller and Huhndorf is a species complex found on decorticated deciduous branches in North America, South America, and Europe, with 360–740 μm tall × 360– 640 μm wide papillate ascomata covered with a dense white to yellowish or pale pinkish tomentum and 33– 60 × 2.5–5 μm ascospores that are cylindrical with a septum and developing rounded, swollen ends with lash-like bipolar gelatinous appendages 7–40 μm long (Miller and Huhndorf 2004a, 2004b).

Lasiosphaeria spermoides

L a siosph a er i a

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Nectria cinnabarina (Tode) Fries Anamorph: Tubercularia vulgaris Tode ex Fries stromata form a base for a dense, cushion-shaped cluster of individual spherical to oval perithecia 0.2– 0.4 mm wide, perithecia bright vermilion to brown-red, darkening in age, on a somewhat lighter-colored stroma; perithecia surface minutely warted, each wart with a small apical, papillate ostiole; conidial stage cushion-like, 1.5–4 mm wide, light orange to pink to bright coral-red. MICROSCOPIC FEATURES: spores 12–19 × 5– 7 μm, elliptic, often curved, with a single septum at the center, often slightly constricted, smooth, irregularly biseriate, hyaline; asci 75– 90 × 9–10 μm, cylindric-clavate, 8-spored; paraphyses absent. OCCURRENCE: scattered or in dense clusters on decaying branches of hardwoods; year-round; northeastern and western North America, distribution limits yet to be determined; common. COMMENTS: Nectria purpurea (Linnaeus) G. W. Wilson and Seaver is a synonym. Neonectria coccinea (Persoon) Rossman and Samuels = Nectria coccinea (Persoon) Fries has smooth perithecia, MACROSCOPIC FEATURES:

Cor a l Spot Fu ngus

grows on bark and wood, especially on scaleinfested beech trees in eastern North America, and has ascospores measuring 12–15 × 5– 6 μm. Neonectria fuckeliana (C. Booth) Castlebury and Rossman is distinguished by growth on conifer bark, has bright red to brownish red smooth perithecia with a distinct ostiole and ascospores 13–16 × 5– 6 μm. Hydropisphaera peziza (Tode: Fries) Dumortier = Nectria peziza (Tode) Fries grows on Polyporus squamosus (Hudson) Fries and Trametes versicolor (Linnaeus) Lloyd as well as on wood and bark. It produces litt le tangerine-colored, globose, thick-walled perithecia (25+ μm thick) that on drying collapse from above to form cups. The perithecia have short orange hairs on the surface that occasionally form triangular fascicles. The ascospores are 12–16 × 5–7 μm, 1-septate, longitudinally striate, and hyaline to a pale straw color. Neither the perithecia nor the stroma of Hydropispharea peziza change color in either lactic acid or KOH, unlike Neonectria species which turn darker in them.

Nectria cinnabarina

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Nectriopsis tubariicola W. Gams Anamorph: Verticillium sp. fruitbody a whitish to ochraceous subiculum that covers the gills of its host and causes gross disfiguration; perithecia pale brown, almost completely immersed in the subiculum and often densely aggregated, flaskshaped, containing asci and spores; no reaction to KOH. MICROSCOPIC FEATURES: spores 12–16 × 3–4 μm, 2-celled, and constricted at the median septum, MACROSCOPIC FEATURES:

fusiform with slightly rounded to truncate ends, fi nely verrucose, hyaline; paraphyses absent. OCCURRENCE: on the gills of Tubaria furfuracea; spring, summer, and fall, also winter in California and the southern parts of its range; widely distributed in North America; occasional. COMMENTS: Fruitbodies of Tubaria furfuracea that are attacked by this fungus often appear to be mummified.

Nectriopsis tubariicola

Nectr iopsis

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Nectriopsis violacea (Fries) Maire Anamorph: Acremonium fungicola (Saccardo) Samuels subiculum white at fi rst, becoming violet to purple and surrounding each perithecium, covering the surface of the aethalia of the host slime mold Fuligo septica; no reaction to KOH. MICROSCOPIC FEATURES: spores (5–)7–8 × 2.5– 3 μm, spinulose, equally 2-celled, not constricted at the septum, obliquely uniseriate with overlapping ends, hyaline; asci 40–75 × 3–5 μm, 8-spored, sessile, apices with a nonchitinoid ring, bases rounded; perithecia 120–390 × 150–310 μm, broadly pyriform; papillae acute, fringed with hairs. MACROSCOPIC FEATURES:

OCCURRENCE: on the slime mold Fuligo septica; all seasons when the slime mold is fruiting; widespread; fairly common. COMMENTS: Samuels (1973) argued that this species is sufficiently similar to Nectria that it should be retained there but in 1999 accepted the name Nectriopsis. It is intermediate between Nectria and Hypomyces. Nectriopsis violacea is very distinctive and widespread, but typically observers think that they are simply looking at old Fuligo septica and not at a mycoparasite of the slime mold.

Nectriopsis violacea

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Ophiocordyceps dipterigena (Berkeley and Broome) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora Anamorph: Hymenostilbe dipterigena Petch fruitbody a capitate stroma with one or more hemispherical to globoid heads on a cylindrical stalk; head 1–2 mm × 0.5–1 mm, stalk 2.5–8 mm long × 0.2– 0.5 mm thick; head rough on the upper surface from apices of the perithecia, orange-cinnamon to cinnamon-brown; stalk orange-cinnamon to light yellow; one to several stromata are att ached to each host, and the host is attached to the substratum by a mycelium that spreads out, forming a yellowish to brownish crust several mm wide. MICROSCOPIC FEATURES: spores fi liform, multiseptate, breaking into 6–12 × 1–1.5 μm segments, cylindric or fusoid, multiseriate; asci 480– 600 × 4– 6 μm with a 4 μm thick cap, cylindric; perithecia 700–1,000 × 240–540 μm, 15–25 μm thick, vertical, ostioles opening on upper surface of the head, completely embedded; the anamorph forms round, sometimes forked, brown synnemata, 4–12 mm × 0.2– 0.5 mm with the conidiogenous cells forming a dense palisade layer, polyblastic, bearing solitary conidia (4– 9 × 2–4 μm) on short denticles. MACROSCOPIC FEATURES:

on fl ies (Diptera species); southeastern United States, Central America, South America, southeast Asia. COMMENTS: Th is species has been known by many names, including Cordyceps dipterigena Berkeley and Broome and Cordyceps muscicola A. Möller. Ophiocordyceps myrmecophila (Cesati) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora has longer, thinner stromata (0.8–4 cm long × 1 mm wide) and is found on mummified ants in the Pacific Northwest, Michigan, Tennessee, China, and Japan. Ophiocordyceps gracilis (Greville) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora is a widely distributed species complex on larvae of Lepidoptera with a mahogany-red to ochraceous punctate head (2–5 × 2–4 mm) on a deep chrome to straw-yellow stalk (1.5– 9 cm × 0.8–2 mm), and usually with a dense growth of yellow mycelial strands around the host. The spores, asci, and perithecia of Ophiocordyceps gracilis (on Lepidoptera larvae) are very difficult to distinguish from Ophiocordyceps dipterigena (on Diptera species). OCCURRENCE:

Ophiocordyceps dipterigena

Ophiocor dyceps

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Ophiocordyceps melolonthae (Tulasne and C. Tulasne) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora Beetle Cor dyceps, R hinoceros-Beetle Cor dyceps

fruitbody consisting of a head and stalk attached to its host, no head present in early developmental stages; head 1–1.5 cm wide, 2–3 cm high, oval, occasionally forking, whitish to bright yellow, punctate from perithecia embedded in an often irregular pattern; flesh thin, whitish to pale yellow; stalk 5–7 cm long, 3–10 mm thick, nearly equal, or clavate to fusiform, smooth, yellowish. MICROSCOPIC FEATURES: spores 4–10 × 1–1.5 μm, elliptic, smooth, fi liform, and multiseptate at fi rst, breaking into 1-celled spores at maturity, hyaline; asci 180–300 × 6–8 μm, cylindric; paraphyses lacking. OCCURRENCE: solitary or several att ached to buried beetle larvae that look like large white grubs; spring, summer, and fall; widely distributed in eastern North America; occasional. COMMENTS: Cordyceps melolonthae (Tulasne and C. Tulasne) Saccardo and Torrubia melolonthae Tulasne and C. Tulasne are synonyms. According to Joseph Spatafora in a note on the website cordyceps.us, “The use of the name Ophiocordyceps melolonthae for species that produce a bright yellow stroma on the larva of the Hercules beetle is probably inaccurate, but further investigation is required.” We have decided to retain this name until further investigations have been completed. Cordyceps militaris (p. 290) is similar, but it has a cylindric to clavate or fusiform, orange-buff to orange, roughened fruitbody that is att ached to the pupae, or less commonly the larvae, of shallowly buried Lepidoptera. MACROSCOPIC FEATURES:

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Ophiocordyceps ravenelii (Berkeley and M. A. Curtis) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora fruitbody a clavate stroma; stroma 4–10 cm long, upper perithecial portion 3–5 mm thick, stalk 0.8–4 mm thick; apex often sterile, fertile upper portion dark purplish brown to chocolate-brown; interior hyaline to brownish; stalk dark brown; perithecia brownish black, protruding from stroma. MICROSCOPIC FEATURES: ascospores 132–252 × 2– 2.5 μm, fi liform, multiseptate, cells 15–30 μm long, sometimes irregularly breaking into part-spores, multiseriate; asci 175–336 × 7–10 μm with a 2.5– 3 μm thick cap, fusoid or clavate; perithecia 300– 500 × 240–380 μm, hemispherical to subcylindric, walls up to 60 μm thick, brownish black, originating in a layer of pseudoparenchyma and pushing through it into and sometimes through an outer palisade-like layer; conidia narrowly obovoid 5– 7 × 2 μm, 1-celled, hyaline, produced on hyaline phialides scattered in the palisade layer. OCCURRENCE: on beetle larvae in the genus Phyllophaga; widespread in eastern North America, ranging from New England to the southern reaches of the Appalachians; seasonal occurrence not well established; uncommon. COMMENTS: Ophiocordyceps ravenelii was previously known as Cordyceps ravenelii Berkeley and M. A. Curtis. Because of its dark color it can be mistaken for Elaphocordyceps ophioglossoides (p. 298), which grows on Elaphomyces. MACROSCOPIC FEATURES:

Ophiocor dyceps

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Ophiocordyceps sphecocephala (Klotzsch ex Berkeley) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora Anamorph: Hymenostilbe sphecocephala fruitbody consisting of a long stalk att ached to its host with a capitate head developing as the fungus matures, the eyes of the Hymenoptera host are often turned the same shade of yellow as the fruiting body of the fungus; head 2–8 mm long, 1.5–2 mm thick, ovoid, obovoid, or subcylindric to fusoid, fi nely roughened due to partly embedded perithecia, light cream to brownish yellow at fi rst, becoming yellowish brown at maturity, longitudinally irregularly wrinkled when dry, frequently with sparse scatterings of fi ne white hairs; flesh thin, fi rm, brown; stalk 2– 9 cm long × 0.3–1.5 mm thick, dry, glabrous, often twisted, colored like the head on the upper portion, often darker below. MICROSCOPIC FEATURES: spores nearly as long as the ascus fi liform and multiseptate at fi rst, breaking into 1-celled part-spores 8–14 × 1.5–2.5 μm, fusoid, smooth, hyaline; asci 500– 600 × 5– 6 μm, 8-spored; conidia 3–10 × 3–4.5 μm, broadly clavate, strongly apiculate. OCCURRENCE: solitary or in groups on wasps, bees, and hornets that are covered by leaves and pine straw; summer and fall; widely distributed in eastern North America; uncommon. MACROSCOPIC FEATURES:

Specimens must be carefully excavated to recover the host organism. Sphaeria sphecocephala Klotzsch ex Berkeley, Torrubia sphecocephala (Klotzsch ex Berkeley) Tulasne and C. Tulasne, and Cordyceps sphecocephala (Klotzsch ex Berkeley) Berkeley and Curtis are synonyms. Ophiocordyceps myrmecophila (Cesati) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora is closely related to Ophiocordyceps sphecocephala but is smaller and found on mummified ants in the Pacific Northwest, Tennessee, Michigan, Europe, and Asia. The stromata are 0.8–4 cm long, the head 1.5– 4 × 1.5–2 mm, and stalk diameter 1 mm. The head is ochraceous and dries longitudinally rugose. The stalk is light yellow. Ophiocordyceps myrmecophila spores are fi liform, multiseptate, at maturity breaking into part-spores 8–10 × 1.5 μm, and multiseriate. The asci are 480–720 × 4– 6 μm and cylindric with a 6 μm cap. The perithecia are 600–1,020 × 190– 300 μm, 20–25 μm thick, and completely embedded obliquely to the surface in a brownish pseudoparenchyma tissue 50 μm thick.

COMMENTS:

Ophiocordyceps sphecocephala

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Ophiocordyceps superficialis (Peck) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora fruitbody a very slender stroma that tapers to a point and is topped with a short cylindrical to globular aggregation of tiny superficial and free perithecia, 1–5 cm long × 0.3– 2.0 mm thick; light gray or grayish brown, black with age, brownish to clear below the surface; perithecia rough and irregular, chocolate-brown. MICROSCOPIC FEATURES: spores fi liform breaking into 1-celled part-spores 14–32 × 1.5–2 μm, multiseriate; asci cylindric, 170–360 × 6– 9 μm, with a 2 μm thick cap; perithecia 320–560 × 250–420 μm, wall up to 90 μm thick, with an inner layer of thin hyaline cells and an outer layer of mostly 20 μm globular cells. OCCURRENCE: on larvae of Coleoptera; seasonal occurrence not well established; eastern North America from Canada to North Carolina and Tennessee. MACROSCOPIC FEATURES:

Cordyceps superficialis (Peck) Saccardo is a synonym. Ophiocordyceps superficialis is often found in herbaria under the name Cordyceps acicularis Ravenel, which is now Ophiocordyceps acicularis (Ravenel) Petch (not illustrated). Ophiocordyceps acicularis is also eastern and also grows on Coleoptera. It is 6–10 cm long × 0.5–1 mm thick with an ochraceous, warm brown or grayish brown stroma, with superficial, free ochraceous to dark brown perithecia scattered to crowded in the upper part of the stroma, and much shorter partspores (up to 6 μm long). Ophiocordyceps michiganensis (Mains) G. H. Sung, J. M. Sung, HywelJones, and Spatafora is another eastern species found on Coleoptera larvae found buried deep in wood and having superficial to free perithecia scattered on the upper part of the stroma. Its stromata are 1.5–2.5 cm long × 0.2– 0.4 mm thick, ochra-

COMMENTS:

Ophiocordyceps superficialis

Ophiocor dyceps

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ceous to ochraceous orange, commonly occur in groups of 3–11, and have long slender part-spores 12–50 × 1–1.5 μm. Two other species with superficial to free perithecia that are found on Lepidoptera are Ophiocordyceps paludosa Mains (not illustrated) and Ophiocordyceps crinalis (Ellis ex Lloyd) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora (not illustrated). Ophiocordyceps paludosa is 5–13 cm × 0.5–1 mm, with 1–3 stromata per host, is gray or brownish gray, and readily distinguished

by perithecia 800–855 × 375–410 μm (about twice as large as the other species with superficial to free perithecia). Ophiocordyceps crinalis is found on Lepidopteran larvae buried in soil and produces numerous (up to about 30) light grayish brown stromata 5 cm × 0.2– 0.3 mm and unusual fi liform ascospores nearly twice as long as the asci, with obscure septation, not breaking into part-spores. None of the species above are known from west of the Mississippi.

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Ophiocordyceps variabilis (Petch) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora Anamorph: Syngiocladium tetanopsis K. T. Hodge, R. A. Humber, and C. A. Wozniak fruitbody a cylindric stroma, acute above, with a terminus that is usually well developed or lateral cushions on thin stalks that almost or entirely encircle the central portion, occasionally multiple fruiting bodies on a host; stromata 0.5–1.7 cm long × 0.5–1 mm thick; fertile area highly variable in shape, extending anywhere from halfway up the stroma to its terminal, partially or fully encircling it, slightly pulvinate, strongly roughened by the apices of the perithecia, ochraceous almost to the apex; sometimes translucent red above the fertile area; stalk coarsely pruinose to furfuraceous, ochraceous; perithecia 0.4 ×

MACROSCOPIC FEATURES:

0.25 mm, fused in groups, flask-shaped with a conical or subcylindric apex; superficial, adhering at the base, sometimes as few as 10 perithecia on a fruiting body. MICROSCOPIC FEATURES: spores fi liform, breaking into 5–10 × 1.5–2 μm part-spores, multiseriate; asci 200–350 × 8–10 μm with a 2.5–4 μm thick cap, cylindric to subclavate; perithecia completely embedded, 330– 600 × 230–400 μm with poorly differentiated walls 25–35 μm thick, in cushions with a thin cortex of brown pseudoparenchyma. OCCURRENCE: on Diptera larvae in rotten wood, turning the larvae host orange-brown so that they

Ophiocordyceps variabilis

Ophiocor dyceps

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are easily mistaken for beetle larvae; season not well established; throughout North America from Canada to Central America. COMMENTS: Cordyceps viperina Mains and Cordyceps ithacanensis Balazy and Bujakiewicz are synonyms. The species is highly variable. Other species with a sterile apex include Ophiocordyceps stylophora (Berkeley and Broome) G. H. Sung, J. M. Sung, Hywel-Jones, and Spatafora (anamorph Hirsutella stylophora Mains), Ophiocordyceps melolonthae (p. 338), Ophiocordyceps macularis Mains (not illustrated), and Ophiocordyceps unilateralis (Tulasne and C. Tulasne) Petch. Ophiocordyceps stylophora has perithecia in a cylindric layer surrounding the stalk. It is colored ochraceous tawny to dark cinnamon-brown, has short brown hairs on the stalk, and the 102–164 × 2–3 μm multiseptate spores do not break into part-spores. The development of the stromata of Ophiocordy-

ceps stylophora is unusual because they start to develop in late summer but do not mature until the following spring, resulting in many collections of immature material. Ophiocordyceps macularis is 2– 3.5 cm × 1 mm, ochraceous-salmon to vinaceousfawn, grows on Coleopterous larvae, has multiseptate spores 60– 90 × 2–3 μm with 6–14 μm cells that do not form part-spores, and is known from Michigan and New York. Ophiocordyceps unilateralis is 0.3–2 cm × 0.2– 0.5 mm, light to dark brown, tomentose, with one to several dark brown to brownish black pulvinate perithecial cushions on the stalk, grows on ants, and has 100–160 × 2.5–3 μm multiseptate spores with 5–14 μm cells that do not break into part-spores. Of this group, only the much larger Ophiocordyceps melolanthae (p. 338) has spores that, like Ophiocordyceps variabilis, break into part-spores.

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Rosellinia subiculata (Schweinitz) Saccardo fruitbody a single perithecium, at fi rst brown then black, britt le, usually surrounded by a sulphur yellow, orange, or creamcolored subiculum. MICROSCOPIC FEATURES: spores 9–13 × 5– 6 μm, elliptic with a full-length germ slit, brown; asci 90– 150 μm × 7– 9 μm, spores in upper 70–80 μm, small amyloid apical ring; perithecia approximately 0.5 mm diameter, papillate ostioles. MACROSCOPIC FEATURES:

on decayed wood; year-round; worldwide distribution. COMMENTS: Rosellinia quercina R. Hartig, composed of small fused mounds of black perithecia, is found on oaks and is distinguished by the absence of a subiculum. Rosellinia quercina is a significant pathogen of young oak seedlings and young beech trees. OCCURRENCE:

Rosellinia subiculata

Rosellinia

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Thuemenella cubispora (Ellis and Holway) Boedijn fruitbody erumpent through bark, lobed or tuberculate, tending to coalesce into elongated to irregularly shaped compound stromata, margin free or raised with a short, stout stalk; up to 10 mm tall × 4 mm wide × 2.5 mm deep; surface bright lemon-yellow, later darkening or turning brown, eventually uniformly dotted with brown punctuations from an accumulation of spores at the ostioles of the perithecia. MICROSCOPIC FEATURES: spores 6–8(–10) × 4– 6 μm, elliptic to cuboid to angular, wall smooth or MACROSCOPIC FEATURES:

fi nely wrinkled, distinctly gutt ulate, uniseriate, initially yellowish then reddish brown at maturity; asci 70–80 × 7–8 μm, unitunicate, cylindrical on a long stalk; perithecia 220–300 × 100–170 μm including neck. OCCURRENCE: on dead wood and bark; known from Tennessee, Iowa, Quebec, and Jamaica; rare. COMMENTS: Chromocreopsis cubispora (Ellis and Holway) Seaver is a synonym (Corlett 1985).

Thuemenella cubispora

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Torrubiella arachnophila var. leiopus (Johnston) Mains Anamorph: Gibellula leiopus (Vuillemin) Mains subiculum yellow to yellowish white, covering the host; perithecia scattered or crowded, at fi rst embedded and later nearly superficial, light yellowish brown to reddish brown. MICROSCOPIC FEATURES: spores 450– 650 × 1.5 μm, fi liform, multiseptate, tardily breaking into partspores 4–10 μm long; asci 450– 660 × 4– 6 μm, thin walled, with a 4–5 μm cap at the apex; perithecia 550–1,200 × 230–350 μm, narrowly ovoid to conoid. OCCURRENCE: on dead spiders; summer and fall; eastern North America. COMMENTS: Cordyceps arachnophila Johnston and Torrubiella gibellulae Petch are synonyms. There are two varieties in eastern North America characterized by two different conidial forms. Torrubiella arachnophila var. pulchra Mains has a distinctive MACROSCOPIC FEATURES:

conidial stage that was known as Gibellula pulchra (Saccardo) Cavara. It is characterized by violaceous synnemata (3– 9 mm × 100–400 μm) that dry yellowish brown to violaceous brown and are covered with a network of brownish asperulate hyphae (150– 600 × 7–12 μm). The hyphae bear 30– 44 μm diameter heads composed of approximately 12 broadly obovoid prophialides (6–12 × 4– 6 μm), each bearing at their apex 6–8 narrowly clavate phialides (6–10 × 2–3 μm) which produce fusoid to fusoid-elliptic conidia (2.5– 6.4 × 1.5–2.5 μm) singly or in short chains. Torrubiella arachnophila var. leiopus Mains (originally spelled pleiopus) is known as Gibellula leiopus (Vuillemin) Mains in the conidial form. It produces tufted synnemata (1.5– 8 mm × 80–300 μm) that are light grayish brown to violaceous brown and dry.

Gibellula leiopus, the anamorph of Torrubiella arachnophila var. leiopus

Tor ru bi ell a

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Xylaria cubensis (Montagne) Fries Anamorph: Xylocoremium flabelliforme (Schweinitz) J. D. Rogers stromata cylindricclavate to clavate, solitary, usually unbranched, sometimes with a short stalk; (1.5–)3–7(–10) cm tall × (0.2–)0.5–1(–2.5) cm wide; bronze to copper color when young, black in age; stalks often arising from reddish base; slightly rough from minute perithecial ostiolar papillae, network of fi ne cracks (by hand lens); perithecia 0.3– 0.5(– 0.8) mm diameter; anamorph on wood with conical black villose bases and somewhat cerebriform fertile portions 5–30 mm tall × 2–15 mm wide, whitish, yellowish, or pinkish. MICROSCOPIC FEATURES: ascospores 7–13 × 3.5– 6 μm, usually with an obvious germ slit, elliptical-inequilateral, one-celled, smooth, brown to dark brown; asci 125–160 μm long × 6–7 μm wide, spore-bearing part 55–75 μm long, amyloid apical ring short cylindrical to inverted hat-shaped, 1.5–2 μm high × 1.5–2 μm broad; conidia 3–7 × 1.5– 4 μm, obovate to elliptical with flattened basal scar, smooth by light microscopy, irregularly and minutely roughened by electron microscopy, pinkish in mass. OCCURRENCE: anamorph coremia late winter through spring (into summer in northern part of MACROSCOPIC FEATURES:

range) followed by teleomorphic stromata that can persist until the following spring; southeastern North America and South America; common. COMMENTS: Xylaria poitei (Léveillé) Fries (not illustrated) is a large, brown, club-shaped Xylaria with ascospores that measure 14–18 × 5.5–7.5 μm, with a long straight germ slit somewhat less than the full length of the ascospore. It is believed to be a member of the Xylaria cubensis group. These species produce conidia on special anamorphic stromata known as coremia (the anamorph of Xylaria poitei is unknown). A new species from the Pacific Northwest with the proposed name Xylaria atropictor (Callan in preparation) produces a greenish gray, brainy Xylocoremium anamorph in the spring (Brenda Callan, personal communication). The name atropictor was chosen because the wood of the alder and bigleaf maple it colonizes becomes covered with a black layer and atropictor means “dark painter.” It has very dark brown, elliptical-inequilateral, 9–10.5 × 5– 6 μm spores with broad apices and a long but less than sporelength, difficult-to-discern germ slit. The ascus apex is amyloid, almost rectangular to slightly tapered, and 2.2 × 2.2 μm.

Xylaria cubensis (club) with Xylocoremium flabelliforme

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Xylaria hypoxylon (Linnaeus) Greville fruitbody up to 10 cm high, very tough, slender, erect, rounded below, often flattened, branched and antler-like on the upper portion, at fi rst gray to whitish and powdery above and black toward the base, eventually black overall and minutely roughened by embedded perithecia on the upper portion; interior white to whitish. MICROSCOPIC FEATURES: spores 12–15 × 4.5– 6 μm, bean-shaped, smooth, with 1–2 oil drops and a straight germ slit that runs nearly the entire length of the spores, uniseriate, blackish brown; asci 100–150 × 8 μm, cylindric, amyloid apical pore, 8-spored; paraphyses fi liform, aseptate, hyaline. OCCURRENCE: scattered or in dense groups or clusters on decaying branches, logs, and stumps of many species of hardwood; year-round; throughout North America; common. COMMENTS: Xylaria longiana Rehm (not illustrated) is nearly identical, grows on decaying wood and the bark of oak, and has spores that measure 9–11 × 4–5 μm. Xylaria cornu-damae (Schweinitz) MACROSCOPIC FEATURES:

Ca ndlesnu ff Fu ngus

Fries is similar and grows on decaying wood and has much larger spores that measure 16–22(–25) × 5– 6 μm. Xylaria persicaria (Schweinitz: Fries) Berkeley and Curtis (not illustrated) is similar but grows on buried peach seeds. Xylaria oxyacanthae Tulasne and C. Tulasne is 5–10 cm high × 1–3 mm wide at the base, upright, slender, fi lamentous to fusiform, sometimes branched, tough, coated with white asexual spores on the upper half and blackish below, eventually black overall, and roughened by embedded perithecia. Xylaria oxyacanthae, an eastern species on buried fruits of hawthorn, hickory, or pecan, has spores that are (8–)10–13(–14) × 4.5– 6 μm and irregularly elliptic with one side flattened, have a straight germ slit that runs almost the entire length of the spore, and are dark brown to blackish. The asci are 100–140 × 6– 9 μm and cylindrical with an amyloid pore. Paraphyses are fi liform and hyaline. Xylaria multiplex (Kunze) Fries has smaller fruitbodies up to 3 cm high and 1–2 mm wide, with spores that have a long germ slit and measure 9.5–11 × 4.5–5 μm.

Xylaria hypoxylon

Xylaria

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Xylaria liquidambaris Rogers, Ju, and San Martín fruitbody up to 6 cm high × 1–3 mm thick, with short or long concolorous stalks, cylindric to long-conic, usually unbranched, brown to black, very tough; interior white, soft; surface smooth or roughened by embedded perithecia. MICROSCOPIC FEATURES: spores 10–15 × 4– 6.5 μm, elliptic-inequilateral to somewhat crescent shaped, with a spiraling germ slit, smooth, brown; asci 170 × 6–7 μm, cylindric, amyloid apical pore. OCCURRENCE: solitary or in groups on the fallen fruits of American sweetgum, Liquidambar styraciflua; fall and early winter; southeastern United States; fairly common. COMMENTS: Xylaria liquidambar is an orthographic variant. The exclusive association with sweetgum fruits makes Xylaria liquidambaris very easy to identify. Xylaria magnoliae J. D. Rogers, commonly known as “cone fl ickers” or “magnolia cone xylaria,” is upright, slender, fi lamentous to fusiMACROSCOPIC FEATURES:

form, up to 12 cm high, up to 5 mm thick at the base, sometimes branched, tough, and coated with white asexual spores when immature, becoming black at maturity when coated with sexual spores produced in embedded perithecia. Xylaria magnoliae, a southeastern species found on decaying magnolia fruits, has spores that are 11–15(–17) × 3– 5(– 6) μm, elliptic-inequilateral to crescent-shaped, smooth, and pale yellow, with a germ slit that is obscure or lacking. Asci are 90–120 × 6–8 μm and cylindric with an amyloid pore. Xylaria filiformis Fries is distinguished by its growth on the dead stems of herbs and ferns. Its spores (12.5–17 × 5– 6.5 μm) are elliptic, some flattened on one side, with a fulllength germ pore. These species all have whitish conidia produced over the entire teleomorphic stromata and turning black in age after the conidia are released and sexual spore production commences later in the year.

Xylaria liquidambaris

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Xylaria polymorpha (Persoon: Mérat) Greville stromata 2– 9 cm high × 1–3 cm thick, unbranched, sometimes fused together at the base to form a hand-like structure, irregularly clavate or fusiform, hard, pale grayish in the spring from a coating of pale conidia of the anamorph, later black, carbonaceous, fi nely roughened, often wrinkled and minutely cracked, very tough; stalk short, indistinct, cylindric; interior white, fibrous-tough; perithecia embedded just beneath the surface. MICROSCOPIC FEATURES: spores 22–30 × 5– 9 μm, fusiform or irregularly elliptic with one side flattened, germ slit straight, one-third to two-thirds the length of the spore, sometimes with 1 oil drop, smooth, uniseriate, dark brown; asci 130–140 × 8 μm, cylindric, amyloid pore, 8-spored; paraphyses fi liform, aseptate. OCCURRENCE: densely clustered or sometimes solitary on or near decaying hardwood stumps; late spring, summer, and fall, nearly year-round in California; widely distributed in North America; common. MACROSCOPIC FEATURES:

Dea d M a n’s Fingers

Xylaria is divided into four sections based on where and when conidia are formed (Rogers 1985). The Xylaria polymorpha group, which includes Xylaria curta Fries (not illustrated) and Xylaria longipes Nitschke, falls into a section in which the conidia are produced from a palisade of conidiogenous cells over entire young teleomorphic stromata. Xylaria longipes Nitschke has a long stalk, typically grows in clusters on decaying branches and stumps of hardwoods, and has spores that measure 13–16 × 5–7.5 μm with unusual germ slits that spiral around the ascospore. Xylaria curta is widespread and distinguished by its short ascospores that are 9–10 × 4.5– 6 μm with a straight germ slit running the full length of the spore. The stromata of Xylaria corniformis Fries (not illustrated), up to 5 cm high × 8 mm wide, arise from a prominent basal bulb of purplish brown felty tomentum and has short dark brown spores 8–10.5 × 4.5– 6 μm with a thin ventral germ slit running the whole length of the spore (Læssøe 1987).

COMMENTS:

Xylaria polymorpha

Xylaria

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Xylaria tentaculata Berkeley and Broome fruitbody 1.6–3.5 cm high, consisting of a stalk and head with a crown of tentacles; stalk 1.6–2.5 cm long × 1.5–3 mm thick, erect, nearly equal or irregular and twisted, frequently curved near the base, surface granularscurfy, gray to blackish, often with bluish tints; flesh white; head 3–10 mm long, 3– 6 mm wide, cylindric, distinctly scurfy, opening at the apex a prominent ostiole surrounded by up to 8 or more tentacles, gray to pinkish gray; tentacles 1.3–4 cm long, up to 0.75 mm thick, partially erect then spreading, tapered toward the tips, sometimes branched, pale gray to pinkish gray, coated with pale gray to whitish powder at maturity. MICROSCOPIC FEATURES: ascospores 19–22 × 7.5– 9 μm, elliptic-inequilateral to crescent shaped, smooth, with 2 large oil drops, often with a cellular appendage on one end and a mucilaginous appendMACROSCOPIC FEATURES:

Fa iry Spa r k lers

age on the other end, with a germ slit that is slightly less than the spore length, partially biseriate, dark brown; asci 135 × 10 μm, amyloid pore. OCCURRENCE: solitary, scattered, or in groups, sometimes clustered, on organic debris such as decaying wood and humus, or among mosses; summer and fall; eastern North America, most commonly collected in the southeastern United States; rare to locally fairly common. COMMENTS: Xylaria tentaculata is easily overlooked and very fragile. The anamorph is responsible for the crown of tentacles that makes this fungus so distinctive. The conidia are produced in more or less sympodial sequence, seceding individually and passively. The teleomorphic stromata produced later in the year never bear conidia (Rogers 1985).

Xylaria tentaculata

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Chapter 6 Leotiomycetes

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he Leotiom ycetes, a long with Sordariomycetes and Pezizomycetes, are the three large nonlichen-forming ascomycetous classes. Leotiomycetes is currently considered to include Rhytismatales, Helotiales, Cytarriales, Erysiphales, Thelebolales, Myxotrichaceae, and Pseudeurotiaceae. To date, no members of Cytarriales have been observed in North America. Species in Erysiphales, Thelebolales, Myxotrichaceae, and Pseudeurotiaceae do not produce fruitbodies large enough to att ract much attention from mushroomers and thus are mentioned here only in passing. The asci found in Leotiomycetes species are unitunicate, cylindrical, thin- and single-walled, and have a pore, not an operculum. The hyaline spores are of various shapes and are released through the apical pore. The Leotiomycetes include nonlichenized fungi generally with a small apothecium. Most species produce many asci per fruitbody, with eight spores per ascus. They usually have paraphyses between the asci. Considerable morphological diversity exists among these fungi, with ascomata varying from highly gelatinous (Bulgaria) to hairy and fleshy or fragile (Hyaloscyphaceae). The species in Erysiphales, Mycotrichaceae, and Thelebolales form tiny, closed fruitbodies (cleistothecia) with one to many asci. Thelebolus stercoreus typically produces just one ascus per fruitbody with up to 2,000 spores in each ascus. It is quite possible that members of the Thelebolales are not monophyletic and that at least some of the species belong in the Pezizales. Many species in Erysiphales, Rhytismatales, and Thelebolales have distinct ecological and nutritional roles, and knowing the ecology or nutrition of these species helps in naming them. Of these three groups, only two families in the Rhytismatales—Cudoniaceae and Rhytismataceae— produce fruitbodies that are likely to att ract the attention of mushroom hunters and naturalists. Rhytismataceae species are mostly endophytes.

T

They are usually unobserved in plants except when they reproduce, forming dark stromata that are typically immersed in the host tissue; some are called tar spots. We include Rhytisma species and Coccomyces species that produce noticeable spots on leaves. The asci are undifferentiated at the apex and are nonamyloid. The ascospores are long and fi liform, and typically have gelatinous sheaths or appendages. The Cudoniaceae family has recently been placed in the Rhytismatales based on DNA evidence. Th is placement is supported by a hymenium covered by a stromatal layer during early stages of development, ascospores with gelatinous sheaths, and simple asci lacking an amyloid pore. Th is combination of features is not found in the Helotiales order, the previous home of Cudoniaceae. Cudoniaceae genera include Cudonia, Spathularia, and Spathulariopsis (Mains 1956a, 1956b; Maas Geesteranus 1972). The Helotiales order occupies a broad range of ecological niches and includes a great many att ractive macrospecies distributed among eleven families, with additional genera and species that are not yet assigned to a family. The fungi in this order include wood rot fungi, root symbionts, endophytes, terrestrial and aquatic saprotrophs, plant pathogens, mycorrhizae, nematode-trapping fungi, and fungal parasites. Many species are known only from the teleomorphic stage. Some anamorphs without a clear teleomorph are suspected to be members of this order. In the Helotiales we include the polyphyletic Helotiaceae family, one member of the brownblack to black highly gelatinous Bulgariaceae family (Bulgaria inquinans), and the closely related Holwaya mucida. We also include two very different members of the Dermateaceae family, Dermea cerasi (which produces erumpent or superficial small, fleshy apothecia with a gelatinized “epithecium” and thick-walled ascospores; Groves 1946) and Chlorosplenium chlora (a tiny yellow cup on rotten wood with simple nongelatinized asco-

Leotiom ycetes

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spores). Mollisia, which has traditionally been included in the Dermateaceae family, is a large heterogeneous genus that will require further work. It belongs elsewhere in the Leotiomycetes. Mollisia cinerea is closely related to the very differentappearing aquatic species Vibrissea truncorum. Within the Leotiomycetes, Cudoniella and Hymenoscyphus may prove to be the basis for a monophyletic Heliotiaceae sensu stricto (Wang et al. 2006b). We also treat Ascotremella faginea, Bulgariella pulla, Coryne dubia, Discinella boudieri, Gelatinodiscus flavidus, Godronia urceolus, Ionomidotis fulvotingens, Phaeohelotium epiphyllum, Roseodiscus rhodoleucus, Roseodiscus subcarneus, and Tatraea macrospora as part of the Heliotiaceae family. Ascocoryne and Neobulgaria (along with Chloroscypha, which is not included in this treatment) form a clade characterized by a gelatinous fruitbody and endophytic habits. Mitrula is the only genus that we include from a probable third (aquatic) clade (Redhead 1977b; Wang, Binder, and Hibbert 2005). The Hemiphacidiaceae, Rustroemiaceae, and Sclerotiniaceae families are closely related and sufficiently different from other Helotiales to justify a new order for this group (Wang et al. 2006b). The Rustroemiaceae family is represented by Lanzia luteovirescens and Rutstroemia bulgarioides. Rutstroemia species have a gelatinous layer on the underside of the apothecium and Lanzia species have an outer layer of hexagonal cells, but no gelatinous layer. Sclerotiniaceae is represented by: Ciboria rufofusca, which grows on the damp scales of fi r cones; Dumontinia tuberosa, which grows from a tuber among the rhizomes of Anemone species (and differs from similar tuberous Sclerotinia species in having a gelatinous layer on the outer portion of the cup); Sclerotinia sulcata and Sclerotinia veratri, which form sclerotia in culms of herbaceous plants; and Encoelia furfuracea, which grows on alder and hazelnut branches (Whetzel 1946). The very different-looking, but closely related, gen-

era Heyderia (represented by Heyderia abietis) and Chlorencoelia (represented by Chlorencoelia versiformis and Chlorencoelia torta) are the only included members of the Hemiphacidiaceae family. Heyderia species colonize conifer needles as endophytes, and fruit on fallen needles. The Hyaloscyphaceae family is represented by Belonidium sulphureum and Hyaloscypha britannica, plus several Lachnellula and Lachnum species (Huhtinen 1989). Lachnum, Belonidium, and Dasyscyphus are so closely related that many authors treat them all as Lachnum species. All three are saprotrophs on wood, herbaceous stems, leaves, cones, cupules, and so on. All have disc- to cupshaped apothecia, and an exterior that is covered with white, whitish, or brown hairs that microscopically are at least partially rough. Lachnellula species fit the above description except that they are saprotrophic, or parasitic, only on coniferous wood or the resinous excretions, and revive after drying out. Hyaloscypha species, saprotrophs on wood and plant debris, are distinguished by lanceolate paraphyses and smooth hairs that may be heavily coated in granular secretions. Bryoglossum (represented here by Bryoglossum gracile) has been removed from the Geoglossaceae and placed with the Hyaloscyphaceae. However, it is distinctly different in appearance, macroscopically resembling a Mitrula species. The Leotiaceae family is represented by the viscid, globular-headed Leotia lubrica and the distinctive green-headed Leotia viscosa. Microglossum, an earth tongue genus represented here by five different species, appears distinctively different than, but closely related to, Leotia. The two form a distinct clade within the Helotiales. Thuemenidium atropurpureum, while macroscopically resembling a species of Geoglossum, is closely related to Microglossum. Several genera in the Helotiales order remain unassigned to a family. Of the large macrofungi, these include Bisporella, Chlorociboria, and Tapesia.

Overview

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Arachnopeziza aurelia (Persoon) Fuckel fruitbody up to 2 mm wide, rounded and cup-shaped when young, becoming shallowly cup-shaped at maturity, stalkless, attached to the substrate by yellowish mycelium; inner surface smooth, pale yellow to golden yellow; outer surface and margin covered with yellow-orange to orange hairs. MICROSCOPIC FEATURES: spores 12–20 × 3–5 μm, elliptic to fusoid, smooth, 1–3-septate when mature, sometimes with rounded hyaline appendages, hyaline; asci clavate, 70– 90 × 8–10 μm; paraphyses fi liform, septate. OCCURRENCE: scattered, in groups, or in clusters, usually on the underside of decaying fallen logs, acorn cups, branches, or leaves; spring, summer, and fall; eastern and central North America, distribution limits and frequency yet to be determined. MACROSCOPIC FEATURES:

COMMENTS: Peziza aurelia Persoon is a synonym. Because of its growth habit on the underside of fallen logs and branches, Arachnopeziza aurelia is seldom seen or collected. Arachnopeziza delicatula Fuckel (not illustrated), reported from eastern North America, is very similar but has larger clavate spores that measure 35–40 × 3.5–4 μm. Arachnopeziza aurata Fuckel, reported from eastern North America, has a yellowish cup with long whitish hairs on the margin and outer surface, and fi liform to slightly clavate multiseptate spores that measure 50–75 × 1.5–3 μm. Arachnopeziza arctostaphyli Cash (not illustrated), reported from California, has a cream-colored cup and fi liform 7-septate spores that measure 65–80 × 2.5–3 μm.

Arachnopeziza aurelia

Leotiom ycetes

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Ascocoryne cylichnium (Tulasne) Korf fruitbody 3–10 mm high, 3–20 mm wide, nearly round when very young, becoming turban- to cushion-shaped with a depressed center, or shallowly cup-shaped, stalkless or with a rudimentary stalk; inner surface slightly to conspicuously wrinkled, reddish purple to violet-pink; flesh violet-pink, rubbery-gelatinous; outer surface reddish purple to violet-pink; margin lobed or wavy, usually darker than the disc. MICROSCOPIC FEATURES: spores 18–30 × 4– 6 μm, elliptic, multiseptate at maturity, smooth, biseriate, hyaline, sometimes forming one to several tiny secondary spores; asci 205–220 × 10–12 μm, amyloid; paraphyses fi liform, cylindric, unbranched. OCCURRENCE: in groups or clusters on decaying wood, especially beech; late summer and fall, also MACROSCOPIC FEATURES:

winter in California; widely distributed in North America; fairly common. COMMENTS: The widely distributed (North America, Europe, and Asia) Ascocoryne sarcoides, also called purple jelly drops, is very similar but has smaller spores (11–18 × 3.5–5 μm, 1-septate at maturity) and branched paraphyses. The anamorph of Ascocoryne sarcoides, Coryne dubia S. F. Gray (not illustrated; Groves and Wilson 1967), is morphologically similar to its teleomorph and often appears fi rst. The conidia of Coryne dubia are 3–3.5 × 1–2 μm, cylindrical, straight or curved. The fungus produces a terphenylquinone with mild antibiotic properties that helps protect trees against invasion by the heart rot fungi.

Ascocoryne cylichnium

A scocory ne

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Ascocoryne turficola (Boudier) Korf fruitbody 0.5–2 cm wide, 1.2–2.5 cm high, cup-shaped or shallowly cupshaped to turbinate, or irregularly pulvinate, typically with a long, stout stalk that tapers to a point at the base; inner surface smooth, pale to dark olive-green, sometimes with vinaceous tints, becoming brownish in age, more or less translucent; margin wavy to lobed and furrowed; outer surface and stalk whitish to pinkish or pale tan when young, becoming vinaceous to brownish in age, gelatinous, translucent; flesh thick, soft, gelatinous, flexible. MICROSCOPIC FEATURES: spores 12–18 × 4– 6.6 μm, elliptic to fusoid, smooth, unicellular or sometimes septate, uniseriate, hyaline; asci cylindric-clavate, 90–120 × 6.4–10 μm; paraphyses cylindrical, septate, mostly unbranched, tips not enlarged. MACROSCOPIC FEATURES:

solitary, in groups, or in clusters with fused bases, among sphagnum mosses in bogs; summer and fall; northeastern North America; rare to uncommon. COMMENTS: Coryne turficola Boudier, Ombrophila turficola (Boudier) Svrček and Sarcoleotia turficola (Boudier) Dennis are synonyms. Both Species fungorum and MycoBank give Sarcoleotia turficola as the accepted name, but Sarcoleotia is a genus in the Geoglossaceae lacking a gelatinous layer, and Ascocoryne turficola has a gelatinous layer. DNA confi rms that the proper placement is in the Leotiomycetes (Bunyard et al. 2008). Ascocoryne turficola is easily overlooked because its olivaceous colors are very similar to those of some sphagnum mosses on which it is growing. OCCURRENCE:

Ascocoryne turficola

Leotiom ycetes

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Ascotremella faginea (Peck) Seaver fruitbody 1–2.5 cm high, 2–4 cm wide, rounded to irregularly rounded, sometimes forming a continuous mass up to 10 cm or more wide, smooth, shiny or dull, brain-like with rounded lobes and deep furrows or sometimes shallowly cup-shaped, pale pink when young, becoming pinkish brown to reddish purple at maturity; flesh rubbery-gelatinous. MICROSCOPIC FEATURES: spores 6–10 × 3.5–5 μm, elliptic, truncate, with 3–4 fi ne longitudinal striations, with 2 oil drops, typically uniseriate, hyaline; MACROSCOPIC FEATURES:

asci subcylindric, 70– 90 × 6–8 μm, inamyloid; paraphyses slender, slightly enlarged at the tips. OCCURRENCE: solitary, scattered, or in groups on decaying hardwood branches and trunks, especially beech; summer and fall, also winter in California; widely distributed in North America; infrequent. COMMENTS: Th is fungus may be confused with some Basidiomycete jelly fungi, but the presence of asci and ascospores rules them out (Gamundí and Dennis 1969).

Ascotremella faginea

Ascotr emella

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Belonidium sulphureum (Persoon) Raitviir MACROSCOPIC FEATURES:

fruitbody 0.5–2 mm wide, cup-shaped at fi rst, becoming shallowly cupshaped at maturity, stalkless, inner surface smooth, grayish white; margin and outer surface densely coated with stiff sulphur-yellow hairs that fold over the inner surface during dry periods and open during wet weather. MICROSCOPIC FEATURES: spores 26–35 × 1.9–3 μm, cylindric and often slightly curved, with indistinct septa when mature, smooth, biseriate, hyaline; asci cylindric-clavate, 80– 90 × 5–10 μm, inamyloid, 8-spored; paraphyses sparse, narrowly lanceolate to fi liform, projecting beyond the asci. OCCURRENCE: in groups or dense clusters on the stems of nett les and other herbaceous plants, especially members of the carrot family (Apiaceae); spring, summer, and fall; eastern North America, western distribution limits yet to be determined; occasional to common. COMMENTS: Dasyscyphus sulphureus (Persoon) Massee, Erinellina nylanderi (Rehm) Seaver, and Lachnella sulphurea (Persoon) Quélet are synonyms. Hyaloscypha britannica Huhtinen is smaller (0.2– 0.7 mm broad), white on both the upper and lower surfaces, and has an underside densely covered with blunt hairs that are frequently coated with amber to reddish brown resin. Its spores are 7–12.5 × 2–3 μm, narrowly elliptic, with distinct oil droplets, often 1-septate. It is gregarious to confluent on bark and decorticated wood of conifers, has

a nearly continuous fruiting period, and is known from Europe and both eastern and western North America in areas with a primarily maritime climate. There are numerous Hyaloscypha species that are found in North America on wood and various kinds of plant debris. The fruitbodies are minute, cup-shaped and white to grayish, yellowish, or brownish. According to Huhtinen (1989), Hyaloscypha aureliella (Nylander) Huhtinen (not illustrated) and Hyaloscypha candida (Starbäck) Boudier (not illustrated) are ecologically very similar to H. britannica. Hyaloscypha aureliella spores are 6.5–10 × 2–3 μm, rarely septate, and rarely with distinct internal droplets. Hyaloscypha candida has distinctive 200 μm long aseptate hairs with hyaline, rather than amber, encrustations, and spores measuring 5.6–7.6 × 1.8–2.2 μm. All three species vary from collection to collection in hair length, the degree of encrustation, and microscopic detail. Hyaloscypha albohyalina (P. Karsten) Boudier, var. albohyalina (P. Karsten) Boudier, var. spiralis (Velenovský) Huhtinen, and var. tigillaris (P. Karsten) Huhtinen, all with spores 7–14.5 × 2–4 μm, grow on either coniferous or deciduous wood. Their 30–50 × 2–5 μm hairs lack apical lumps of resin and appear minutely to distinctly rough in Melzer’s reagent. See also Arachnopeziza aurelia (p. 356), which has elliptical to fusoid spores 12– 20 × 3–5 μm.

Leotiom ycetes

360

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Belonidium sulphureum

Beloni diu m

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Bisporella citrina (Batsch: Fries) Korf and Carpenter Lemon Drops, Y ellow Fa iry Cups

fruitbody 1–3 mm wide, shallowly cup-shaped, overlapping, stalkless or with a rudimentary stalk; inner surface smooth, bright lemon-yellow to golden yellow; outer surface pale yellow. MICROSCOPIC FEATURES: spores 9–14 × 3–5 μm, elliptic, 0–1-septate, smooth, with several to many oil drops, hyaline; asci 75–135 × 7–10 μm, clavate, pore weakly amyloid or nonamyloid; paraphyses fi liform with internal droplets. OCCURRENCE: in groups or dense clusters on decaying deciduous wood; summer and fall; widely distributed in North America; common. COMMENTS: Helotium citrinum (Hedwig) Fries and Bisporella claroflava (Greville) Lizoñ and Korf are synonyms. Bisporella subpallida (Rehm) Dennis is distinguished by distinctly shorter asepMACROSCOPIC FEATURES:

tate spores (6– 9 × 2.5–3 μm) and smaller weakly amyloid asci (50– 65 × 4.5–5 μm). The spores of Bisporella sulfurina (Quélet) S. E. Carpenter are narrow (9–10 × 2 μm), the asci are intermediate in size (60– 90 × 4–4.5 μm) and nonamyloid, the fertile layer is sulphur-yellow, and it grows on both deciduous wood and the stromata of old pyrenomycetes. Hymenoscyphus and Phaeohelotium species are open initially and are easily confused with Bisporella species except that both of these genera have an amyloid pore at the tip of the ascus. Pithya cupressina (p. 225) is a somewhat similar, bright orange, operculate species. Numerous species in the genera Belonidium, Hyaloscypha, Lachnellula, and Lachnum usually have at least a short stalk and have hairs on the underside.

Bisporella citrina

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Bryoglossum gracile (P. Karsten) Redhead fruitbody an irregular recurved to flat or globose cap on a stalk; cap 0.1– 0.6 cm broad and stalk 1–3 cm tall; cap smooth to convoluted, pale to bright orange, orange-tan, or orange-brown; margin generally free from the stalk; stalk 0.05– 0.1 cm thick, smooth, nearly translucent, flecked with white, paler than upper sporebearing surface of the cap. MICROSCOPIC FEATURES: spores 10–14 × 2–3.5 μm, oblong-fusiform to fusiform, slightly curved, 1-septate when mature, smooth, biseriate, and hyaline; asci 65–80 × 5–8 μm, 8-spored, clavate, amyloid; paraphyses fi liform, 1.5–2 μm thick, septate, straight or sometimes forked. OCCURRENCE: gregarious on moist living mosses (parasitic on Paludella squarrosa) on soil and rocks MACROSCOPIC FEATURES:

in alpine and arctic environments; late summer and fall; widespread. COMMENTS: Mitrula gracilis P. Karsten and Gymnomitrula gracilis (P. Karsten) S. Imai are synonyms. Bryoglossum rehmii (Bresàdola) Ohenoja (not illustrated) is the only other Bryoglossum species. It is associated with mosses in coniferous forests and is distinguished by a pale yellow cap, a margin that is not free from the stalk, and slightly smaller spores (9–13 × 2.5–3 μm). Mitrula borealis Redhead (not illustrated) is a dull orange-yellow northern boreal species associated with living mosses and liverworts. It has 0–1-septate spores (10.5–18 × 2.5–5 μm) with a gelatinous sheath. Cudoniella clavus (p. 375) and Mitrula species fruit on organic material in water or very wet habitats.

Bryoglossum gracile

Bryoglossum

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Bulgaria inquinans (Persoon) Fries fruitbody 2–4 cm wide, 1.6–3.5 cm high, nearly round to top-shaped when young, becoming shallowly cup-shaped in age, stalkless or with a rudimentary stalk; inner surface black, smooth, shiny; outer surface blackish brown to brown and scurfy; flesh rubbery-gelatinous. MICROSCOPIC FEATURES: spores 9–16 × 6– 7 μm, kidney-shaped to elliptic, smooth, of two color forms in each ascus: upper four (mature) brown, lower four (immature) hyaline and typically very poorly developed; asci 95–200 × 9–10 μm, clavate, amyloid, 8-spored; paraphyses slender, with sinuous tips. MACROSCOPIC FEATURES:

Bl ack Jelly Drops

in groups or clusters on decaying hardwoods, especially oak; summer and fall, yearround in California; widely distributed in North America; uncommon to locally common. COMMENTS: Peziza inquinans Persoon and Phaeobulgaria inquinans (Persoon) Nannfeldt are synonyms. Bulgariella pulla P. Karsten, which grows on deciduous branches, especially birch, has a diameter of around 0.3 cm, is dark green to black, and is gelatinous. The spores are dark brown, aseptate, and elliptic, measuring 10–15 × 7.5–8.5 μm. The asci are nonamyloid. OCCURRENCE:

Bulgaria inquinans

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Catinella olivacea (Batsch) Boudier fruitbody saucer-shaped, flat when old, broadly attached to the substrate or embedded, 0.2–1.5 cm broad; upper surface smooth, a gelatinous layer when mature, blackish olive, browner towards the margin; margin vertically furrowed, farinose-furfuraceous, ochraceous to olive-brown; outer surface turns dark purplebrown on application of KOH. MICROSCOPIC FEATURES: spores 7.5–10.5 × 3.5– 4.5 μm, oval-cylindric and concave in the middle of both sides, with 2 oil droplets (resembling the connecting link on a bicycle chain), smooth to minutely verrucose, uniseriate, slightly brownish to darkly pigmented; asci 90–120 × 6–8 μm, 8-spored, inoperculate, nonamyloid; paraphyses cylindrical, septate, slightly clavate at tip, outer coating at tips sloughs off to contribute to the formation of an amorphous, brown, gelatinized pseudoepithecium. MACROSCOPIC FEATURES:

on and in wet rott ing wood and rotting resupinate polypores; late fall; widespread; considered rare but probably usually overlooked since it is often inside rott ing wood or on the bottom of logs and covered by moss (Greif et al. 2007). COMMENTS: Originally named Peziza olivacea Batsch, this species was successively transferred to five other genera in the late 1800s. Cultural studies of the developmental stages clearly show it to be a Dothidiomycete, although its exact relationship to other fungi remains unclear (Greif et al. 2007). Greif et al. (2007) postulate that the formation of a gelatinous pseudoepithecium, as well as associated spore-rich gelatinous droplets, facilitates the dispersal of spores by utilizing passing microarthropods. The typically covered location of the fruitbodies does not facilitate spore dispersal. OCCURRENCE:

Catinella olivacea

Catinella

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Chlorencoelia versiformis (Persoon) J. R. Dixon MACROSCOPIC FEATURES: fruitbody 1–3 cm wide,

shallowly cup-shaped with a short stalk; inner surface olive-green to olive-brown or orange-brown; outer surface dull reddish brown, velvety; stalk 3– 6 mm long, tapered downward, dull reddish brown. MICROSCOPIC FEATURES: spores 9–16 × 2.1–4 μm, cylindrical to allantoid or fusiform, smooth, with 0–4 oil drops, often pseudoseptate, hyaline; asci clavate, 80–140 × 5–7 μm, cylindrical clavate with a long stalk, strongly amyloid; paraphyses slender, sometimes enlarged at the tips, containing greenish matter. OCCURRENCE: scattered or in groups on decaying wood; summer and fall; eastern and central North America, British Columbia; uncommon.

COMMENTS: Chlorociboria versiformis (Persoon) Seaver, Chlorosplenium versiforme (Persoon) P. Karsten, Midotis versiformis (Persoon) Seaver, and Peziza versiformis Persoon are synonyms. The widespread Chlorencoelia torta (Schweinitz) Dixon is often confused with Chlorencoelia versiformis (Persoon) J. R. Dixon, but it is distinguished by hyphae of the tomentum that are stalked and subglobose to clavate rather than fi liform to subclavate, and spores that are irregularly elliptic (5.6–) 9–11(–12) × 2–4 μm versus narrowly elliptic to curved.

Chlorencoelia versiformis

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Chlorociboria aeruginascens ssp. aeruginascens (Nylander) Kanouse ex C. S. Ramamurthi, Korf, and L. R. Batra Blue-Sta in Fu ngus, Gr een-Sta in Fu ngus

fruitbody 3–8 mm wide, cup-shaped to nearly flat and shallowly cupshaped; inner surface smooth, blue-green, sometimes tinted yellow; outer surface blue-green, fi nely roughened; stalk 3– 6 mm long, tapered downward, frequently off-center, blue-green. MICROSCOPIC FEATURES: spores 5– 7(–10) × 1– 2.5 μm, irregularly fusiform or elliptic, bigutt ulate with an oil drop at each end, smooth, biseriate or irregularly crowded, hyaline; asci 45–50 × 3–4 μm, clavate; paraphyses fi liform, slightly enlarged at the tips. OCCURRENCE: in groups or clusters on decaying hardwood; spring, summer, and fall, also winter MACROSCOPIC FEATURES:

and spring in the Pacific Northwest and California; widely distributed in North America; common. COMMENTS: The mycelium of this fungus stains the substrate blue-green. Chlorociboria aeruginascens (Nylander) Kanouse, Chlorosplenium aeruginascens (Nylander) P. Karsten, and Peziza aeruginascens Nylander are synonyms (Ramamurthi, Korf, and Batra 1957). Chlorociboria aeruginosa is nearly identical but has spores that measure 8–15 × 2–4 μm. The stalk is generally central, forming a very symmetrical cup compared to the generally off-center stalk and asymmetrical cup of Chlorociboria aeruginascens.

Chlorociboria aeruginascens ssp. aeruginascens

Ch lorocibor i a

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Chlorosplenium chlora (Schweinitz) Curtis fruitbody 1–3 mm wide, shallowly cup-shaped, stalkless; inner surface yellow when young, becoming yellowish green in age, smooth; outer surface bright yellow. MICROSCOPIC FEATURES: spores 7– 9 × 1.5–2 μm, narrowly elliptic to fusiform, smooth, biseriate, hyaline; asci 40–50 × 6–8 μm, clavate; paraphyses fi liform, tips not enlarged. OCCURRENCE: scattered, in groups, or in clusters on decaying wood; summer and fall; eastern and central North America; infrequent. MACROSCOPIC FEATURES:

COMMENTS: Helotium chlora (Schweinitz) Morgan and Peziza chlora Schweinitz are synonyms. Chlorosplenium olivaceum Seaver (not illustrated), known from Georgia and North Carolina, has a sage-green to pea-green upper surface and simple hyaline spores measuring 9–10 × 3–4 μm.

Chlorosplenium chlora

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Ciboria rufofusca (O. Weberbauer) Saccardo fruitbody initially bladder-like, becoming goblet-shaped to flat, 0.3– 1.0(–1.5) cm broad; upper surface smooth, orange to chestnut-brown; margin can be wavy and split, fi nely dusted with white when young; underside concolorous with upper surface, sometimes frosted with white; stalk slim, 0.2–1.5 × 0.07– 0.15 cm, brownish, darker towards the base.

MACROSCOPIC FEATURES:

MICROSCOPIC FEATURES: spores 4– 7.5 × 2–3.5 μm, elliptic, smooth, sometimes with two small droplets, uniseriate, hyaline; asci 75–80 × 5–5.5 μm, amyloid, 8-spored; paraphyses fi liform, barely thickened toward tips, without septa. OCCURRENCE: single to several per scale on damp fallen fi r cones; spring; Pacific Northwest and northern Europe; uncommon.

Ciboria rufofusca

Ci bor i a

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COMMENTS:

Ciboria amentacea (Balbis) Fuckel (not illustrated), the catkin cup, is widespread in Europe and North America on catkins of alder and willow in spring and early summer. It forms a light brown to yellowish brown shallow cup 0.5–1.2 cm wide, and has a slender stalk 1–5 cm long. Spores are 7.5–13 × 4– 6 μm, elliptic or somewhat inequilateral, uniseriate, and hyaline. Asci reach 110–120 × 7–10 μm, and are 8-spored, with an amyloid pore. Paraphyses are 1–2 μm wide, thread-like, and clavate, up to 5 μm wide at tip. Ciboria caucus (Rebentish) Fuckel (not illustrated) is an almost indistinguishable species on fallen catkins of poplars and hazelnuts. Other Ciboria species grow on fruits, acorns, cones, bulbils, or withered leaves. All Ciboria species are brownish, have a slender stalk, and have unicellular, elliptic, hyaline spores. There are roughly twenty

species in North America still listed in the genus. Other former Ciboria species can now be found in Hymenoscyphus, Rutstroemia, and Monilinia, plus eight other genera that have received one or two transfers from Ciboria. Monilinia species arise from a hollow or compact pseudosclerotium incorporating the fungus and host tissues within the fruits of ericaceous and rosaceous hosts (Honey 1936; Barr 1993). Otherwise they are quite similar to Ciboria species. Rutstroemia species, which grow on stromatized patches of host tissue, are distinguished by a distinct layer of hyphae embedded in a gelatinous matrix on the outside of the cup, and their spores can be multicellular (White 1941). Hymenoscyphus species may or may not have a stalk, their upper surface can be white, yellow, ochraceous, grayish, brown, or rose, and their spores are 0–1-septate. See also Tatraea macrospora (p. 413).

Leotiom ycetes

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Coccomyces dentatus (K. Schumacher) Saccardo apothecium embedded in host leaf tissue, typically about 1 mm diameter in a thin, dark brownish stroma that opens by splitting into four or five rays, exposing the grey or yellowish hymenium. MICROSCOPIC FEATURES: spores 45–55 × 1.5–2 μm, lying parallel in the ascus; asci 8-spored; paraphyses numerous, slender, 1–2 μm. OCCURRENCE: several on dead leaves of hardwood trees and shrubs; mainly in autumn but also in spring; found throughout North America and Europe; common. COMMENTS: There are several species of Coccomyces in North America, but it is difficult to fi nd much information about the genus. They belong to the MACROSCOPIC FEATURES:

Toothed Ta r Spot

Rhytismatales and cause bleaching of leaves via lignin decomposition, similarly to species in the genus Rhytisma. The stromata form black spots on the leaves. Rhytisma species have several lineate wrinkled apothecia in each stroma, while Coccomyces species have one apothecium per stroma. The Coccomyces fungus infects and can kill twigs, and the fruitbodies are seen on dead leaves. In Coccomyces coronatus (Schumacher) De Notaris (not illustrated), the stroma is dark brown to shiny black, circular to broadly elliptic, up to 3 mm across, opening by numerous fi ne rays to expose the yellowish hymenium of the apothecium, and the spores are 30–75 × 2–3.5 μm, lying parallel in the ascus.

Coccomyces dentatus

Coccom yces

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Crocicreas coronatum (Bulliard) S. E. Carpenter fruitbody deeply cupshaped, with or without a short stalk; cup 2–5 mm broad, white to pale yellowish; margin mostly with long pointed teeth; stalk 0.1– 0.3 cm, equal. MICROSCOPIC FEATURES: spores 15–18 × 3.5–4 μm, elliptic, 0–1-septate, with 2 oil droplets, multiseriate, hyaline; asci 80–100 × 6–8 μm, with an amyloid pore, 8–spored; paraphyses 2 μm broad, cylindric to clavate, at most slightly exceeding the asci. OCCURRENCE: on herbaceous stems or woody substrates; autumn; widely distributed in northern Europe but not generally reported from North America, although it has been reported from Newfoundland and Québec. MACROSCOPIC FEATURES:

Synonyms include Calycella alba Patouillard, Cyathicula coronata (Bulliard) De Notarius, and Hymenoscyphus coronatus. Species Fungorum lists 81 records of species and varieties of Crocicreas. Members of this genus are sessile to stipitate, and have an apothecium that is urceolate or cupulate, with an even to dentate margin and colors varying from white to yellowish or pinkish or brown. The spores may have up to 7 septa and the ascus pore may or may not be amyloid. The species are saprotrophs on herbaceous stems and wood. Most are less than 2 mm across.

COMMENTS:

Crocicreas coronatum

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Cudonia circinans (Persoon) Fries fruitbody consisting of a head and stalk; head 5–20 mm wide, irregularly rounded and flattened, smooth to folded or lobed with an inrolled margin, dry; flesh thin, cartilaginous, pale ochre to pale brown; stalk 2–4.5 cm long, up to 6 mm thick, more or less equal, cylindric or compressed, colored like the head above, reddish brown toward the base. MICROSCOPIC FEATURES: spores 30–45 × 1.8– 2.5 μm, narrow, thread-like, sometimes curved, multiseptate, smooth, hyaline; asci 85–130 × 8– 10 μm, clavate with a narrowed apex, inamyloid, 8-spored; paraphyses fi liform, strongly curved near the apex, often branched, with slightly thickened tips. OCCURRENCE: in groups or clusters on the ground or in needle litter, occasionally on well-decayed wood, under conifers; summer, fall, and early winter; widely distributed in North America; occasional to fairly common. MACROSCOPIC FEATURES:

Leotia circinans Persoon is a synonym. Cudonia lutea (Peck) Saccardo, commonly known as yellow cudonia and found in leaf litter under hardwoods in summer and fall in eastern North America, is nearly identical but has a pale yellow to pale brownish orange head, and has larger needle-like, multiseptate, smooth, hyaline spores that measure 40–75 × 1.8–2.5 μm. The asci are 100– 170 × 10–12 μm, clavate with a narrowed apex, and 8-spored. Paraphyses are fi liform, branched, and not thickened but strongly circinate at the tips. Leotia lubrica (p. 394) is similar but has a slippery, gelatinous, ochre head and stalk. The spores are fusiform with numerous gutt ules and 5–7 septae at maturity, and measure 18–28 × 4– 6 μm. Cudonia grisea Mains (not illustrated) fruits in the spring and has spores that are 18–22 × 1.5–2 μm.

COMMENTS:

Cudonia circinans

Cu doni a

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Cudoniella acicularis (Bulliard) J. Schröter fruitbody fleshy, with a cap and stalk; cap 1–3 mm broad, convex, margin involute, white throughout, grayish or brownish with age; stalk 0.5–1.2 cm × 0.3– 0.5 mm, round, white then brownish in age. MICROSCOPIC FEATURES: spores 15–22 × 4–5 μm, becoming 1-septate or up to 3-septate at maturity; fusiform or inequilateral, biseriate, hyaline; asci 80–120 × 10–13 μm, cylindric-clavate with the apex somewhat conical, inamyloid, 8-spored; paraphyses straight, slender, swelling to 3–5 μm at the tip. OCCURRENCE: scattered to gregarious on rott ing trunks and cut surfaces of tree stumps, especially MACROSCOPIC FEATURES:

oaks; fall; widespread, although not reported from western North America; common. COMMENTS: Cudoniella borealis Linder (not illustrated) is a northern Canadian species that looks a lot like Cudoniella acicularis but has smaller spores (8–14 × 2–2.5 μm) and is associated with mosses. Neocudoniella radicella (p. 402), a similar but much smaller translucent white gelatinous species (2–3 mm tall × 1–2 mm broad), is ubiquitous in the boreal forests of Canada and is att ached to spruce rootlets (Kohn, Summerbell, and Malloch 1986). See further comments under Cudoniella clavus (p. 375).

Cudoniella acicularis

Leotiom ycetes

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Cudoniella clavus (Albertini and Schweinitz) Dennis var. clavus fruitbody a disc and stalk; disc 0.4–1.2 cm across, concave, becoming shield-like and then strongly convex (like squatt y gumdrops) with a reflexed margin, smooth, more or less cream-colored, sometimes with grayish, ocher, or violaceous tints; underside of the disc concolorous with the upper side; stalk short to long enough to lift the disc clear of water when the substrate is underwater, 1 mm wide, widened toward apex, slightly tomentose, watery white, becoming dark brown to blackish toward the base. MICROSCOPIC FEATURES: spores 10–17 × 3–5 μm, oblong-fusiform, often narrower at one end, nonseptate, without droplets, uniseriate or irregularly biseriate, hyaline; asci up to 115 × 10 μm, varying greatly in length, cylindric-clavate, inamyloid, 8-spored; paraphyses fi liform, septate, sometimes forked. OCCURRENCE: single to gregarious on wet, rott ing twigs, dead stems and leaves, cones, in ditches and swamps, often in running water; from snowmelt through fall; widespread; common. COMMENTS: The combination of its aquatic habitat and gumdrop shape make Cudoniella clavus a very distinctive species. Cudoniella aquatica (Libert) Saccardo, Ombrophila clavus (Albertini and Schweinitz) Cooke, and Helotium clavus (Albertini and Schweinitz) Gillet are some of the synonyms. Cudoniella clavus var. grandis (Boudier) Dennis grows subsessile on humus in swamps, and its disc is up to 2 cm across. Cudoniella species are distinguished from other members of the Leotiaceae by a convex nongelatinous apothecium that is

not closed when young, hyaline spores, and nonamyloid asci. The apothecia are like those of Hymenoscyphus, but typically have a convex hymenium reminiscent of Leotia. Cudoniella acicularis (p. 374) is the other common Cudoniella in North America. Neocudoniella albiceps (Peck) Korf is a rare species very similar in size, aspect, and sometimes color (the head can be white or rose), but the stalk is gelatinous and translucent to tan to brown or sometimes rose, and the spores are much smaller (5–8 × 2–3 μm).

Cu doni ell a

375

MACROSCOPIC FEATURES:

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Dasyscyphella nivea (R. Hedwig) Raitviir fruitbody cup-shaped with a stalk; 0.5–2 mm broad; upper surface white to cream; exterior white with white hairs often drying orange from encrusting resin. MICROSCOPIC FEATURES: spores 6.5–11 × 1.5–3 μm, elliptic to fusoid, hyaline; asci 30– 60 × 3.5–5.5 μm, cylindric with an apical pore, amyloid, 8-spored; paraphyses 2–2.5 μm broad, exceeding asci by 5– 25 μm, cylindric to narrowly lanceolate; hairs 90– 120 × 2.5–3 μm, slightly capitate, 3–4 μm at apex, multiseptate, fi nely granulose but 1–3 apical cells smooth, amorphous lumps of orange resin often present that dissolve in KOH. OCCURRENCE: gregarious on hardwoods, especially oaks. MACROSCOPIC FEATURES:

COMMENTS: Synonyms include Dasyscyphus niveus (R. Hedwig) Saccardo, Lachnella nivea (R. Hedwig) W. Phillips, and Lachnum niveum (R. Hedwig) P. Karsten. Dasyscyphella montana Raitviir (not illustrated) is very similar, even smaller in stature and with spores about two-thirds the length of Dasyscyphella nivea. Both Dasyscyphella species often have yellowish KOH-soluble crystals along the hairs. Cistella species have exterior hairs that are smooth in the basal portion. Lachnum species and Lachnellula species have hairs that are granular throughout their length. The distinctive feature of Dasyscyphella nivea and Dasyscyphella montana is the absence of granulation on the 1–3 terminal cell(s) of the hairs.

Dasyscyphella nivea

Leotiom ycetes

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Dermea cerasi (Persoon) Fries fruitbody erumpent, gregarious, separate or sometimes cespitose, stalk absent; 1–3 mm wide × 1–1.5 mm high; at fi rst greenish yellow or brownish to yellowish brown, furfuraceous, fi nally black and glabrous, hard, leathery to horny in consistency, becoming more fleshy-leathery when moist; margin at fi rst thick, raised, furfuraceous, fi nally glabrous and disappearing; openings of the pycnidial cavities often appearing as yellowish wrinkles around the margin. MICROSCOPIC FEATURES: ascospores 12–25 × 4–7.5 μm, elliptic-fusiform, straight or slightly MACROSCOPIC FEATURES:

curved, 1–4-celled, irregularly biseriate, hyaline to yellowish brown; asci 75–150 × 10–15 μm, cylindric-clavate, tapering below into a short stalk, amyloid, 8-spored; paraphyses hyaline, fi liform, septate, simple or branched, 1.5–2.0 μm wide, the tips slightly swollen up to 4 μm and glued together, forming a yellowish epithecium. OCCURRENCE: grows on species of Prunus, erupting through the bark; observable year-round; widespread in North America and Europe. COMMENTS: The genus Dermea contains over twenty species of plant parasites.

Dermea cerasi

Der m e a

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Dumontinia tuberosa (Hedwig) L. M. Kohn fruitbody cup-shaped, becoming plane in age, with a stalk, 1–3 cm broad, uniformly brown, grows from buried sclerotia 1.5– 4 cm × 0.5–2.5 cm. MICROSCOPIC FEATURES: spores 12–17 × 6– 9 μm, elliptic, bigutt ulate, hyaline; asci 140–175 × 8– 10 μm; paraphyses cylindrical, enlarged to 3 μm at tip; brick-shaped hyphae of outer excipulum and loose hyphae of inner zone are in a gelatinous matrix. OCCURRENCE: solitary to several among rhizomes of Anemone nemorosa, Anemone ranunculoides, Anemone trifolia, and probably other Anemone species; spring; wherever Anemone species are growing. COMMENTS: Dumontinia tuberosa is the only member of the genus. Whetzelinia tuberosa (Hedwig) Korf and Dumont, Sclerotinia tuberosa (Hedwig) MACROSCOPIC FEATURES:

A nemone Cup

Fuckel, and Hymenoscyphus tuberosus (Bulliard) W. Phillips are synonyms. Sclerotinia species also arise from a tuber borne in soil, but the cells of the outer layer of the cup are not in a gelatinous matrix. Stromatinia rapulum (Bulliard) Boudier (not illustrated) looks very similar macroscopically and microscopically to Dumontinia tuberosa, but arises from a black stroma on buried blackened rhizomes, corms, and bulbs of monocotyledons. The cells of the outer and inner surface of Stromatinia rapulum are not embedded in a gelatinous matrix. Monilinia vaccinii-corymbosi (J. M. Reade) Honey (not illustrated), the blueberry cup (spores 14–18 × 9– 10 μm), is similar but grows from the mummified fallen fruits of highbush blueberry and is found in bogs and wet soil from Massachusett s to North Carolina.

Dumontinia tuberosa

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Encoelia furfuracea (Roth) P. Karsten fruitbody erumpent in clusters, 1–2 cm wide, at fi rst cushion-shaped or irregular; inner surface of young specimens completely covered by the infolded outer surface, eventually splitt ing open to form an irregular cup or shallow cup with a deeply indented and incurved margin; inner surface smooth, yellowish brown to reddish brown; outer surface scurfy and pale yellowish brown. MICROSCOPIC FEATURES: spores 6–12 × 2–4 μm, narrowly elliptic or allantoid with rounded ends, MACROSCOPIC FEATURES:

Scur f y A lder Cup

aseptate, often with two or more small oil drops, biseriate, hyaline; asci 60–80 × 6–7 μm, narrowly clavate with a slender stalk, amyloid pore, 8-spored; paraphyses slender, somewhat enlarged at the tips. OCCURRENCE: emerging singly or in clusters on alder or hazel branches; fall, winter, and early summer; widely distributed in North America; uncommon. COMMENTS: Cenangium furfuraceum (Roth) De Notaris and Peziza furfuracea Roth are synonyms.

Encoelia furfuracea

Encoeli a

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Gelatinodiscus flavidus Kanouse and A. H. Smith fruitbodies top-shaped with a stalk, 0.1– 0.5 cm wide × 0.1– 0.4 cm high; the spore-bearing upper surface initially concave, becoming flat to convex at maturity, bright yellow; stalk entirely gelatinous, bright translucent to opaque yellow with short yellow fuzz at the base, shrinking markedly when drying, turning dark olive. MICROSCOPIC FEATURES: spores 26–34 × 9–11 μm, oblong-elliptic with one broader end, bigutt ulate with large oil drops, smooth, hyaline to yellow, brown when mature; asci 8-spored with a broad, amyloid apical pore; paraphyses 1.5 μm at apex, slender, irregularly branched, curved at the apices, hyaline. MACROSCOPIC FEATURES:

on dead yellow cedar foliage, twigs, and cones; spring, near or under snow banks; known from the coastal regions of the Pacific Northwest; not common. COMMENTS: Th is litt le species may be overlooked by someone thinking it is a very small, young Guepiniopsis, a Basidiomycete. Gelatinodiscus is a monotypic genus in the Heliotiaceae family. Kriegeria alutipes (Phillips) Seaver, found in western North America on the dead foliage of incense cedar, is usually yellowish to pale brown, drying almost black, with fusoid multinucleate spores measuring 18–20 × 5–8 μm. OCCURRENCE:

Gelatinodiscus flavidus

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Godronia urceolus (Albertini and Schweinitz) P. Karsten fruitbody globose, becoming cup-shaped, without a stalk, 0.5–1.5 mm broad; upper surface dark greenish olive to olivaceous black; outer surface black; flesh thick, with a horn-like texture. MICROSCOPIC FEATURES: ascospores 50– 75 × 1.5 μm, fi liform, multispetate, multiseriate, smooth, hyaline; asci 90–130 × 6–8 μm, cylindric, amyloid pore, 8-spored; paraphyses cylindric, at most slightly exceeding the asci; conidia fusoid, 1-septate, 15 × 4 μm. OCCURRENCE: solitary or in small clusters, erumpent through bark or on bare wood, on birch, alder, Ribes, Viburnum, and some ericaceous shrubs; visible year-round; probably throughout North AmerMACROSCOPIC FEATURES:

ica, defi nitely across the northern-tier states and Canada; common but seldom noticed. COMMENTS: Godronia cassandrae Peck (not illustrated), found on Prunus, some ericaceous shrubs, and on Vaccinum, has slightly shorter, distinctly wider spores (50–70 × 2–3 μm). It has been associated with considerable economic damage by causing end rot on cranberries in storage. Godronia cassandrae has a whitish to greenish gray upper surface and a furfuraceous tawny to dark olivebrown exterior. There are well over a dozen species of Godronia in North America. They usually have a dark brown-black exterior with a paler fertile upper layer and are usually seated on a basal stroma. Some are gelatinous or waxy, most are leathery to

Godronia urceolus

Godron i a

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britt le, with the fruitbody sometimes open but often closed when young. All grow on wood or herbaceous substrates. Discocainia (Rhytismataceae) species are distinguished by nonamyloid asci. Their fruitbodies are closed until the spores start to mature. Like many Godronia species they have fi liform spores greater than 30 μm long. The ascospores often bud off from secondary spores. The very similar Gremmeniella species, found on twigs

of conifers, also have nonamyloid asci, fruitbodies that are initially cup-shaped, and elliptic to clavate spores that are greater than 25 μm long, 2– 3 septate, and bud with small secondary spores. Cenangium species growing on conifers and Encoelia species growing on hardwoods have aseptate, cylindrical to fusoid spores; see Encoelia furfuracea (p. 379).

Leotiom ycetes

382

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Heyderia abietis (Fries) Link

Spruce Needle Beacon

fruitbody small, up to 3 cm high, consisting of a head and a slender stalk; head up to 3 mm high × 2 mm wide, cylindric to somewhat clavate, smooth, dull yellow to pale yellow-brown; stalk up to 3 cm long × about 1 mm thick, smooth, colored like the head on the upper portion and dark brown toward the base, cylindric, slightly scurfy; brown basal mycelium spreads out into the conifer needles that are its substrate. MICROSCOPIC FEATURES: spores 10–17 × 1.7–2.5 μm, cylindric-fusiform, with several oil drops, smooth, MACROSCOPIC FEATURES:

hyaline; asci 50–70 × 6–8 μm, amyloid pore, 8-spored; paraphyses fi liform with slightly clavate tips, brownish. OCCURRENCE: in groups on fallen conifer needles, especially spruce; late summer and fall; western and northeastern North America; fairly common. COMMENTS: Although fairly common, this fungus is most likely overlooked because of its small size. Gymnomitrula abietis (Fries) S. Imai and Mitrula abietis Fries are synonyms.

Heyderia abietis

H e y der i a

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Holwaya mucida (Schulzer) Korf and Abawi Anamorph: Crinula caliciiformis (Fries) Fries teleomorph 7–16 mm wide, shallowly cup-shaped with a short to prominent stalk; inner surface smooth, shiny or dull, black, often perforated at the center; outer surface smooth to slightly scurfy, dull, black; stalk up to 1 cm long, tapered downward, black. Anamorph erect, clavate, with a head supported by a stalk; head 2–4 mm wide × 2–4 mm tall, elliptic to rounded, smooth, shiny to dull, viscid when fresh, white to gray; stalk 6–20 mm high × 2–4 mm thick, shiny to dull, scurfy, black. MICROSCOPIC FEATURES: ascospores highly variable in length, 30–75 × 3–4 μm, fi liform to fi liformMACROSCOPIC FEATURES:

cylindric, more or less fasciculate, 14–20-septate at maturity, smooth, often with oil drops, hyaline; asci 120–200 × 10–12 μm, narrowly clavate, nonamyloid, 8-pored; paraphyses fi liform, with globose apices; conidia approximately 3 × 1 μm, elliptic, smooth, hyaline. OCCURRENCE: scattered or in groups arising from cracks in the bark of fallen hardwood trees, especially basswood; late summer, fall, and early winter; eastern North America; occasional. COMMENTS: Holwaya gigantea E. J. Durand, Holwaya leptosperma (Peck) E. J. Durand, and Patellaria leptosperma Peck are synonyms.

Holwaya mucida

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Hymenoscyphus fr uctigenus (Bulliard) Gray fruitbody consisting of a tiny shallow cup and a conspicuous stalk; cup up to 4 mm wide with an entire margin; inner surface smooth, pale yellow; outer surface smooth, colored like the cup or paler; stalk up to 15 mm long, up to 1 mm thick, smooth, shiny when wet, colored like the cup. MICROSCOPIC FEATURES: spores 12–18 × 3–5 μm, irregularly fusiform, smooth, often with two to several oil droplets, sometimes septate, hyaline; asci 80–100 × 7– 9 μm, amyloid pore, 8-spored; paraphMACROSCOPIC FEATURES:

Acor n Cup Fu ngus

yses fi liform, sometimes forked, with slightly enlarged tips. OCCURRENCE: scattered or in groups on acorn cups and beechnut husks; summer and fall; northeastern North America, distribution range and frequency yet to be determined. COMMENTS: Ciboria fr uctigena (Bulliard) Killermann and Helotium fr uctigenum (Bulliard) Fuckel are synonyms. The tiny shallow cup, conspicuous stalk, and growth on acorn cups and beechnut husks make this fungus easy to identify.

Hymenoscyphus fr uctigenus

H y m enosc y phus

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Hymenoscyphus virgultorum (Oeder) W. Phillips MACROSCOPIC FEATURES:

fruitbody at fi rst closed, opening to a shallow cup with a stalk, sometimes becoming convex, 0.05– 0.5 cm wide; upper surface downy, bright yellow to golden yellow, drying ochraceous; underside whitish to pale yellowish; stalk the same length as, or two to three times longer than, the cup diameter, cylindric, sometimes downy toward base, whitish or yellowish. MICROSCOPIC FEATURES: spores 15–22 × 3–5 μm, fusoid or subclavate, partially biseriate, usually containing 2 oil droplets with several smaller ones, occasionally spuriously septate; asci 100–140 × 8– 11 μm, clavate, with an amyloid pore, 8-spored; paraphyses up to 3 μm broad, fi liform, slightly enlarged in upper part. OCCURRENCE: scattered to gregarious on dead hardwood, twigs, partially buried sticks and roots; widespread in North America and Europe; common. COMMENTS: Hymenoscyphus calyculus (Sowerby) W. Phillips is listed in Species fungorum as the accepted name, but it is a misapplied name (Lizoň 1992). According to MycoBank, Hymenoscyphus calyculus is actually a synonym of Cyathicula cyathoidea (Bulliard) Thümen (not illustrated). Cyathicula cyathoidea is 0.3–1.5 mm broad, cup-shaped, and stalked. It has an upper surface that is pigmented brown, grows on herbaceous stems, leaves, or woody substrata, and has spores 9–14 × 2–2.5 μm. The genus Hymenoscyphus is represented by dozens of species in North America. All are sapro-

trophs on wood, herbaceous stems, leaves, mosses, cones, fruits, or cupules. The fruitbodies are cupto cushion-shaped, sessile to stalked, and softfleshed. The fertile upper surface is white, yellow, ochraceous, grayish, brown, rose, or red. The asci have amyloid pores and contain cylindric to fusoid spores that are 0–1-septate, hyaline, and smooth; see also Hymenoscyphus fr uctigenus (p. 385). Roseodiscus rhodoleucus (Fries) Baral = Hymenoscyphus rhodoleucus (Fries) W. Phillips has a rose upper surface and fruits on dead stems of Equisetum. Hymenoscyphus albidus (Roberge ex Desmazières) W. Phillips (not illustrated) fruits from blackened spots on petioles of Fraxinus, is a whitish to yellowish disk 1–2.5 mm broad with a 2 mm long stalk and has spores 13–18 × 3–5 μm. Hymenoscyphus vernus (Boudier) Dennis (not illustrated) fruits on wood, has a yellowish upper surface, a stalk that is dark brown to black at the base, aseptate spores 8–12 × 3–4 μm, and a pore that is not clearly amyloid. Phaeohelotium species are orange to brown and stipitate and grow on wood, with the outer portion of the cup made up of large, thin-walled hyphae. Bisporella species are sessile or short-stipitate; white, yellow, or orange; and have thick-walled hyphae in the outer portion of the cup, versus the thin-walled hyphae found in Hymenoscyphus. Bryoscyphus marchantiae (Berkeley) Spooner is a small (1–1.5 mm) brown species parasitic on a liverwort, with 0–1-septate, elliptic to rhomboid spores measuring 12–15.5 × 6–7 μm.

Leotiom ycetes

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Hymenoscyphus virgultorum

H y m enosc y phus

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Ionomidotis fulvotingens (Berkeley and M. A. Curtis) Cash fruitbody globular and irregularly cup-shaped to umbilicate, subsessile to stalked, 0.1–1 cm wide, brown-green to reddish brown to dark olive, outer surface and recurved margin slightly darker; a distinct stalk sometimes arising from a blackish crust-like stroma; flesh thin, subgelatinous, with soluble pigment that becomes reddish brown in KOH. MICROSCOPIC FEATURES: spores 5–8 × 1–2 μm, cylindrical to elliptic, slightly allantoid, smooth, nonMACROSCOPIC FEATURES:

septate with 2 oil drops, biseriate, hyaline; asci 40– 50 × 3.5–5 μm, inamyloid, 8-spored; paraphyses fi liform, forked, sometimes branched at the apex. OCCURRENCE: erumpent through bark, often solitary to fasciculate, in bark crevices of deciduous trees; fall and winter; eastern North America; rare. COMMENTS: Th is distinctive species is unlikely to be confused with anything else.

Ionomidotis fulvotingens

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Lachnellula agassizii (Berkeley and Curtis) Dennis fruitbody consisting of a shallow cup, up to 5 mm wide, with a rudimentary white stalk; cup rounded at fi rst and soon expanding; upper surface smooth, bright orange-yellow to yellow; outer surface and margin covered with a dense layer of flexible white hairs. MICROSCOPIC FEATURES: spores 6–10 × 3–4 μm, narrowly elliptic, smooth to fi nely verrucose, uniseriate or biseriate, hyaline; asci 60– 95 × 3–4 μm, cylindric to subcylindric, 8-spored; paraphyses fi liform, with clavate tips. OCCURRENCE: in groups or clusters on the bark of decaying conifers; summer and fall; western and northeastern North America; occasional. COMMENTS: Dasyscyphus agassizii (Berkeley and Curtis) Saccardo and Lacnella agassizii (Berkeley and Curtis) Seaver are synonyms. Lachnellula species are saprophytic or parasitic on coniferous wood or resinous excretions. Lachnellula occidenMACROSCOPIC FEATURES:

talis (G. G. Hahn and Ayers) Dharne (not illustrated), found on larch and sometimes spruce and other conifers, is similar, differing by its fruitbody having an upper surface that is salmon-orange and larger spores that measure 10–18 × 4–7 μm. Lachnellula willkommii (Hartig) Dennis (not illustrated), a larch parasite, is also similar, but the upper surface of its fruitbody is yellow to orange and its spores are larger (15–24 × 6–10 μm). Lachnellula suecica (de Bary ex Fuckel) Nannfeldt, found on larch, pine, fi r, and juniper, is also similar, differing because the upper surface of its fruitbody is yelloworange and its globose spores measure 4.5–5 μm. Lachnellula gallica (P. Karsten and Hariot) Dennis, a fi r saprophyte, has broader spores than L. agassizii (8–18 × 5– 6.5 μm), while Lachnellula calyciformis (Batsch) Dharne, a saprophyte of fi r, spruce, and pine, has small elliptic spores (4.5– 6.5 × 2.5–3 μm).

Lachnellula agassizii

Lachnellula

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Lachnellula arida (W. Phillips) Dennis MACROSCOPIC FEATURES:

fruitbody cup- to plateshaped, slightly cushion-like when old, 0.3– 0.7 cm broad, with a short stalk; upper surface smooth, cream to buff to yellow or gold-yellow; margin often wavy, set with bristly short brown hairs, often hyaline-tipped; outer surface thickly set with brown hairs; shrivels in dry conditions, revives after a rain. MICROSCOPIC FEATURES: spores 6.5– 9 × 3.5–5 μm, elliptic, smooth, sometimes with 1 oil droplet, uniseriate, hyaline; asci 55– 60 × 7.5–8 μm, 8-spored, inamyloid; paraphyses cylindric, tips with slight clavate thickenings, sometimes with small outgrowths; hairs up to 190 × 6–8 μm, dark brown, somewhat thick-walled, coarsely encrusted, tips rounded, with distinct septa. OCCURRENCE: clustered on the bark of downed montane conifers; fruits during the spring, present year-round, revives after a rain; western North America, Europe; common in the mountains, dry habitats. COMMENTS: Dasyscyphus arida (W. Phillips) Saccardo (as Dasyscypha) and Lachnella arida (W. Phillips) Seaver are two of the synonyms. Species in both Dasyscyphus and Lachnella are morphologically similar. Lachnellula arida is distinguished by its brown exterior and hairs, relatively broad spores, growth on the wood of fi r species,

and growth in often arid montane regions. Dasyscyphus pini (Brunchorst) G. G. Hahn and Ayers (not illustrated) is also morphologically similar. It has brown exterior hairs, typically an orangeyellow fertile upper surface, and grows on 5-needle pines above 3,000 feet elevation in the western mountains of North America, and in Europe. Its spores are typically 1-septate at maturity but can be 2–3-septate, and measure 15–20 × 5– 6 μm. Lachnellula flavovirens (Bresàdola) Dennis, another conifer species, has brown hairs and smooth elliptic spores measuring 10–13 × 3.5–5.5 μm. Brunnipila calyculiformis (Schumacher) Baral = Lachnum calyculiforme (Schumacher) P. Karsten has brown hairs and is distinguished by its growth on hardwoods and by having fusoid spores measuring 8–14 × 2 μm. Velutarina rufo-olivacea (Albertini and Schweinitz ex Persoon) Korf = Lachnella rufo-olivacea W. Phillips, found on hardwoods from New Jersey to Oregon and in Europe, is distinguished by a dark green upper surface and being 0.1– 0.3 cm across, erumpent, and cup-shaped with an inrolled margin. It has a dense coat of hairs and rusty brown powder giving a mealy appearance, spores that are 12–15 × 6–8 μm, elliptic, and containing 1–2 large oil droplets, and asci that are 120– 160 × 10–15 μm with an amyloid pore.

Leotiom ycetes

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Lachnellula arida

Lachnellula

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Lachnum bicolor (Bulliard) P. Karsten fruitbody closed to cupshaped when young, expanding to almost flat in age, 0.1– 0.2 cm wide, sessile or with a short stalk; upper surface smooth, egg-yellow to orangeyellow, margin thickly covered with long white hairs, strongly inrolled and concealing the upper surface when dried. MICROSCOPIC FEATURES: spores 6–12 × 1.5–2 μm, fusiform to fusiform-clavate, straight or slightly curved, smooth, biseriate, hyaline; asci 40–50 × 4.5– 6 μm, cylindric-clavate, with an amyloid pore, 8-spored; paraphyses 4–5 μm broad, projecting beyond the asci 10–15 μm, lanceolate; hairs 130–200 × 4 μm, septate, with fi ne granular encrustation and sometimes a tuft of crystals at tip. OCCURRENCE: gregarious to cespitose on old canes of Rubus species and branches of hardwoods; summer; widespread in North America and Europe; common. MACROSCOPIC FEATURES:

Dasyscypha bicolor Fuckel and Lachnella bicolor (Bulliard) W. Phillips are synonyms. The genus Lachnum is represented by more than 30 species in North America. Most have a short stalk. All grow on wood, herbaceous stems, leaves, and cones. They are very small, with hairs on the margin of the fertile surface. The fertile upper surface can be white, yellow, orange, reddish, or brown. The hairs are white, whitish, or brown, septate, and encrusted with minute granules, and sometimes have apical crystals. Lachnellula species are very similar but are revivable, while the fruitbodies of Lachnum species are ephemeral. Dasyscyphella nivea (p. 376) can be distinguished from Lachnum and Lachnellula species by examining the terminal 1–3 cells of the hairs. The terminal 1– 3 cells are smooth in Dasyscyphella but not in Lachnum or Lachnellula.

COMMENTS:

Lachnum bicolor

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Lachnum virgineum (Batsch) P. Karsten fruitbody 1.5–4 mm wide, up to 6 mm high, cup-shaped with a distinct stalk; inner surface white, smooth; outer surface white to creamy white, covered by a dense layer of white hairs; stalk 1.5–3 mm long, covered with white hairs. MICROSCOPIC FEATURES: spores 6–10 × 1.5–2.5 μm, spindle-shaped to clavate, smooth, hyaline; asci 45– 60 × 4–5 μm, clavate-cylindric, amyloid pore, 8-spored; paraphyses lanceolate, extending far beyond the asci. OCCURRENCE: scattered, in groups, or in clusters on decaying stems, branches, cones, beechnut MACROSCOPIC FEATURES:

Sta lk ed H a iry Fa iry Cup

burrs, and other debris; spring through early winter; widely distributed in North America; occasional to fairly common. COMMENTS: Dasyscyphus virgineus S. F. Gray is given by MycoBank as the preferred name. Lachnella virginia (Batsch) W. Phillip and Peziza virginea Batsch are synonyms. Lachnum pudibundum (Quélet) J. Schröter is distinguished by its growth on deciduous wood, having shorter hyaline or pale brownish hairs (45– 60 × 4 μm), and aging and drying reddish brown. Its spore size, 7– 9 × 1.5–2.5 μm, is indistinguishable from that of Lachnum virgineum. See also Dasyscyphella nivea (p. 376).

Lachnum virgineum

Lachnum

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Leotia lubrica (Scopoli) Persoon

Ochr e Jelly Club, Jelly Ba bies

MACROSCOPIC FEATURES:

fruitbody consisting of a cap and stalk; cap 6–30 mm wide, irregularly rounded and flattened, smooth to distinctly furrowed or brain-like, with a strongly inrolled margin, moist to viscid, gelatinous, pale dull yellow to orange-yellow, sometimes with green or olive tints; stalk 2–5 cm long, 6–10 mm thick, enlarged downward, slippery, smooth or nearly so, pale dull yellow to orange-yellow. MICROSCOPIC FEATURES: spores 18–23(–28) × 4– 6 μm, cylindric-oblong to fusiform, often curved, multiseptate at maturity, with 3–8 oil drops, biseriate above, uniseriate below, smooth, hyaline; asci 130–160 × 10–12 μm, narrowly clavate, nonamyloid, 8-spored; paraphyses fi liform, branched, tips clavate to pyriform and agglutinated by amorphous matter, hyaline. OCCURRENCE: in groups or clusters on the ground or sometimes on well-decayed wood under conifer and hardwood trees; summer and fall, also winter in the southern and southwestern parts of its range; throughout North America; fairly common. COMMENTS: Helvella lubrica Scopoli is a synonym. Coryne atrovirens (Persoon) Saccardo, distinguished by a pea-green to bluish green cap and a pea-green to bluish green or pale green to whitish stalk, appears to be Leotia lubrica parasitized by

an anamorph that produces single rounded conidia 4– 6 μm in diameter without interfering with the ability of Leotia lubrica to produce normal spores and paraphyses. Coryne atrovirens is easily confused with Microglossum viride (Persoon) Gillet. Microglossum viride is easily distinguished by an ascus with an amyloid pore. It also lacks the rounded, 4– 6 μm diameter conidia, and its 0–4-septate ascospores measuring 12–22 × 5– 6 μm are shorter than Leotia lubrica spores. Leotia viscosa Fries is also similar but it has a dark green cap and a pale greenish yellow to yellowish orange stalk. Cudonia circinans (p. 373) is also similar but its flesh is not gelatinous and the multiseptate spores are threadlike, measuring 30–45 × 1.8–2.5 μm. In eastern North America, Leotia lubrica is sometimes attacked by the parasite Hypomyces leotiicola Rogerson and Samuels, which produces white mycelium on the hymenium and sometimes the stalk of the host and perithecia in the host tissue. Hypomyces leotiicola produces three kinds of spores: hyaline, elliptical to oblong conidia measuring (7–)12.5– 16(–23) × (2.5–)3–4(–5) μm; pale green, globose aleu riospores measuring (17–)19 × 22(–25) μm; and hyaline, naviculate to elliptical ascospores measuring (6–)7.5– 9.5(–11) × 2–3.5(–5) μm (Rogerson and Samuels 1985).

Leotiom ycetes

394

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Leotia lubrica

Leotia

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Leotia viscosa Fries

Gr een-Ca pped Jelly Ba by, Gr een-Hea ded Jelly Club

fruitbody consisting of a cap and stalk; cap 6–20 mm wide, irregularly rounded and flattened, smooth to distinctly furrowed or brain-like, with a distinctly inrolled margin, moist to viscid, gelatinous, medium to dark green, sometimes nearly black; stalk 2–4 cm long, 6–10 mm thick, enlarged downward or nearly equal, slippery, smooth, pale greenish yellow to yellowish orange, often dotted with greenish granules, especially near the apex. MICROSCOPIC FEATURES: spores 18–20 × 5– 6 μm, narrowly elliptic to fusiform with rounded ends, MACROSCOPIC FEATURES:

straight or curved, multiseptate at maturity, smooth, hyaline; asci 120–160 × 8–11 μm, inamyloid, 8-spored; paraphyses fi liform, branched, apices pyriform, septate at maturity. OCCURRENCE: in groups or clusters on soil or among mosses in woods, also on sand dunes; summer and fall, also winter and spring in the southern and southwestern parts of its range; throughout North America; occasional to frequent. COMMENTS: Th is species is common on old sand dunes along the Atlantic and Gulf Coasts during fall and winter.

Leotia viscosa

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Mitrula elegans (Berkeley) Fries

Swa mp Beacon

fruitbody 2–5 cm high, consisting of a head and stalk; head 3–12 mm wide, 6–20 mm high, fusiform to elliptic, pear-shaped, irregularly rounded or lobed, somewhat gelatinous, smooth to slightly wrinkled, shiny, translucent yellow to orange, becoming dull orange in age; stalk 2–4 cm long, 1.5–3 mm thick, enlarged slightly downward, smooth, shiny, whitish to pale translucent gray, sometimes tinted pink. MICROSCOPIC FEATURES: spores 11–17.5 × (1.5–)2– 2.5(–3) μm, narrowly elliptic to cylindric or slightly clavate, septum sometimes present, smooth, lacking a gelatinous sheath, hyaline; asci 115–123 × 5– 7.5 μm, elongate-clavate, amyloid pore, 8-spored; paraphyses fi liform, slightly enlarged at the tips, hyaline. OCCURRENCE: scattered or in groups in shallow water on decaying leaves, needles, twigs, and debris in woodlands and bogs; spring, summer, and fall; northeastern North America, south to Georgia and MACROSCOPIC FEATURES:

Tennessee, west to Oregon and British Columbia; occasional to locally common. COMMENTS: Th is species has been incorrectly identified in some field guides as Mitrula paludosa Fries, a species that has broader spores (12.5– 18.5 × 2.5–3.7 μm) and to date is known only from northeastern Canada and from Europe. Leotia elegans Berkeley and Microglossum elegans (Berkeley) Underwood are synonyms. Mitrula lunulatospora (p. 398) is similar but has a pale watery yellow to pinkish yellow or pale pink head and crescentshaped to allantoid or boat-shaped spores, usually with a gelatinous sheath. Mitrula borealis Redhead (not illustrated) is a northern species that grows in wet areas, usually on living mosses and liverworts. The size and color of its fruitbody and the size and shape of the spores are virtually identical to those of M. elegans, but the spores of M. borealis have a gelatinous sheath.

Mitrula elegans

Mitrula

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Mitrula lunulatospora Redhead fruitbody 2.5–5 cm high, consisting of a head and a stalk; head 3–10 mm wide, 6–20 mm high, oval to pear-shaped or clavate to irregularly rounded, somewhat gelatinous, smooth to slightly wrinkled, shiny, pale watery yellow to pinkish yellow or pale pink; stalk 2–4 cm long, 1–3 mm thick, enlarged slightly downward, smooth, shiny, whitish to pale translucent gray, sometimes tinted pink. MICROSCOPIC FEATURES: spores 11–19 × (2–)2.5– 3.5(–4) μm, lunate to allantoid or cymbiform, sepMACROSCOPIC FEATURES:

tum sometimes present, smooth, usually with a gelatinous sheath, hyaline; asci 91–135 × 6.5–8.5 μm, elongate-clavate, amyloid pore, 8-spored; paraphyses fi liform, tips slightly enlarged, hyaline. OCCURRENCE: scattered or in groups in shallow water on decaying scales, leaves, twigs, and cones in woodlands and bogs; spring and summer; eastern North America; infrequent. COMMENTS: See Mitrula elegans (p. 397).

Mitrula lunulatospora

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Mollisia cinerea (Batsch) P. Karsten fruitbody 0.5–3 mm broad, nearly globose to cup-shaped when young, expanding to nearly flat or convex with age, sometimes becoming distorted and irregularly undulating, without a stalk, sometimes anchored by a mat of hyaline to brownish mycelium; upper surface gray to yellowish gray, sometimes becoming nearly black; underside grayish to brownish, covered with fi ne down; margin often whitish in young stages. MICROSCOPIC FEATURES: spores 6–10 × 2–3 μm, elliptic, sometimes slightly curved, often with several oil droplets, smooth, biseriate, hyaline; asci 45–70 × 5–7 μm, 8-spored, with an amyloid pore; MACROSCOPIC FEATURES:

paraphyses fi liform, cylindric, with blunt ends (3– 5 μm at tips). OCCURRENCE: gregarious to crowded on damp rotting hardwood, especially oak and beech; yearround; widespread in North America and Europe; common. COMMENTS: there are probably at least 40 Mollisia species found on plant debris and decaying wood in North America. All authors we have read agree that Mollisia is a difficult and poorly understood genus and is probably polyphyletic. All members are small, most less than 1 mm in diameter. The fertile upper surface is usually dark-colored but can

Mollisia cinerea

Mollisi a

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be whitish, gray, bluish, or orange-yellow. The cells of the outer wall surface are usually thin-walled, more or less globose, and dark-colored. To identify to species you need to study microscopic features and be able to identify the plant host. Mollisia cinerea is the type for the genus and is by far the most common species. Mollisia ventosa P. Karsten, found on rott ing wood, has a yellowish gray upper surface, a pale margin, a blackish underside, and larger spores (10–20 × 2–3.5 μm) that are 1-septate to possibly 3-septate and cylindrical or slightly allantoid (Dennis 1950). Mollisia discolor (Montagne) Phillips, found erumpent in clusters from cracks in the bark on dead twigs, especially oak, has a light gray interior, a dark brown exterior, and nonseptate spores that have 2 oil drops and measure 10–13 × 2–2.5 μm. Tapesia fusca (Persoon) Fuckel is

found on dead wood and bark like Mollisia cinerea; it can only be distinguished by its slightly larger spores (8–14 × 1.8–3 μm) that lack oil droplets and its subiculum of thick-walled, dark brown hyphae measuring 4–7 μm wide. Belonopsis graminea (P. Karsten) Saccardo and Sydow, which occurs on culms of small grasses in Europe and North America, has 3-septate spores that measure 20–30 × 3– 4 μm, and the 8-spored asci have an amyloid pore (Dennis 1950). Patellariopsis clavispora (Berkeley and Broome) Dennis, found on deciduous wood and bark, is up to 2 mm in diameter, has black apothecia with white flesh, exterior hyphae heavily encrusted with dark brown pigment, and spores that are biseriate, clavate, hyaline, 27–38 × 4.5 μm, and 3–5-septate.

Leotiom ycetes

400

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Neobulgaria pura var. pura (Persoon) Petrak fruitbody 1–2.5 cm high, 1–3 cm wide, forming dense clusters up to 10 cm in diameter; cushion- to top-shaped or shallowly cup-shaped, often distorted, rubbery-gelatinous, smooth, shiny, pale pink to pinkish brown or yellowish brown. MICROSCOPIC FEATURES: spores 7– 9 × 3.5–4.5 μm, elliptic, smooth, with 2 oil drops, uniseriate, hyaline; asci 70– 95 × 7–8 μm, clavate, amyloid pore, MACROSCOPIC FEATURES:

8-spored; paraphyses cylindrical, aseptate, with slightly enlarged tips. OCCURRENCE: in groups or clusters on decaying hardwoods, especially beech; summer and fall; eastern North America; occasional. COMMENTS: Bulgaria pura (Persoon) Fries, Ombrophila pura (Persoon) Quélet, and Peziza pura Persoon are all synonyms.

Neobulgaria pura var. pura

Neobu lga r i a

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Neocudoniella radicella Kohn, Summerbell, and Malloch fruitbody 0.2–2.5 mm wide, consisting of a turbinate to involute creamy white translucent cap on a translucent stalk; cap with a well-defi ned margin, tapering to form a concolorous stalk 2–3 mm long × 0.15– 0.25 mm wide. MICROSCOPIC FEATURES: spores 5– 9 × 1.5–2.2 μm, narrowly elliptic, biseriate, hyaline; asci 33–50 × 3.5–5.2 μm, apex thickened to about 2 μm, ascus pore channel wall moderately amyloid (especially with KOH pretreatment), 8-spored; paraphyses filiform, sparse, not exceeding the asci. MACROSCOPIC FEATURES:

on fi ne rootlets of spruce among Sphagnum mosses; boreal forests across Canada and probably the northern portions of North America, as far south as Oregon in Sphagnum bogs. COMMENTS: Neocudoniella albiceps (Peck) Korf = Leotia albiceps (Peck) Mains is colored tan to brown or rose or sometimes translucent white. It is a much larger, gelatinous species with somewhat fatter spores (5–8 × 2–3 μm) that grows on rott ing wood. It is rare. See also Roseodiscus subcarneus (p. 407) and Cudoniella clavus (p. 375). OCCURRENCE:

Neocudoniella radicella

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Pachycudonia monticola (Mains) S. Imai fruitbody a thick cap on a stalk, 3–10 cm tall; cap 1.0–3.0 cm broad, with shape variably convex, irregularly hemispheric, laterally compressed or subspathulate; surface is wrinkled, fleshy-leathery, pinkish cinnamon or pinkish buff or grayish brown; stalk 0.5– 0.7 cm thick in lower part, somewhat narrower in upper part, hollow when old, bald, brown to grayish purple-brown. MICROSCOPIC FEATURES: spores 18–24(–28) × 2 μm, acicular, 1–celled or rarely with a single septum, hyaline; asci 90–100 × 8–10 μm, clavate, narrower in lower part, 8-spored, spores in upper onehalf to two-thirds; paraphyses fi liform, curved to hooked at the apices, hyaline. MACROSCOPIC FEATURES:

single, gregarious, or cespitose on conifer debris or rott ing wood; usually in spring and summer, at times on the edge of snow banks; a common element of the snow bank flora of the western mountains. COMMENTS: Cudonia monticola Mains is a synonym. Cudonia grisea Mains (not illustrated), a spring species known only from the Pacific Northwest, has slightly smaller spores (18–22 × 1.5–2 μm), a drab to dark gray head, and a smoky to brown stalk. See also Cudonia circinans (p. 373) and Leotia lubrica (p. 394). OCCURRENCE:

Pachycudonia monticola

Pa c h y c u d o n i a

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Phaeohelotium epiphyllum (Persoon) Hengstmengel var. epiphyllum fruitbody up to 0.4 cm wide, cup-shaped to convex, stalk exceedingly short when present, upper spore-bearing surface concave to flat or sometimes convex, bright yellow to orange or occasionally dull red throughout, flesh soft to waxy. MICROSCOPIC FEATURES: spores 15–20 × 3.5–5 μm, fusoid, slightly curved, granular, often with 2–3 oil droplets, sometimes appearing 1-septate, uniseriate in upper part, biseriate in lower part; asci 90–130 × 9–12 μm, clavate, amyloid pore, 8-spored; paraphyses 2–3 μm wide, straight, sometimes branched, scarcely thickened in upper part. MACROSCOPIC FEATURES:

single to scattered on fallen leaves, rarely on needles; throughout North America and Europe. COMMENTS: Hymenoscyphus epiphyllus (Persoon) Rehm apud Kauff man is listed in MycoBank as the preferred name. Phaeohelotium species have large, thin-walled hyphae in the outer part of the cup, while Hymenoscyphus species have normal-sized, thin-walled hyphae and Bisporella species have thick-walled hyphae in the outer part. OCCURRENCE:

Phaeohelotium epiphyllum var. epiphyllum

Leotiom ycetes

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Podophacidium xanthomelum (Persoon) Kavina fruitbody lens-shaped to irregularly disk-shaped, lacking a stalk, 0.1– 0.4 cm wide; upper surface smooth, sulphur-yellow to gold-yellow with a slightly olive tint; margin distinctly brown-lobed and notched, surrounded with a dark brown or almost black laciniate border. MICROSCOPIC FEATURES: spores 10–17 × 5– 6 μm, elliptic, smooth, with 2 oil drops, the ends slightly narrowed, irregularly uniseriate, hyaline; asci 90– 150 × 7–11 μm, cylindroclavate with long stalks, rather broad amyloid pore, 8-spored; paraphyses filiform, sparsely septate, tips forked and bent, about 2.5 μm wide at tips. OCCURRENCE: gregarious or tightly clustered on bare, needle-covered, or moss-covered ground in coniferous forests, more rarely in hardwood forests; late summer to autumn; across the northern part of North America and Europe; fairly common. COMMENTS: There are many old synonyms; the current name has been in use since 1936. MACROSCOPIC FEATURES:

Podoph aci diu m

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Rhytisma americanum Hudler and Banik fruitbody typically circular, 5–12 mm wide, a slightly raised tar-like stroma on maple leaves; surface dry, somewhat shiny or dull, black, marked by a serpentine pattern of alternating ridges and depressions. MICROSCOPIC FEATURES: spores 50–88 × 1.5– 2.5 μm, fi liform, smooth, aseptate, hyaline; asci 60– 100 μm long; paraphyses fi liform, tips slightly to conspicuously curved, or sometimes straight, not enlarged. OCCURRENCE: solitary, scattered, or in groups, sometimes coalescing, on leaves of red, silver, or sugar maple; fall and early winter; northeastern United States and eastern Canada where the host trees are present; very common. MACROSCOPIC FEATURES:

Ta r Spot Of M a ple

Rhytisma acerinum (Persoon) Fries (not illustrated) is similar but it grows on Norway maple. It also forms black spots on leaves, the surface of its fruitbody is roughened by discrete pimple-like projections, and it has larger spores that measure 60–106 × 2–2.5 μm. Rhytisma punctatum (Persoon) Fries is on bigleaf maple leaves in the Pacific Northwest and mountain maple leaves in eastern Canada. Rhytisma punctatum is distinguished by smaller spores that measure 30–40 × 1.5–2 μm. Rhytisma salicinum (Persoon) Fries is found on willow leaves and has spores that measure 60– 90 × 1.5–3 μm. COMMENTS:

Rhytisma americanum

Leotiom ycetes

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Roseodiscus subcarneus (Saccardo) Baral fruitbody 0.4–1.5 mm wide × 1 mm tall, shallowly cup-shaped to saucershaped with a distinct stalk; interior surface whitish with a distinct pale pink tint, similar in color to the exterior surface and the stalk when dried, or sometimes becoming a more opaque creamy yellow; margin smooth, elevated, obtuse, sometimes becoming irregular; stalk 0.6–1.2 mm × 0.2– 0.35 mm, abruptly delimited from cup, nearly smooth under a hand lens, concolorous with the disc, pale ochraceous toward base. MICROSCOPIC FEATURES: spores 4.5–8.5 × 2–3 μm (in KOH about 6–8 × 2–2.3 μm), of variable length, more or less strongly clavate, pyriform or even somewhat cuneate, smooth, biseriate, hyaline; asci 52– 66 × 6.5–7.5(–10) μm (in KOH 42–57 × 5.5– 7.5 μm), 8-spored, apex strongly conic, apical ring amyloid, pore thick in optical section, widest in its MACROSCOPIC FEATURES:

upper part, which extends to the very apex, 0.6– 1 μm high × 1 μm wide, base attenuated or rather broad, arising from croziers; paraphyses cylindric to slightly clavate, 3–7 μm shorter than the asci, terminal cell 16–40 × 3–4 μm, when fresh multigutt ulate in the upper 25–40 μm. OCCURRENCE: scattered to gregarious in bogs and swamps, growing on and apparently killing various species of liverworts and mosses; late summer and early fall; North America and Europe; common but not often observed. COMMENTS: Hymenoscyphus subcarneus (Saccardo) O. Kuntze, Phialea subcarnea Saccardo, and Helotium destructor (Peck) W. L. White are synonyms. Roseodiscus subcarneus is a mountainous to subalpine, and probably boreal, species most easily found by locating the dead patches of liverworts or mosses it causes by growing on them. Other dis-

Roseodiscus subcarneus

Roseodiscus

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tinctive features include its pink-colored, cupshaped fruitbody and relatively long stalk. Roseodiscus rhodoleucus (Fries) Baral (spores 9–12 × 5– 6 μm, elliptic, aseptate) and Roseodiscus equisetinus (Velenovský) Baral (not illustrated) (spores 9.5–13 × 2.5–3.5 μm, 1-septate, pointed at one end at maturity) grow on dead Equisetum and are otherwise very similar to Roseodiscus subcarneus. All three species had previously been placed in Hymenoscyphus. However, the ascus of Hymenoscyphus is truncate, and the pore is a narrow ring that shows up as two very thin amyloid lines that stop short of the ascus apex. The pore on Roseodiscus is like that of Calycina, being thick in optical section and extending to the very tip of the ascus, becoming wider at the apex forming a T shape on the conical ascus apex. Stamnaria is another genus

that may produce pink fruitbodies on dead Equisetum. It has a Calycina-like pore, but the fruitbody is distinguished by a gelatinized outer layer of the cup. The DNA evidence confi rms a close relationship between Roseodiscus, Calycina, and Stamnaria but not Hymenoscyphus (Zhang and Zhuang 2002). Phaeohelotium carneum (Fries) Hengstmengel (= Hymenoscyphus subcarneus [Schumacher] J. Schröter) (not illustrated) is a different species found in the northeast on dead mosses and sometimes rotten wood. Phaeohelotium carneum is up to 1 mm wide, shallowly cup-shaped, stalkless or with a rudimentary stalk, smooth, soft , very thin, somewhat translucent, and whitish to pale purplish pink throughout with smooth hyaline spores 9–13 × 2.5– 4 μm (Hengstmengel 2009).

Leotiom ycetes

408

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Rutstroemia bulgarioides (Rabenhorst) P. Karsten fruitbody 0.25–1 cm wide, concave, expanding to subdiscoid or shallowly cup-shaped, sometimes repand, subsessile, or with a stalk; upper surface brownish black or with an olive tint; underside concolorous with the upper surface; stalk length scarcely exceeding one-half the diameter of the fruitbody, slightly lacunose. MICROSCOPIC FEATURES: spores 6– 9.5 × 3–5 μm, irregularly elliptic, hyaline; asci 85–100 × 5–7 μm, clavate, amyloid, 8-spored; paraphyses fi liform, septate. OCCURRENCE: gregarious on spruce cones; spring; eastern North America, Europe; not uncommon. COMMENTS: Chlorociboria bulgarioides (Rabenhorst) P. Karsten and Ciboria bulgarioides (RabenMACROSCOPIC FEATURES:

horst) Baral are synonyms. Rutstroemia species are somewhat cup-shaped, brownish or sometimes reddish brown or yellowish brown, and stipitate, and arise from stromatized patches of host tissues that often look like an uneven black line around the fungus. The outer layer of the cup has a prominent layer of gelatinized long-celled hyphae. Most species fruit summer and fall on herbaceous debris and wood. Lanzia luteovirescens (Roberge ex Desmazières) Dumont and Korf grows on black stromatized hardwood leaf stalks. The fruitbody is pale greenish yellow overall, consisting of a 0.1– 0.3 cm wide cup on an approximately 1 cm stalk. Spores measure 12–15 × 5–7 μm, and the pores of the asci are amyloid.

Rutstroemia bulgarioides

Ru tstroe m i a

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Sclerotinia sulcata (Roberge ex Desmazières) Whetzel MACROSCOPIC FEATURES:

fruitbody a cup with a relatively long stalk arising from a fusiform, longitudinally furrowed sclerotium; apothecia up to 1 cm broad, externally smooth, fawn-colored; stalk slender, smooth, cylindrical; sclerotium dark brown to black, measuring up to 2.5 × 0.3– 0.4 cm, but varying greatly in size according to the host. MICROSCOPIC FEATURES: spores 9–17 × 5–8 μm, ovoid or inequilateral, uniseriate; asci up to 180 × 11 μm, 8-spored, cylindric-clavate, with an amyloid pore at the apex; paraphyses cylindrical and slightly enlarged at the tip. OCCURRENCE: powdery white microconidial sporodochia fi rst appear on the upper parts of infected culms of Carex species; the sclerotia form inside the culms and are liberated at maturity, producing apothecia the following spring; known from Europe and North America; widespread, common. COMMENTS: Th is species occurs on sedges (Carex species) in swampy areas in many parts of North America. The powdery asexual phase can be observed on young stems of sedges as soon as they start to grow in the spring, with sclerotia forming as early as June then dislodging from the stems at maturity. Usually there is just one sclerotium per stem, but sometimes two or three are observed. The apothecia appear the following spring. Sclerotinia veratri Cash and Davidson forms a 0.3–1 cm grayish brown cupulate apothecium with inrolled margins and a reddish brown underside on a 0.5–

1 cm stalk. The fruitbody emerges from a flat elliptical to irregularly elongate reddish brown to black sclerotia embedded in the stems of California false hellebore in wet montane areas of western North America. Sclerotinia veratri spores are 15–18 × 5– 6.5 μm, oblong-elliptical, with 2 oil drops, irregularly uniseriate, and hyaline. The asci are 140– 150 × 11–13 μm, 8-spored, cylindrical, and attenuated near base, with the wall thickened at the apex. Paraphyses are 2–2.5 μm at the apex, fi liform, septate, simple or branched near base, and pale brown. The anamorph appears to be a species of Botrytis with spores measuring 13–17 × 5–7.5 μm. Species of Sclerotinia always arise from a sclerotium. Another western species, Sclerotinia coloradensis Cash and Davidson (not illustrated), produces sclerotia on stems and seed pods of Pedicularis (louseworts) and produces 2–3.5 mm diameter pale brown cups on a brown 4–7 × 0.5 mm stalk, with spores measuring 10–12 × 4–5 μm. Sclerotinia sclerotiorum (Libert) DeBary (not illustrated) is widespread and infects many herbaceous plants, including Mertensia (alpine bluebells), Aconitum (monkshood), and Delphinium (larkspur). Sclerotinia sclerotiorum produces 0.2–1 cm brownish disks with a 0.4–3 × 0.1– 0.2 cm stalk from a variably shaped tuberoid sclerotium that is 0.4–2.5 × 0.2– 1 cm, with spores measuring 9–13.5 × 4–6 μm. See Dumontinia tuberosa (p. 378) for comments about other similar species.

Leotiom ycetes

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Sclerotinia sulcata

Sclerotinia

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Spathularia flavida Persoon fruitbody up to 6 cm high, consisting of a fan-shaped head and a distinct stalk; head 1–3.5 cm high, 1–4 cm wide, flattened, fan-shaped, wavy to lobed or irregularly folded, dry, glabrous, yellow-ochre to deep yellow; stalk 1–2.5 cm long, 4–20 mm thick, tapered toward the base, dry, smooth, whitish. MICROSCOPIC FEATURES: spores 30– 75(– 95) × 1.5– 3 μm, fusiform, multiseptate, smooth, hyaline; asci 85–125 × 8–13 μm, clavate, inamyloid, 8-spored; paraphyses slender, forked, with curved to spiral tips. OCCURRENCE: scattered or in groups, on the ground or among mosses in conifer woods; spring, summer, and fall in northern California, summer and fall in the rest of its range; widely distributed in North America; occasional. COMMENTS: Spathularia rufa Schmidel is very similar to but less robust than S. flavida, growing among mosses in conifer forests in eastern North America, and it has an ochraceous head that is MACROSCOPIC FEATURES:

folded and a dirty grayish or yellowish brown stalk. Its spores measure 32–50 × 1.5–2 μm. Spathulariopsis velutipes (Cooke and Farlow ex Cooke) Maas Geesteranus, an eastern species known as the velvet-stalked fairy fan or velvety fairy fan, is also similar. Spathulariopsis velutipes has a fan- to spoon-shaped, yellowish to brownish yellow head 1–3 cm wide, is up to 8 cm high, with a distinct stalk that is 2–4 cm long × 6–15 mm thick, narrowed downward or nearly equal, solid, velvety, and reddish brown, with a distinctive dense, orange, basal mycelium. Its spores are 30–45 × 1.5– 2 μm, needle-like, multiseptate, smooth, and hyaline. Asci are 80–105 × 10 μm, clavate, and 8-spored. Paraphyses are fi liform, strongly curved or coiled at the tips, branched, and hyaline. Microglossum rufum (p. 427) has a yellow-orange to orange, spoonto tongue-shaped or cylindric head, and smaller spores.

Spathularia flavida

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Tatraea macrospora (Peck) Baral fruitbody 0.5–1.5 cm wide, up to 1.5 cm high, convex, often somewhat depressed in the center, becoming saucer-shaped when dry; margin not elevated in mature discs, acute, turning upward on drying; upper surface dirty white or grayish, drying brownish; underside minutely furfuraceous, colored like the upper surface; stalk short to long, smooth or minutely furfuraceous, tapered narrower at base. MICROSCOPIC FEATURES: spores 22–34 × 6–8 μm, narrow oblong-fusoid, typically somewhat flattened on one side and tending to be slightly pointed toward the ends, sometimes containing a row of 4–8 large oil drops, a septum fi nally appearing between the drops, uniseriate with ends slightly overlapping to somewhat biseriate, tubules appearing typically at the ends, often on the intercalary cells from which are extruded spermatia until all content is used, leaving only the collapsed spore wall; asci 155–185 × 11–16 μm, cylindric; paraphyses 2–3 μm wide, unbranched, hyaline, not enlarged at tip. MACROSCOPIC FEATURES:

single or more rarely in crowded clusters, on hard parts of decaying stumps, logs, and fallen limbs of hardwood species, also reported on conifer wood; widespread; common. COMMENTS: Rutstroemia macrospora (Peck) Kanouse and Ciboria peckiana (Cooke) Korf are synonyms (Korf 1971). The elongate spores of Tatraea macrospora are distinctive. Macroscopically similar species, all with amyloid asci, are found in genera including Rutstroemia (characterized by a distinct layer of hyphae on the outer part of the cup embedded in a gelatinous matrix), Ciboria (characterized by stalked fruitbodies that grow from a stroma on catkins, fruits, and leaves), Sclerotinia (characterized by tiny sausage-shaped spores), Monilinia (characterized by fruitbodies that arise from a compact pseudosclerotium on shriveled fruits), and Hymenoscyphus (characterized by not growing from a stroma and growing on a wide range of substrates, including wood, stems, fruits, and cupules). OCCURRENCE:

Tatraea macrospora

Tatr a e a

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Vibrissea truncorum (Albertini and Schweinitz) Fries fruitbody consisting of a head and stalk; head 1.5– 6 mm wide, hemispheric to convex with an inrolled margin, smooth, color yellow, orange, or reddish orange; stalk 6–15 mm long × 0.75–1.5 mm thick, nearly equal, slightly roughened, white to pale translucent gray. MICROSCOPIC FEATURES: spores of highly variable length, 120–250 × 1–1.5 μm, needle-like, multiseptate, hyaline; asci 200–325 × 5– 6 μm, inamyloid, 8-spored; paraphyses fi liform, occasionally forked, with somewhat clavate tips. MACROSCOPIC FEATURES:

Water Club

scattered, in groups, or in clusters on partially or completely submerged branches in cold streams; spring and early summer; widely distributed across northern North America, south to West Virginia and higher elevations in the South; infrequent to locally common. COMMENTS: Leotia truncorum Albertini and Schweinitz is a synonym (Schumacher 1976). Vibrissea foliorum Thaxter (not illustrated) is very similar but grows on dead oak leaves, acorn cups, and so on in wet areas and has multiseptate, needle-like spores that measure 85–100 μm long. OCCURRENCE:

Vibrissea truncorum

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Chapter 7 Eurotiomycetes

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urotiom ycetes produce a cleistothecium from which they produce their spores. There are nine orders, two of which, the Eurotiales and the Onygenales, contain mushrooms that att ract the attention of mushroom hunters. The Eurotiales are known as the blue and green molds. There are three families, forty-nine genera, and over nine hundred species. The well-known genera Aspergillus and Penicillium are placed in the Eurotiales. The family Elaphomycetaceae has two genera, one

of which, Elaphomyces, produces hypogeous fungi likely to att ract the attention of mushroom hunters (see the section on hypogeous fungi, chapter 3). The Onygenales are known mostly for the fact that many members are emerging human pathogens that take advantage of people with depressed immune systems or immune system disorders. One genus, Onygena, has five known species that grow on bones or feathers, and we have included two representatives.

Eu rotiom ycetes

416

E

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Onygena corvina Albertini and Schweinitz fruitbody a globose head on a stalk; head 1–2 mm broad, roughfurfuraceous, ocher to light brown; stalk 5–25 mm tall × 1–2 mm thick, cylindric, rather thickened toward the base, usually smooth, whitish. MICROSCOPIC FEATURES: spores 6–8 × 2.5–3 μm, cylindric-elliptic, sometimes curved, smooth, with 1–2 oil droplets, light brown; asci 10–16 × 10–12 μm, nearly spherical, 8-spored, inamyloid, developing within the head of the fruitbody on a hyphal meshwork; paraphyses not observed. MACROSCOPIC FEATURES:

solitary to clustered; on a wide range of keratinous animal remains, including bones, owl pellets, animal hair, remnants of sheep wool, and feathers; summer to fall; widespread; common. COMMENTS: Onygena equina (Willdenow) Persoon is very similar and grows on the shed horns of cows and sheep and on rotten hooves of horses and elk. Onygena equina has broader spores (4–5.5 μm), a larger head (2–4 mm), a shorter stalk (3– 6 mm tall × 2 mm thick), and fruits in spring and summer. OCCURRENCE:

Onygena corvina

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Chapter 8 Geoglossaceae

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he Geoglossace a e fa mily (which includes macroscopic ascomycetes in the genera Geoglossum, Microglossum, Thuemenidium, Trichoglossum, and Sarcoleotia) has long been a part of the Leotiomycetes (Durand 1908, 1921; Mains 1940b, 1954a; Maas Geesteranus 1964; Grund and Harrison 1967). Numerous studies (e.g., Wang et al. 2006b; Eriksson 2005) have led to the exclusion of the Geoglossaceae from the Leotiomycetes, though a new placement remains unclear. The Geoglossaceae may constitute a new class of fungi. In addition to DNA-based evidence, many of the species have brown to black spores rather than the hyaline spores found in members of the Leotiomycetes. Geoglossum species can have either brown or initially hyaline spores. The brown-spored species can be viscid (in which case the paraphyses cover the entire stalk in a dense layer; see Geoglossum difforme and Geoglossum glutinosum) or nonviscid (in which case the paraphyses are confi ned to the hymenium). There are many nonviscid brown-spored Geoglossum species, separated fi rst by spores that soon become septate and colored (e.g., Geoglossum umbratile, Geoglossum glabrum, Geoglossum cookeanum, and Geoglossum simile) versus species with spores that are only colored and septate when fully mature; see Geoglossum fallax. If the spores are hyaline and the septation develops early in spore development, see Geoglossum intermedium. All Geoglossum species are 2– 6 cm tall with a flattened clavate head, have black or dark brown coloration, and have a smooth, pubescent, rugulose, or squamulose stalk that is not very distinctly delimited from the hymenium. Their asci are 8-spored (rarely with 4– 6 spores) and have an amyloid apical pore. Their spores are clavate to subfusoid and their paraphyses are straight or curved or coiled, hyaline or colored, and discrete or agglutinated in a brown gelatinous matter.

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Trichoglossum species are quickly distinguished from the very similar Geoglossum species by the presence of setae that make the fruitbody look hairy, especially when using a hand lens. Trichoglossum species are brownish black and the hymenium is not distinctly delimited from the stalk. The asci are large and 4- or 8-spored, with an amyloid apical pore. The spores are brown, with 3–15 septa, and the paraphyses are brownish and straight or somewhat curved (Sinden and Fitzpatrick 1930). Microglossum species are the general size and shape of either a Geoglossum or a Trichoglossum species, and the hymenium is not distinctly delimited from the stalk. Microglossum species are green, brown, reddish brown, or yellowish orange. They have an ascus pore that is amyloid and spores that are biseriate, hyaline, cylindric-oblong, and septate. Their paraphyses are hyaline and straight or slightly curved. Sarcoleotia is a small genus with only Sarcoleotia globosa (Sommerfelt: Fries) Korf known from North America. We have never found a North American image of Sarcoleotia globosa, a species with a somewhat globose brown to black head (1.5– 3.5 mm tall × 1.5–8 mm wide) that is distinctly delimited from the 0.3–2.5 cm long × 0.05– 0.2 cm thick dark grayish brown stalk. The hyaline spores are 18–36 × 3–5 μm, clavate, slightly curved or straight, smooth, with 0–1(–3) septa. Asci have a distinctly amyloid pore, and paraphyses are straight to hooked. Thuemenidium species are distinguished by hyaline spores and are represented by two species, Thuemenidium atropurpureum and Thuemenidium arenarium, in northern North America. Unfortunately, Thuemenidium is polyphyletic, with Thuemenidium atropurpureum a member of the Leotiaceae and Thuemenidium arenarium a member of the Geoglossaceae (Ohenoja et al. 2010).

Overview

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Geoglossum difforme Fries

Com mon Ea rth Tongue

MACROSCOPIC FEATURES:

fruitbody 3–7 cm high, black, dull when young then smooth and shiny at maturity, viscid, becoming gelatinous when wet, with a head and stalk; head 3–10 mm wide × 2– 3.5 cm high, lance- to spoon-shaped or tongueshaped to cylindric, compressed at the center or irregularly folded, not appearing minutely spiny when examined with a hand lens and not sharply differentiated from the stalk; flesh tough, brown; stalk 2.5–4.5 cm long, 1.5– 6 mm thick, nearly equal or enlarging in either direction, slightly roughened or smooth. MICROSCOPIC FEATURES: spores 90–120 × 6– 7 μm, needle-like, mostly 12–15-septate, smooth, arranged in a parallel fascicle in the ascus, brownish; asci 240–275 × 18–25 μm, clavate, with an amyloid pore, 8-spored; paraphyses slender, coiled and twisted, septate, with slightly enlarged tips, brownish, covering the entire stalk; setae absent. OCCURRENCE: scattered or in groups on soil, decaying wood, and conifer needles or among mosses; summer and fall, eastern North America; fairly common. COMMENTS: The nearly identical viscid black earth tongue, Geoglossum glutinosum Persoon, is widely

distributed in North America and is another viscid Geoglossum species with paraphyses that cover the stalk as well as the fertile area. It is distinguished by shorter (55–)60– 90(–100) × 4– 6 μm, 0– 7-septate spores. Many collections of Geoglossum glutinosum are mostly 3-septate while others are mostly 7-septate when mature, while Geoglossum difforme spores are mostly 12–15-septate. The paraphyses terminate in broad ovoid cells. Geoglossum peckianum Cooke is considered by Mains (1954a) and Seaver (1978a) to be a synonym of Geoglossum difforme, although both Index fungorum and MycoBank list it as a distinct species. Geoglossum affine (Durand) Lloyd (not illustrated), a rare eastern species, is also gelatinous and has paraphyses that continue down the stalk, but the spores are much shorter (40–)45– 62 × 5– 6 μm and are mostly 7-septate; the spores are also more clavate than those of Geoglossum glutinosum and the paraphyses are more septate and irregular. Several species of Trichoglossum are also similar but they have setae and appear minutely spiny when examined with a hand lens.

Geoglossacea e

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Geoglossum difforme

Geoglossum

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Geoglossum simile Peck

Com mon Bl ack Ea rth Tongue

fruitbody 2–7 cm long, compressed-clavate with a stalk; fertile region 2– 12 mm wide, 1/6 to 1/2 of the total fruitbody length, brownish black or black; stalk 1–5 mm thick, round, usually squamulose, sometimes minutely pubescent or smooth, brownish black or black.

MACROSCOPIC FEATURES:

MICROSCOPIC FEATURES: spores (60–)75– 90(–105) × 6– 9 μm, clavate, straight, or frequently somewhat curved, mostly 7-septate or rarely fewer, multiseriate, dark brown; asci 170–200 × 20–22 μm, clavate, apical pore amyloid, 8-spored; paraphyses exceeding the asci, straight or somewhat curved

Geoglossum simile

Geoglossacea e

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above, closely and abundantly septate above with elliptic or obovoid 2-celled segments, constricted at septa, upper cells at most slightly enlarged. OCCURRENCE: scattered to cespitose in swamps, bogs, and well-drained soils on humus, on rotten wood, and in mosses; summer; known from eastern North America and Europe; by far the most common Geoglossum species. COMMENTS: Geoglossum glabrum var. simile (Peck) S. Imai is a synonym. A question among mycologists has been how broadly to interpret Geoglossum glabrum Persoon, another species with 7-septate brown spores. Geoglossum glabrum has paraphyses that are somewhat longer than the asci and much constricted at the septa. The paraphyses are terminated by one to a few dark brown elliptic or globoid cells that can reach a diameter of 15 μm and are agglutinated. The paraphyses of Geoglossum simile separate easily and the terminal cells are not enlarged. Geoglossum pygmaeum W. R. Gerard (not illustrated), known from Tennessee, is a diminutive species 1.5–2 cm tall × 1–2 mm wide. It is brownish black, has stalks that measure about 0.5 mm thick, and its spores measure 120–180 × 6 μm and are 15-septate when mature. The paraphyses are longer than the asci, and apices are slightly curved, slightly constricted at septa, and clavate. Geoglossum pumilum Winter (not illustrated) is another diminutive Geoglossum species that is frequently overlooked. It is distinguished by shorter, sometimes slightly curved spores (90–130 × 5– 6 μm) and by paraphyses that are longer than the asci, apices at most slightly constricted at the septa and moderately brown. Geoglossum fallax E. J. Durand, a common species of NE and NW North America, Scandinavia, Japan, and China, is no-

table for its brown (rather than black) fertile surface color, a fi nely scaly stalk, and generally hyaline spores (45–110 × 5–7 μm) that often leave a white dusting on the fruitbody, only tardily becoming septate and turning smoky. Its spores are clavate-cylindric, straight, or curved, at fi rst continuous and multigutt ulate, then 3-septate and finally with 7–12 septa. Paraphyses are 5– 6 μm thick, cylindric, sparingly to moderately septate, slightly constricted at septa or lacking constrictions, and usually strongly curved or coiled in the upper part. Geoglossum intermedium E. J. Durand (not illustrated), found in northeastern North America, has 0–7-septate spores that measure 40–75 × 6 μm. Geoglossum cohaerens Durand (not illustrated), known from New Jersey, has a scaly stalk and 3– 4-septate spores that measure (30–)38–40(–55) × 5– 6 μm and are hyaline initially, becoming pale brown. It has paraphyses that are longer than the asci, cohering with amorphous matter to form a brown epithecium above the asci. Mains (1954a) notes that Nannfeldt (1942) suggested that Geoglossum cohaerens is close to or identical to Geoglossum littorale (Rostrup) Nannfeldt, a Scandinavian species found on the shores of oligotrophic lakes in association with herbs such as Littorella. Geoglossum alveolatum (E. J. Durand ex Rehm) E. J. Durand, found in Michigan, New York, and possibly Idaho, has 0–15-septate, usually hyaline or rarely light brown spores that measure (50–)65– 85(– 95) × 4–5 μm. It appears to be closely related to Geoglossum intermedium because the species has tufted setose hairs on the stalk and abundant dark amorphous matter that agglutinates the tips of the paraphyses.

Geoglossum

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Geoglossum umbratile Saccardo fruitbody 0.5–7 cm long, somewhat compressed-clavate with a stalk; fertile area 1–5 mm wide, 1/3 to 1/2 the total length of the fruitbody, dark brown to black; stalk 0.5–2 mm thick, round, sometimes somewhat viscid, smooth to pubescent and sometimes squamulose, brown to black. MICROSCOPIC FEATURES: spores 30– 90 × 4.5– 6.5 μm, clavate, straight or somewhat curved, mostly 7-septate, multiseriate, dark brown; asci 125–180 × 16–20 μm, mostly 8-spored, pore amyloid; paraphyses longer than the asci, straight or strongly curved near the tips, cylindric, often slightly enlarged at apex, terminal cell cylindric to clavate, with litt le or no constriction at the septa. OCCURRENCE: scattered or in small groups, on well-drained or boggy soil, among sphagnum and other mosses or on rotten conifer logs; summer; found in both eastern and western North America; second-most-common Geoglossum species. MACROSCOPIC FEATURES:

Geoglossum nigritum (Fries) Cooke is considered to be a misapplied name for this species in North America and refers to a Clavaria. Geoglossum umbratile is closely related to both Geoglossum simile (p. 423) and Geoglossum glabrum Persoon, which differ in having paraphyses that are closely septate in the upper portions. The spore lengths in Geoglossum umbratile are highly variable, even from the same locations but in different years, and the size of the fruitbodies is also highly variable. Some collections of Geoglossum umbratile are viscid; see Geoglossum difforme (p. 420) for a discussion of glutinous species. Some fruitbodies are brown rather than black. Geoglossum umbratile paraphyses septa are not constricted. Similar Geoglossum species with constrictions at the septa of the paraphyses are Geoglossum simile and Geoglossum pygmaeum W. R. Gerard (not illustrated).

COMMENTS:

Geoglossum umbratile

Geoglossacea e

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Microglossum olivaceum (Persoon) Gillet fruitbody up to 6 cm high, consisting of a head and stalk; head cylindric to oval or irregularly clavate, smooth, dull or shiny, sometimes longitudinally furrowed or somewhat flattened, greenish brown to olive-brown or brownish yellow; stalk cylindric or sometimes flattened, with or without a groove, smooth, shiny, pale olive-brown to brownish yellow. MICROSCOPIC FEATURES: spores 12–18 × 4– 6 μm, fusiform to oblong, straight or curved, smooth, 0– 3-septate, biseriate above, uniseriate below, hyaline; asci 75–100 × 9–10 μm, cylindric-clavate, amyloid pore, 8-spored; paraphyses fi liform, often branched, tips not enlarged, hyaline. MACROSCOPIC FEATURES:

Oli v e Ea rth Tongue

solitary, scattered, in groups or clusters on the ground in woods; summer, fall, and early winter, also winter and early spring in California; widely distributed in North America; uncommon to rare. COMMENTS: Geoglossum olivaceum Persoon and Microglossum contortum Peck are synonyms. Microglossum fumosum (Peck) E. J. Durand is very similar but its fruitbody is smoky yellowish to yellowish brown. Microglossum viride Fries, a widely distributed species found on the ground or among mosses and commonly known as green earth tongue, is similar but it has an olive-green to dark green or bluOCCURRENCE:

Microglossum olivaceum

M icroglossu m

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ish green fruitbody, spores that are 12–22 × 5– 6 μm, cylindric-oblong to oblong-elliptic, straight or curved, smooth, 0–4-septate, biseriate, and hyaline. Its asci are 110–150 × 8–10 μm, clavatecylindric with an amyloid pore, and 8-spored, and its paraphyses are fi liform, branched, slightly clavate, and tinged green. Geoglossum viride Persoon and Leptoglossum alabamense Underwood are synonyms.

Microglossum olivaceisquamulosum Grund and Harrison (not illustrated), reported from Nova Scotia, forms greenish brown to olivaceous fruitbodies that sometimes have a yellowish tint, and subcylindric to suballantoid, multigutt ulate spores that measure 18–41 × 4– 6 μm.

Geoglossacea e

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Microglossum rufum (Schweinitz) Underwood

Or a nge Ea rth Tongue

fruitbody up to 6 cm high, consisting of a head and a stalk; head spoonto tongue-shaped or cylindric, compressed at the center or longitudinally furrowed, smooth, dull or shiny, yellow-orange to orange; stalk nearly equal, scaly-granular, yellow to orange-yellow. MICROSCOPIC FEATURES: spores 18–38 × 4– 6 μm, allantoid to fusiform, aseptate when young, becoming 5–10-septate in age, smooth, biseriate, hyaline; asci 100–140 × 10–12 μm, clavate, amyloid pore, 8-spored; paraphyses fi liform, apices slightly clavate, strongly curved. OCCURRENCE: scattered, in groups or clusters on humus, decaying wood, or among mosses; summer and fall; widely distributed in North America; fairly common. COMMENTS: Geoglossum luteum Peck and Leptoglossum luteum Saccardo are synonyms. Microglossum fumosum (Peck) E. J. Durand, found on the ground or on well-decayed wood in Washington state, New England, and eastern Canada, is very similar but its fruitbody is smoky yellowish to yellowish brown. Its spores are (16–)20–50 × 4– 6 μm, cylindric to needle-like, aseptate when young, becoming 7–15-septate at maturity, curved or straight, smooth, biseriate, and hyaline; asci are 100–150 × 10–12 μm, clavate, the apex narrowed with an amyloid pore, 8-spored; and paraphyses are fi liform, septate, and strongly curved above and slightly thickened at the tips. Leptoglossum fumosum Peck is a synonym. Microglossum longisporum Durand (not illustrated) occurs in eastern North America and is also

very similar, but its fruitbody is cinnamon-brown and it forms spores of two sizes in the ascus. The fi rst 2 (very rarely 3 or 4) lie side-by-side and are nearly as long as the ascus and are hyaline, smooth, cylindric or a litt le broader in the middle (60–100 × 4–5 μm), while the remaining (4–)6 spores are near the tip of the ascus and are 12–18 × 3 μm. Also compare with Spathularia flavida (p. 412), which has a yellow-ochre to deep yellow, fan-shaped to folded head, a whitish stalk, and longer spores.

M icroglossu m

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MACROSCOPIC FEATURES:

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Thuemenidium atropurpureum (Batsch ex Fries) O. Kuntze fruitbody up to 6 cm high, clavate with a compressed head on a stalk, black or purplish brown; stalk slightly squamulose. MICROSCOPIC FEATURES: spores 20–35 × 5– 6 μm, 6-septate at maturity, cylindric, tapering towards ends; asci 100–120 × 10–12 μm, biseriate, 8-spored, apical pore amyloid; paraphyses agglutinated at the apices to form a vinaceous-brown epithecium. OCCURRENCE: solitary to several in meadows and grassy forests; fall; eastern North America and the Pacific Northwest. COMMENTS: The distinctive features of Thuemenidium atropurpureum are the paraphyses agglutinated at the apices into a vinaceous-brown epithecium, and 20–35 × 5– 6 μm spores that are 6-septate at maturity, taper towards the ends, and remain hyMACROSCOPIC FEATURES:

aline. Thuemenidium atropurpureum, while considered allied to Microglossum (Ohenoja et al. 2010), belongs with the Leiotiomycetes. Geoglossum atropurpureum (Batsch) Persoon is a synonym. Thuemenidium arenarium (Rostrup) Korf = Geoglossum arenarium (Rostrup) Lloyd is the only other northern species in the genus Thuemenidium. However, the two species are not closely related and DNA evidence points to a proper placement of Thuemenidium arenarium in the Geoglossaceae with Geoglossum arenarium as the accepted name. The apices of the paraphyses are not agglutinated, and the spores are one-celled (25–40 × 4– 6 μm). It can be found in large numbers on sand dunes and sandy shores.

Thuemenidium atropurpureum

Geoglossacea e

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Trichoglossum hirsutum (Persoon) Boudier H a iry Ea rth Tongue, V elv et y Bl ack Ea rth Tongue

fruitbody 4–8 cm high, consisting of a head that tapers into a stalk; head elliptic to oval or fusiform, often compressed or flattened with a lengthwise furrow, hollow, dark sooty brown to black, surface dry and minutely velvety; flesh tough, brown; stalk about 3 mm thick, nearly equal or slightly compressed, often curved, tough, colored like the head, surface densely velvety. MICROSCOPIC FEATURES: spores (80–)100– 150(–160) × 6–7 μm, cylindric to cylindric-clavate and thread-like, 15-septate at maturity, smooth, arranged in a parallel fascicle, brown; asci 210–225 × 20–22 μm, broadly clavate, with an apical amyloid pore, 8-spored; paraphyses cylindric, septate, usually curved or coiled at the slightly enlarged tips, brown; setae up to 225 μm long, sharply pointed, brown. OCCURRENCE: solitary or in groups or clusters on soil, on well-decayed wood, or among mosses, typically in woods but sometimes in open areas; summer and fall, also collected during winter and spring in the Pacific Northwest and California; widely distributed throughout North America; common. COMMENTS: Similar species of Trichoglossum must be identified microscopically. Geoglossum species are very similar but they have smooth fruitbodies, lack setae, and also must be identified microscopically. Trichoglossum variabile (E. J. Durand) Nannfeldt (not illustrated) is distinguished by mostly 9–14-septate spores (80–150 × 6 μm) that taper at both ends. Trichoglossum velutipes (Peck) E. J. Durand is a widespread species with 4-spored asci;

the spores are (90–)110–145(–160) × 6–7 μm and mostly 9-septate with a range of 7–11(–13) septa. Trichoglossum walteri (Berkeley) E. J. Durand is the most common of the species and is distinguished by its mostly 7-septate spores, which are elliptical to cylindric (70–100 × 5.5–7 μm). Trichoglossum farlowii (Cooke) Durand usually has 3-septate spores, but they can be 1–5-septate, rarely 6-septate, and are 30– 90 × 4.5–10 μm.

Tr ichoglossum

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MACROSCOPIC FEATURES:

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Chapter 9 Neolectomycetes

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he cl ass Neolectom ycetes contains just one genus, Neolecta (order Neolectales, family Neolectaceae) with two species known from North America.

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Overview

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Neolecta irregularis (Peck) Korf and J. K. Rogers Ir r egu l a r Ea rth Tongue

fruitbody up to 7.5 cm high, clavate to spathulate or highly irregular, often twisted or contorted, compressed, obtuse, sometimes lobed, stuffed or hollow; upper fertile portion smooth, yellow to orange-yellow; lower sterile portion tapered downward and forming a stalklike base, smooth, pale yellow to white; flesh fairly tough, white to pale yellow. MICROSCOPIC FEATURES: spores 5.5–10 × 3.5–5 μm, elliptic, often slightly reniform, smooth, uniseriate, hyaline; asci 100–135 × 5–7 μm, clavate-cylindric, 8-spored, amyloid; paraphyses absent. OCCURRENCE: scattered or in groups on the ground, among mosses or on needle duff in conifer woods; summer and fall; widely distributed in North America; occasional to common. MACROSCOPIC FEATURES:

reported as edible, but not recommended. COMMENTS: Ascocorynium irregulare (Peck) S. Ito and S. Imai, Geoglossum irregulare Peck, Mitrula irregularis (Peck) Durand, and Spragueola irregularis (Peck) Nannfeldt are synonyms. Neolecta vitellina is similar but forms regularly clavate and generally narrower fruitbodies with spores that measure 5.5– 9 × 3–4 μm and asci 53–75 μm long (Redhead 1977a; Landvik et al. 2003). Globose to subglobose conidiospores 1.75–2 μm in diameter are regularly associated with Neolecta vitellina but not observed with Neolecta irregularis. EDIBILIT Y:

Neolecta irregularis

Neolectom ycetes

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Chapter 10 Orbiliomycetes

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he Or bili ace a e fa mily, formerly considered a part of the Leotiomycetes, now have been placed in their own class, the Orbiliomycetes. The Orbiliomycetes comprise one order, Orbiliales, with one family, Orbiliaceae, composed of 12 genera. Most species do not have dense structural tissue that produces fruitbodies. The macroscopic species likely to att ract the attention of mushroom hunters and naturalists have small diskshaped apothecia that are typically convex, translucent, and brightly colored. The two distinctive macroscopic genera are Hyalorbilia and Orbilia. Many mitosporic (anamorphic) Orbiliomycetes, including Arthrobotrys species, are car-

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nivorous. The most common way these species trap their prey is to produce hyphal outgrowths that curve to form a complex network of adhesive loops. A second trapping mechanism involves the formation of closely spaced adhesive knobs on the hyphae. Some species form three-celled rings on a stalk attached to the hyphae. Some rings are nonconstricting, simply entangling their prey, but the most dramatic rings swell rapidly inward on contact with a nematode, “lassoing” the victim with such force that the nematode is sometimes nearly severed. In all cases, the nematode thrashes briefly but soon is still, whereupon other hyphae grow into the prey and digest it.

Or bi liom ycetes

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Orbilia luteorubella (Nylander) Karsten fruitbody nearly plane, often depressed in the center, stalk absent; 0.2– 1.5 mm diameter; smooth and waxy, semitranslucent, initially pallid, then orange, pale yellowish red, or yellowish brown, drying yellowish. MICROSCOPIC FEATURES: spores 6–12 × 1–1.5 μm, fusoid, fi liform, biseriate, hyaline; asci 30–40 × 4–4.5 μm, cylindric-clavate, 8–spored, nonamyloid; paraphyses fi liform, the apices clavate to subspheroid, 2–2.5 μm at tip. OCCURRENCE: on rotten wood of hardwoods; summer and fall; eastern North America, Europe; common. COMMENTS: Orbilia xanthostigma (Fries) Fries is orange or golden yellow, and the ascospores are uniseriate (versus biseriate), much shorter (3–4 × 1–2 μm), and have 2 oil drops. Orbilia leucostigma (Fries) Fries (not illustrated) differs from Orbilia xanthostigma only in being translucent white, and it is much less common. Orbilia piloboloides Haines and Egger, found in northeastern North America on hardwood bark (especially dead American elm), is yellow to orangish yellow with a whitish external coating most noticeable as a white fringe at the cup margin. Orbilia piloboloides has distinctive shortnaviculate to fusoid spores (8–10 × 2–2.5 μm) and typical Orbilia paraphyses that are fi liform 0.9– 1.2 μm diameter below, swollen at the apex to a distinct spherical, thick-walled knob 1.1–1.8 μm diameter, subtended by a thin-walled pyriform swelling 2.0–3.3 μm diameter. Orbilia delicatula (P. Karsten) P. Karsten has spores (2.5–3 × 1 μm) that are curved in a semicircle. The golden yellow Orbilia inflatula (P. Karsten) P. Karsten (= Hyalorbilia inflatula [P. Karsten] Baral and G. Marson) differs from O. luteorubella in being surrounded on the substrate by conspicuous white anchoring hyphae like a rudimentary but conspicuous subiculum, and the fi liform paraphyses are a bit clavate but not knob-headed as in most species of Orbilia. Orbilia

inflatula is found on woody substrata and old fungi, especially pyrenomycetes in North America and Europe. It has an epithecium and a strongly developed hymenal gel above the hymenium. The inseparable paraphyses are agglutinated in the hymenal gel, the spores are 4.5–7 × 0.5–1 μm, cylindric or rod-shaped, and the nonamyloid asci are 24–30 × 3–4.5 μm. There are probably close to 20 species of these thin, colorful litt le semitranslucent to waxylooking cups in North America. They are saprotrophic on wood, herbaceous stems, and old pyrenomycetes. Some are claimed to be lichenized. The group is understudied and in need of a modern revision (Hansen and Knudsen 2000).

Or bi li a

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MACROSCOPIC FEATURES:

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Orbilia luteorubella

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Chapter 11 Dothideomycetes

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s of 2012, the Dothideomycetes comprised 11 orders, 90 families, 1300 genera, and more than 19,000 known species. Reproductively, most species in this huge and diverse group of Ascomycetes reproduce via flask-like pseudothecia. The asci are two-layered, and the inner layer bursts through the outer layer somewhat like a jack-in-a-box to release the spores. The best known members of the class are important plant pathogens, including Phaeosphaeria nodorum (E. Müller) Hedjaroude, a major fungal pathogen of wheat and a model for fungicide development, and Venturia inaequalis (Cooke) G. Winter, the cause of apple scab disease. However, the majority of described Dothideomycetes are endophytes or saprobes growing on woody debris, decaying leaves, or dung. A few are lichens. We include just two species. One is Apiosporina morbosa, in the Ven-

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turiaceae family of the Pleosporales, a group that includes many destructive plant pathogens. The other is a sooty mold fungus, Scorias spongiosa from the family Capnodiaceae, order Capnodiales. The Capnodiales incorporate plant and human pathogens, endophytes, saprobes, and epiphytes, with many different nutritional modes. Sooty molds are epiphytes, forming harmless masses of black cells on plant leaves. Scorias spongiosa is associated with the honeydew of aphids that feed on plant sap. However, other members of the Capnodiales are able to parasitize humans and animals or are causal agents of economically important crop and tree diseases (e.g., members of the genus Mycosphaerella). There is a strong possibility that Catinella olivacea will be transferred from the Leotiomycetes to the Dothidiomycetes (Greif et al. 2007).

Overview

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Apiosporina morbosa (Schweinitz) Arx fruitbody 3.5–14 cm long, 1–2.5 cm thick, fusiform to clavate or irregularly elongated; outer surface hard, initially olivegreen soon becoming black, carbonaceous, fi nely roughened, typically furrowed and cracked, stalkless; flesh white when very young, soon black and britt le; perithecia embedded near the surface in a single layer. MICROSCOPIC FEATURES: spores 13–19 × 4– 7.5 μm, narrowly elliptic to clavate, 1–3-septate, smooth, pale yellowish brown; asci 50–75 × 10–15 μm, claMACROSCOPIC FEATURES:

Bl ack K not of Cher ry

vate, bitunicate; pseudoparaphyses septate, hyaline. OCCURRENCE: solitary or several, clasping and enveloping branches and twigs of cherry and plum trees; year-round; northern United States and Canada; very common. COMMENTS: Th is fungus is a destructive pathogen of plum trees and, to a lesser extent, cherry trees. Dibotryon morbosum (Schweinitz) Theissen and Sydow is a synonym.

Apiosporina morbosa

Dot h i deom ycet e s

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Scorias spongiosa (Schweinitz) Fries fruitbody a spongelike loose mycelial subiculum up to 15 or more cm broad and 6 or more cm tall on twigs and leaves of American beech (Fagus grandifolia), cream to buff and resembling a large yellow sponge with a gelatinous texture when young and in the asexual phase, eventually black in the sexual phase at maturity. MICROSCOPIC FEATURES: ascospores 12–15 × 2.5– 3 μm at maturity, yellowish hyaline with 3 septa, biseriate; asci 40–45 × 7 μm, 8–spored, obovateclavate, thick-walled, appearing as croziers in a pseudothecium on proliferating ascogenous hyphae; conidia produced in phialides; a dark brown pigment in the inner wall of conidiogenous hyphae becomes compacted and forms locules into which conidia are exuded in a slime-like matrix. MACROSCOPIC FEATURES:

Beech A phid Poop-Eater

exclusively on branches and leaves of American beech trees. The yellow stage fi rst appears during summer and fall and persists through winter and into spring, when the black stage develops; eastern North America; common. COMMENTS: Scorias spongiosa is one of the Capnodiales sooty molds that grow on insect honeydews or sugary plant exudates. Though most of the sooty molds that grow on leaves and conifer needles are quite small and thin, this species can be as large as a football and is distinctive. It can be found under colonies of beech tree aphids. The molds occur on the surface of the plant and cause no harm except possibly to reduce photosynthesis by blocking sunlight. OCCURRENCE:

Scorias spongiosa

Scor i a s

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Chapter 12 Taphrinomycotina

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he Ta phr inom ycotina subphylum (class Taphrinomycetes, order Taphrinales, family Taphrinaceae) contains only one really noticeable genus, Taphrina. Taphrina species are dimorphic plant parasites (Ray 1939; Mix 1947). They produce both a yeast state and a fi lamentous state that cause deformations in above-ground plant leaves, cat-

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kins, and branches. They do not form fruitbodies. Examples include witches’ brooms, leaf curls, and alder tongues. The Taphrinomycotina subphylum also contains the order Pneumocystidomycetes, with its single genus Pneumocystis. The species that causes Pneumocystis pneumonia in humans is classified in this genus.

Overview

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Taphrina occidentalis W. W. Ray

A lder Tongues

MACROSCOPIC FEATURES:

produces a hypertrophic condition of the bract, resulting in long, twisted, tongue-like outgrowths that are initially yellowish green, later becoming yellowish red, red, or purplish, and drying brown. MICROSCOPIC FEATURES: spores 2– 6 × 1.5–4 μm, elliptic, numerous; asci 35–55 × 10–20 μm, basal cells 8–17 × 12–22 μm (wider than long). OCCURRENCE: on female catkins of Alnus rubra and several other species of alders; summer; western North America; common. COMMENTS: Taphrina robinsoniana Giesenhagen is the common alder tongue of eastern North America. However, recent DNA reports suggest that Taphrina robinsoniana may be a complex of macroscopically and microscopically similar species (Rodrigues and Fonesca 2003). Taphrina robinsoniana has basal cells that are longer than wide and has shorter, narrower asci (23–39 × 7–11.5 μm) that are typically 8–spored, with the elliptical spores (2– 6 × 1.5–4 μm) sometimes budding in the ascus to form a polysporic condition. Taphrina alni (Berkeley and Broome) Gjærum (= Taphrina amento-

rum [Sadebeck] Rostrup) (not illustrated) is the most common European alder tongue. It has been found on Alnus rubra in Alaska. Taphrina alni has numerous spores per ascus, asci measuring 34–81 × 9–18 μm, and lacks basal cells. Taphrina occidentalis is distinguished from Taphrina alni by the presence of basal cells at the base of the asci. Its asci (35–55 × 10–20 μm) cannot easily be distinguished from those of Taphrina alni because they overlap in size. Many Taphrina species produce witches’ brooms, including Taphrina epiphylla (Sadebeck) Sadebeck (not illustrated) (asci 52 × 11–23 μm) on Alnus incana, Taphrina betulina Rostrup (not illustrated) (asci 21– 96 × 11–23 μm, with basal cells) on Betula pubescens, and Taphrina nana Johanson (not illustrated) (asci 29–85 × 9–29 μm, without basal cells) on Betula nana. Other Taphrina species, such as Taphrina populina (Fries) Fries (not illustrated), cause leaf spots. Taphrina populina produces golden yellow leaf spots on cottonwood and aspen, and has asci measuring 30–122 × 13–30 μm and globose spores that measure 4– 6.5 × 4–5 μm.

Ta phr inom ycotina

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Taphrina occidentalis

Ta phr ina

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Glossary

aborted: arrested in development and only partly formed acicular: needle-shaped acropetal succession: youngest at the tip acute: pointed or sharp-edged; compare with obtuse adnate: broadly att ached aerial: reaching upward into the air aethalia: fruitbodies of slime molds in which the plasmodia become aggregated into more or less pulvinate masses agaric: a gilled mushroom agglutinated: fi rmly att ached as if glued together; often used in reference to fibrils aleuriospore: an asexual, nonmotile spore allantoid: curved and sausage-shaped alveolate: having alveoli alveoli: small cavities or pit-like depressions (such as those on the head of a morel) amoeboid: resembling an amoeba amorphous: lacking a specific shape amyloid: staining blue, blue-gray, gray, or nearly black when exposed to iodine, such as in Melzer’s reagent or Lugol’s solution; compare with dextrinoid anamorph: the asexual or imperfect state of a fungus, characterized by the presence of conidia or absence of spores anastomosing: describes structures that split and rejoin in network fashion, like a braided stream angular: having distinct angles angular-oval: predominately oval with some angles annulate: bearing a ring apex: the uppermost part of something, such as a stalk, ascus, paraphysis, or spore; plural apices apical: occurring at the apex of a structure apical ring: thickening of the wall around the tip of an ascus

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apiculate: having one or more short projections apiculus: a short projection; plural apiculi aporhynchous: having a beak apothecium: a type of ascocarp characteristic of the discomycetes; an open-cup-shaped fruitbody with exposed hymenium; plural apothecia appressed: flattened onto a surface apud: in the work of; used in citing the work of an author contained in the publication of another author aquatic: pertaining to water arcto-boreal: pertaining to the arctic and cold northern regions areolar: pertaining to an area marked out on a surface and separated from other areas by color, cracks, or other markings arid: having litt le or no rainfall aromatic: having an aroma that is pleasant and often distinctive ascocarp: fruitbody of an ascomycete (= ascoma) ascoma: fruitbody of an ascomycete (= ascocarp); plural ascomata Ascomycete: a member of the Ascomycota Ascomycota: a phylum of fungi, characterized by production of sexual spores within an ascus ascospore: sexual spore produced in an ascus ascostroma: an undifferentiated mass of tissue or stroma, on which or in which the asci are developed; plural ascostromata ascus: a sac-like, usually elongated, cell in which sexual spores are formed; plural asci aseptate: not divided by cross-walls asexual: reproducing through mitosis only (without meiosis) asperulate: roughened with minute spines or warts; usually in reference to a spore

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attenuate: to reduce in thickness, or become gradually narrow avellaneous (color): variously interpreted from drab to pale grayish brown to hazel basal: at the lower end; nearest to the point of att achment base: the lowest portion of a stalk, cup, or other structure basidiomes: fruitbodies of members of the Basidiomycota Basidiomycete: a member of the large group of fungi, the Basidiomycota, bearing sexually produced spores on a basidium Basidiomycota: a phylum of fungi, characterized by production of spores on a basidium basidiospore: a spore formed on a basidium basidium: a microscopic club-shaped structure on which members of the Basidiomycota form spores beak: a somewhat pointed extension resembling the bill of a bird bicellular: having two cells bifurcate: forking by twos bigutt ulate: having two oil drops binomial: having two names bioethanol: ethanol produced by fermentation of plant starch biseriate: arranged in two rows in the ascus; compare with uniseriate and multiseriate bitunicate: type of ascus characterized by having a double wall; the outer wall is inelastic and ruptures at maturity, allowing the extensible inner wall to protrude and discharge the spores. boleticolous: pertaining to or associated with boletes; living on boletes boreal: pertaining to cold northern regions bryophytes: mosses, liverworts, and hornworts buff (color): pale creamy yellow, brownish yellow, or grayish yellow bulbil: a small bulb-like structure, especially in the axil of a leaf or at the base of a stem, that may form a new plant bulbous: enlarged at the base cadaver: the body of a dead organism calcareous: containing or characteristic of calcium, calcium carbonate, or limestone

capitate: having a head or small knob at the tip carbonaceous, carbonous: black and britt le, like charcoal carnivorous: feeding on animals carotenoid: having yellow, orange, or red pigments cartilaginous: fi rm and tough but pliable catenulate: stuck together in chains cellular: composed of more or less spherical cells cellulase: any of several enzymes that break down cellulose cellulose: a polysaccharide found in cell walls of some species cellulosic: pertaining to cellulose cerebriform: highly convoluted, appearing brainlike cespitose: growth form (habit) in which stalks of several fruitbodies arise very close together but are not fused cf.: abbreviation for conferre; states that a determination is uncertain or provisional chamber: a defi ned enclosed hollow space chambered: having two or more defi ned enclosed hollow spaces chitin: a nitrogen-containing polysaccharide found in many species of fungi chitinase: an enzyme that breaks down chitin chlamydospore: a thick-walled spore produced asexually ciliate: fringed with hairs, eyelash-like circinate: ring-shaped; rolled up in the form of a coil with the tip in the center circumboreal: distributed throughout the far northern regions citrine (color): a color variously interpreted as a light greenish yellow or sometimes lemon yellow citron (color): variously defi ned as greenish yellow or lemon yellow with olive tones clade: a group of organisms believed to have evolved from a common ancestor clavate: club-shaped clay: a stiff, sticky, fi ne-grained earth, often forming an impermeable layer in the soil clayey: having a large percentage of small clay particles cleistothecium: closed ascocarp that opens by rupturing; plural cleistothecia

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cm: abbreviation for centimeter, a unit of measurement in the metric system equal to 1/100 meter, approximately 3/8 inch coalescent: grown together and fused into a single unit coevolved: the evolutionary state of two or more species that have undergone a series of reciprocal adaptive changes in relation to each other coleopterous: of or pertaining to beetles columella: a column of sterile tissue that penetrates the gleba of some truffle-like species compressed (stalk shape): flattened in cross section compressed-spherical: round and somewhat flattened concave: curved inward like the interior of a circle concentric: circles or arcs that extend outward from a common center concolorous: having the same color as something else confluent: merging together conic: cone-shaped conical-papillate: cone-shaped and having a small nipple-shaped projection conidia: asexual spores, produced by mitosis rather than meiosis conidiogenous: producing or giving rise to conidia conidiophore: a specialized hypha that bears conidia conifer: a cone-bearing, usually evergreen, tree such as pine, spruce, or Douglas fi r conoid: cone-shaped contorted: twisted or bent out of the normal shape convex: curved outward like the exterior of a circle convoluted: intricately folded or twisted convolutions: intricate folds or twists coremia: a special anamorphic stroma that produces conidia cortex: rind; outer layer cosmopolitan: pertinent or common to the whole world cotton blue: dye or stain used microscopically to test for color changes; see cyanophilic crenate, crenulate: scalloped, fi nely scalloped; usually in reference to the edge of an apothecium crescent: having a curved sickle shape crescentic: pertaining to a curved sickle shape crimped: compressed into small folds or ridges

crozier: the crook or hook on ascogenous hyphae, prior to the formation of the asci culm: stem of any plant cuneate: wedge-shaped cupulate: cup-shaped cyanophilic: staining blue with the dye cotton blue cylindric: having nearly the same diameter throughout the length cylindric-capitate: having nearly the same diameter throughout and a head or small knob at one end cylindric-clavate: having nearly the same diameter throughout the length but slightly club-shaped cylindric-elliptic: having nearly the same diameter throughout the length with rounded ends and somewhat curved sides cylindric-oblong: having nearly the same diameter throughout but longer than wide and with somewhat flattened ends cylindroclavate: having nearly the same diameter throughout but also somewhat club-shaped cymbiform: shaped like a boat cytoplasm: the contents of a cell exclusive of the nucleus de Bary bubbles: air bubbles or water-vapor bubbles in ascospores that sometimes appear when oil droplets disappear deciduous: shedding leaves annually decorticated: without bark dehiscent: splitt ing open delimiting: determining the limits or boundaries of something deliquescing: disappearing as if by melting dendritic: having a branched appearance dentate, denticulate: toothed, fi nely toothed denticles: small tooth-like projections dextrinoid: staining dark red to reddish brown when exposed to iodine, such as with Melzer’s reagent or Lugol’s solution; compare with amyloid dichotomous: split in two, or forking, like a wishbone or tuning fork dictyospore: a spore having both longitudinal and transverse septa diminutive: of small stature dimorphic: having two forms disarticulate: separating into distinct units

Glossary

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disc: = disk discoid: shaped like a shallow dish disk: saucer-shaped, like a flat circular plate distal: located away from the point of origin or att achment dresden brown: brown with a tiny hint of orange duff: plant litter making up the surface layer of forest soil; usually applied in conifer forests eccentric: away from the center, off-center echinate: covered with pointed spines echinulate: covered with small pointed spines echinulate-papillate: having small pointed spines on a small nipple-shaped elevation ectal excipulum: the outermost portion of the excipulum; lies outside of the medullary excipulum eff used: spread out, expanded; often having a number of separately erumpent perithecia egutt ulate: without oil drops Elata clade: a section of black morels evolved from a common ancestor ellipsoid, elliptic: elongated with rounded ends and curved sides; usually in reference to spores ellipsoid-inequilateral: elongated with rounded ends and unequal sides elliptic-fusoid: more or less spindle-shaped and somewhat elliptical elongate: longer in relation to the width encrustation: a crust-like or hard coating encrusted: having a crust-like or hard coating endophyte: an organism growing inside another organism endophytic: pertaining to an organism growing inside another organism enfolded: held within limits or wrapped up within entire: even, smooth, not toothed or broken up; usually in reference to the margin or edge of an apothecium ephemeral: lasting for a very short time epidermis: the outer layer of a surface epigeous: fruitbodies that grow on or above ground; compare with hypogeous epiphyte: growing on the outside of a plant, but not as a parasite epispore: the fundamental layer of a spore wall that usually gives rise to ornamentation; usually the thickest layer episporium: an epispore

epithecium: the layer above the asci; usually formed by the tips of the paraphyses epithet: the second half of a species’ scientific name (the fi rst half being the genus to which the species belongs) equal: of the same diameter throughout; usually in reference to a stalk ericaceous: denoting plants of the heath family erumpent: breaking through or bursting out Esculenta clade: a section of yellow morels evolved from a common ancestor evanescent: quickly fading or disappearing excipular: pertaining to an excipulum excipulum: the outer, sterile portion of an apothecium; often treated as consisting of medullary and ectal layers exosporium: the layer that covers the epispore and becomes the outermost layer after the perispore disappears exude: to gradually ooze or discharge farinose: covered with mealy or flour-like particles farinose-furfuraceous: covered with flour-like and coarsely granular particles fascicle: group or bundle fasciculate: arranged in groups or bundles fermentation: the process of converting sugar into alcohol in the absence of oxygen fertile: spore-bearing fertile surface: layer of tissue on which the sporebearing structures are produced (= hymenium) fiber: a hair-like structure fibril: a slender fiber fibrillose: composed of fibrils fibrous: tough and stringy fi lamentous: composed of fi laments fi liform: long and very slender fissured: split or cracked to form a long, narrow opening flexuous: bent alternately in opposite directions floccose, flocculose: with tuft s, or small tuft s, of loose cottony material fluted: having longitudinal ridges or ribs free: not att ached fruitbody: macroscopic sexual reproductive structure produced by certain fungi; also known as sporocarp, carpophore, ascocarp, ascoma, basidiome, basidiocarp, or mushroom

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fuliginous (color): smoky or sooty grayish brown, dark and dusky fulvous (color): reddish cinnamon; colored like a red fox furfuraceous: covered with coarsely granular or branlike particles; = scurfy furrow: line, ridge, or groove furrowed: marked with lines, ridges, or grooves fuscous (color): dark brownish gray to brownish black fusiform: spindle-shaped fusiform-clavate: fundamentally club-shaped but somewhat spindle-shaped fusiform-elliptic: elliptic and somewhat spindleshaped fusoid: tapering from the middle toward both ends; somewhat spindle-shaped fusoid-elliptical: fundamentally elliptic but somewhat spindle-shaped gangrenous: relating to death and decomposition of body tissue gelatinous: having a jellylike consistency genera: plural of genus genus: a formally designated group of similar, presumably closely related, species germination: the fi rst process in the transformation of a spore to some other structure in the life cycle of a fungus germination cleft: a fi ssure or crack in the spore wall that facilitates germination germ slit: a fi ssure or crack in the spore wall that facilitates germination glabrous: smooth, naked gleba: a fleshy, gelatinous or powdery spore-bearing inner mass globoid: pertaining to a sphere globose: round like a sphere globose-ovate: almost-round to somewhat egg-shaped globose-papillate: round with a small, nipple-shaped projection globular: spherical or nearly so granulose: covered with small sugar-like particles gregarious (habit): fruitbodies occurring close together in a more or less dense group gristly: resembling cartilage in meat gutt ulate: containing oil drops

gutt ule: oil drop habit: morphological form or manner of growth, such as solitary, scattered, gregarious, or clustered habitat: place where an organism lives haploid: having one-half the number of chromosomes hardwood: flowering trees with broad leaves, such as alder, beech, birch, hickory, and oak; may be deciduous or evergreen; (= broadleaf) hemicellulase: an enzyme that breaks down hemicellulose hemicellulose: a polysaccharide found in plant cell walls that is less complex than cellulose hemispheric: pertaining to half of a sphere herbaceous: describing plants that have no (or very litt le) woody tissue. Most herbaceous plants live for only one growing season. herbarium: a permanent organized repository for dried, or otherwise preserved, fungus or plant specimens herbivore: organism that consumes plants hexagonal: having six angles hilum: the point of att achment on a spore hirsute: covered with long, stiff hairs hispid: covered with stiff hairs or bristles hoary: having a whitish to grayish sheen, like frost homogenous: composed of uniform tissue or the same substance horny: tough and hard; resembling animal horn humus: the organic component of soil hyaline: transparent and nearly colorless hydrazine: a toxic alkaline, colorless, volatile liquid used in fuels for rocket and jet engines hymenium: the spore-producing, or fertile, tissue of a fruitbody, such as the layer of asci and paraphyses lining the inner surface of an apothecium; plural hymenia hypertrophic: pertaining to an overgrowth or excess enlargement hypha: long tube-like element that, with many others, makes up the body (mycelium) of a fi lamentous fungus; may or may not be septate; plural hyphae hypogeous: growing below ground; compare with epigeous immunocompromised: having an immune system that is damaged or weakened inamyloid: remaining colorless or yellowish in Melzer’s reagent

Glossary

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incised: as if cut into inclusions: structures or substances contained within incurved: bent or depressed inward indehiscent: not splitt ing open to release spores indeterminate: without restrictions, or unlimited inequilateral: having unequal sides inflorescence: the complete flowerhead of a plant inoperculate: a type of ascus in which the pore through which the spores are released has no lid; compare with operculate insipid: lacking flavor or interest interactive: acting on each other intercalary: between the apex and the base involute: inrolled, especially in reference to the margin of an apothecium keratin: a tough, insoluble protein that is the chief component of hair, nails, feathers, horns, and hooves keratinous: composed of keratin KOH: chemical formula for potassium hydroxide, usually in a 3-5 percent aqueous solution; used as a mounting medium for microscopic observations and, both microscopically and macroscopically, to test for color changes labyrinth: an intricate structure consisting of interconnecting passages; a maze labyrinthine: relating to or constituting a labyrinth lacerate: cut or torn and sometimes ragged laciniate: torn or coarsely cut lactic acid: a chemical compound used macroscopically to test for color changes; chemical symbol C3H6O3 lacunose: covered with pits or indentations lageniform: flask- or gourd-shaped lanceolate: lance-shaped; widened and then tapering to a point lateral: pertaining to the side lenticular: shaped like a double convex lens lignin: a complex organic chemical deposited in the cell walls of many plants, making them rigid and woody lineate: arranged as or marked with lines or streaks loam: fertile soil composed of a mixture of clay, sand, and silt loamy: having the characteristics of loam lobe: a rounded division

lobing: having lobes locule: a cavity, especially one occurring in a stroma within which the hymenium is produced Longibrachiatum clade: one of two remaining sections, following phylogenetic analysis, now used for the classification of Trichoderma species longitudinal: running along the length rather than across lubricous: slightly greasy or slippery to the touch lunate: crescent- or half-moon-shaped macrofungi: moderately large to large species of fungi macroscopic: visible with the unaided eye, or at low magnification, such as obtained with a 10-power hand lens macrospecies: see macrofungi marbled: marked with streaks of a different color margin: edge; especially in reference to an apothecium maritime: of or relating to the sea maze: an intricate structure consisting of interconnecting passages; a labyrinth median: approximately in the middle medullary excipulum: the innermost portion of the excipulum; lies between the hymenium (or hypothecium, if differentiated) and ectal excipulum meiosis: a type of cell division in which a parent cell produces four haploid nuclei Melzer’s reagent: an iodine-containing solution used, both microscopically and macroscopically, to test for color changes; see amyloid, nonamyloid = inamyloid, and dextrinoid μm: abbreviation for micrometer, a unit of measurement in the metric system equal to one-millionth of a meter; informally (and decreasingly) referred to as a micron microscopic: visible only with a microscope mitosis: a type of cell division in which a parent cell produces two daughter cells, each genetically identical to the parent cell mitre: a tall, pointed, cleft cap worn by a bishop mm: abbreviation for millimeter, a unit of measurement in the metric system equal to 1/10 centimeter and 1/1000 meter, or approximately 1/25 inch moniliform: cylindrical with regular constrictions, like a string of beads monomorphic: existing in only one form or shape

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monophyletic: arising from one ancestral group montane: pertaining to mountainous areas morphological: having the form and structure of an organism or other object mott led: marked with spots or smears of one or more different colors mucilage: a slimy or sticky substance mucilaginous: slimy or sticky multicellular: composed of more than one cell multichambered: having several to many chambers multifurcate: branched repeatedly multigutt ulate: having several to many oil drops multinucleate: having several to many nuclei multiseptate: having several to many cross-walls multiseriate: arranged in several to many rows in the ascus; compare with biseriate and uniseriate mummified: shriveled and dried up mushroom: a loosely defi ned term for the large conspicuous fruitbodies of certain fungi, often confi ned to those with fleshy texture, those with cap and stalk, or those bearing gills mycelium: the body of a fi lamentous fungus, composed of a network of complexly branched hyphae; plural mycelia mycologist: one who studies fungi mycology: the study of fungi mycoparasite: a fungal parasite on another fungus mycorrhizae: symbiotic associations of fungi and the roots of plants mycorrhizal: pertaining to a nearly universal mutualistic symbiosis between the hyphae of fungi and the roots of plants mycosis: a fungal infection; plural mycoses mycota: all the species of fungi that inhabit a given area navicular, naviculate: boat-shaped, in two senses— as seen from above with fusiform or rounded ends like the diatom Navicula, or as seen from the side, inequilaterally fusiform or more or less keeled necrotrophic: deriving food from dead cells, tissues, or their products nodulose: with small bumps (nodules); knobby nom. prov.: abbreviation for nomen provisorium, a term meaning “provisional name” nonamyloid: remaining colorless or yellowish in Melzer’s reagent

nonapiculate: lacking short projections nonchitinoid: lacking chitin nonseptate: lacking cross-walls; = aseptate obclavate: the opposite of clavate; having a thinner distal end obconic: reversely conical, as an inverted cone oblique: slanting, not parallel or at right angles to another line or surface oblong: longer than wide, with more or less parallel sides and somewhat flattened ends oblong-ellipsoidal: longer than wide, with rounded ends and curved sides oblong-elliptical: longer than wide, with rounded ends and curved sides oblong-fusiform: longer than wide and somewhat spindle-shaped oboval, obovate, obovoid: inversely ovate (eggshaped), with broadest portion uppermost obpyriform: thickest above the middle; opposite of pear-shaped obtuse: blunt, not pointed; compare with acute ocher (color): pale brownish orange-yellow; also spelled ochre ochraceous: having an ocher color oligotrophic: relatively low in plant nutrients olivaceous: having an olive green color opaque: not allowing light to pass through operculate: a type of ascus in which the pore through which the spores are released has a lid (operculum), like the hatch of a ship; compare with inoperculate operculum: a lid like the hatch of a ship; plural opercula ordinal: pertaining to something in a specific series organic: derived from living matter orifice: an opening ornamentation: the surface features of a nonsmooth spore, such as spines, warts, ridges, or wrinkles ostiolar: pertaining to an ostiole ostiole: a mouth or opening; used to refer to the canal in the tip of a perithecium and to the opening of a pycnidium oval: egg-shaped oval-cylindric: egg-shaped but having nearly the same diameter throughout ovate, ovoid: egg-shaped

Glossary

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palisade: an arrangement of elongated, perpendicular cells pallid: a very pale shade of any color, almost white papilla: a small, nipple-shaped elevation; plural papillae papillate: having a papilla papyraceous: resembling paper parallel: being the same distance apart at all points paraphysis: sterile hyphal ends that, along with the asci, comprise the hymenium of apothecia; plural paraphyses parasite: an organism that obtains its nutrients from another living organism (its host), causing a negative impact as a result parasitized: having one organism living in or on another organism and deriving food from it parenchyma: the soft tissue of higher plants that consists of thin-walled cells part-spore: a single aseptate spore detached from a larger multiseptate spore pathogen: a parasitic or disease-causing organism pathogenic: able to cause disease perennial: lasting or remaining active for many years peridial: pertaining to a peridium peridium: the outermost layer of a fruitbody such as a truffle or puffball peripheral: related to or located on the outer boundary or margin perispore: the outermost membrane layer that covers all the other layers of the spore wall and often disappears as the spore matures perithecial: pertaining to a perithecium perithecium: a flask-like fruitbody characteristic of some ascomycetes; usually many are embedded in a mass of fungus or host tissue; plural perithecia persistent: retaining its shape or position for a long time petiole: the stalk that joins a leaf to a stem phallic: resembling an erect penis phialide: a flask-shaped structure that bears conidia phragmospore: a spore with two or more transverse septa in one plane phragmosporous: having two or more transverse septa phylogenetic tree: a branching diagram showing evolutionary relationships

phytopathogenic: having the ability to cause disease in plants pinard yellow (color): a light greenish yellow planar: relating to or forming a plane or flat surface plane: a flat or nearly flat surface plasmodia: aggregates of protoplasm that exhibit amoeboid motion pleomorphic: having more that one form or shape pleurorhynchous: having a lateral beak polar: relating to opposite ends polyblastic: having multiseptate spores, or giving rise to many spores polygonal: having many angles polyhedral: many-sided polymorphic: having several to many shapes polyphyletic: arising from several ancestral groups polysaccharide: a complex sugar polysporic: many-spored pore: small opening, such as occurs at the tip of an ascus or perithecium predator: an animal that normally preys on and consumes other animals prophialide: a structure that produces a phialide protoplasm: the living part of a cell, including the nucleus, cytoplasm, and organelles protunicate: a type of ascus that is thin-walled, typically globose to broadly clavate, and passively releases spores by disintegration of the ascus wall proximal: located near the point of origin or att achment pruinose: appearing to be covered with a fi ne powder pseudoapiculi: projections on a spore that do not function as att achment structures pseudoepithecium: an outer layer above the paraphyses that is usually gelatinized and often sloughs off pseudoparaphyses: vertical hyphal thread-like fi laments att ached to the inside top of a fruitbody, growing downward and att aching to the inside base pseudoparenchyma: mass of tissue having the form of parenchyma cells, which are nearly isodiametric and look like crowded soap bubbles when in mass pseudorhiza: a root-like extension that forms a union between the fruitbody and mycelium

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pseudosclerotium: a mass of substratum held together by mycelium to form a defi nite body resembling a sclerotium pseudoseptate: having a membrane that deceptively appears like a septum pseudothecia: a globose perithecium-like fruitbody pubescent: covered with short, soft , fi ne hairs pulvinate: cushion-shaped punctate: covered with tiny scales or spots pungent: having a strongly sharp taste or smell pustulate: appearing as though covered with blisters pustules: pimple-like or blister-like elevations pycnidial: pertaining to a pycnidium pycnidium: more or less closed cavity in which conidia are produced; plural pycnidia pyramidal: resembling a pyramid pyrenomycetes: an informal term for the Sordariomycetes pyriform: pear-shaped rays: recurved and often triangular portions of a fruitbody that has split open recurved: curved backward reflexed: bent downward and outward reniform: kidney-shaped repand: having a margin that is wavy and turned back or elevated resinous: sticky and soft or hard, like pitch or tar resupinate: lying flat on the surface reticulate: having a net-like pattern reticulate-echinulate: having a net-like pattern and small pointed spines reticulum: net-like system of ridges; often in reference to a spore revivable: able to resume natural shape and functions when placed in moist conditions rhizome: a horizontal underground stem rhizomorph: a group of thick, rope-like strands of hyphae growing together as a single organized unit rhomboid: a four-sided structure of which only the opposite sides and angles are equal rhomboidal: pertaining to a rhomboid ribbed: having a pattern of raised bands riparian: relating to wetlands adjacent to rivers and streams rosaceous: denoting plants of the rose family rudimentary: immature or poorly developed

rugose, rugulose: wrinkled, fi nely wrinkled saccate: shaped like a sack Salicaceae: willow family of flowering plants saprotroph: an organism that obtains its nutrition by decomposing dead organic material saprotrophic: pertaining to a saprotroph scabrous: covered with scales scale: an erect, flattened, or recurved projection or torn portion of a surface scalloped: edged with a series of curved projections scattered (habit): fruitbodies occurring closely enough together to be considered a group, but farther apart than for gregarious Sclerotiniaceae: a family in the Leotiomycetes with species that often form sclerotia sclerotium: a hard, compact mass of hyphae, and sometimes soil, that is a resting stage for some fungi, and from which a fruitbody may arise; plural sclerotia scolecospore: a long fi liform spore having septa in one plane sculptured: having raised or incised markings such as reticulation, warts, or spines scurfy: covered with coarsely granular or bran-like particles; = furfuraceous scutellate: shaped like a small shield SEM: scanning electron microscopy semi-eff used: partly spread out sensu auct.: in the sense of various authors; sensu auctorum sensu lato: in the broadest sense of sensu stricto: in the strictest sense of septate: divided by cross-walls (septa) septum: a cross-wall, usually in reference to a spore or hypha; plural septa serpentine: winding, resembling a snakeform or movement sessile: att ached at the base, without any sort of stalk seta: a usually thick-walled, yellow or brown bristle or hair-like element; plural setae setose: having setae sigmoid: S-shaped sinuous: wavy sinus: a bent surface or curve solitary (habit): fruitbodies occurring singly, without others nearby

Glossary

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sordid: dirty or dingy spathulate: ovate with a narrowed base, shaped like a spoon or spatula species: the lowest category in the classification of organisms (although some species are further divided into subspecies, varieties, or forms); variously defi ned, but can be considered to represent a single kind or type of organism spherical: more or less round, shaped like a ball spinose, spinulose: covered with spines or small fi ne spines spiraling: winding up or down around a fi xed central axis spore: single- or sometimes multi-celled, sexual or asexual reproductive propagule formed by fungi and plants such as ferns and mosses; functionally similar to the seeds of gymnosperms and flowering plants sporodochium: a spore mass supported by a superficial cushion-like mass of short conidiophores; plural sprodochia sporulate: to produce and release spores squamules: small scales squamulose: covered with small scales (squamules) ssp.: abbreviation for subspecies stalk: the structure that arises from the substrate and supports the head or cap of a fruitbody stature: overall form (such as tall and slender, or short and stocky) stellate: star-shaped sterigma: tiny slender stalks on which basidiospores are borne; plural sterigmata sterile: infertile stipitate: having a stalk striate: having fi ne parallel or radiating lines or grooves; often in reference to a spore striations: fi ne parallel or radiating lines or grooves stroma: cushion-like tissue on or in which the fruitbodies are produced; plural stromata stromatized: made into a stroma subacute: somewhat pointed or sharp-edged suballantoid: somewhat sausage-shaped subalpine: located on higher mountain slopes just below the tree line subangular: somewhat angular subcapitate: having somewhat of a head or small knob subcespitose: somewhat or nearly cespitose

subclade: a subgroup of a clade subclavate: somewhat club-shaped subconic: somewhat cone-shaped subcylindric: approximately cylindrical subcymbiform: somewhat boat-shaped subdiscoid: somewhat like a saucer or plate subellipsoidal: somewhat elongated with rounded ends and curved sides subfusiform: somewhat spindle-shaped subfusoid: somewhat spindle-shaped subgelatinous: somewhat gelatinous subglobose: nearly round subhyaline: almost transparent and nearly colorless subiculum: a more or less dense felty or cobwebby mat of hyphae covering the substrate and from which the fruitbodies arise subimmersed: partially covered by water, or partially embedded in something submedian: located just below the middle submerged: covered over by water suboperculate: between truly operculate and inoperculate subparallel: nearly parallel but diverging or converging slightly subreticulate: having a somewhat net-like pattern subsessile: att ached at the base by an inconspicuous stalk subspathulate: shaped somewhat like a spoon subspheroid: shaped somewhat like a sphere substipitate: having a very short att achment substrate, substratum: substance on or within which the vegetative body of the fungus is growing and from which the mushroom arises; usually soil, woody material, or leaf litter, but can be such things as insects or other mushrooms; plural substrata subterranean: existing underground subtruncate: appearing somewhat cut off subturbinate: top-shaped with a somewhat flattened apex sulcate: having parallel or radial grooves, the depth of which is greater than in striate and less than in plicate superficial: occurring at or on the surface symbionts: organisms that live together sympodial (of conidiogenous cells): having a new apex that originates below and to one side of a pre-

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vious apex after the main axis has produced a terminal spore synnemata: erect reproductive structures composed of conidiophores that fuse together, forming strands that resemble a stalk of wheat synonym: an alternate name for an organism tawny (color): dull yellowish brown, like a lion’s coat teleomorph: the sexual or perfect state of a fungus, characterized by the presence of spores formed through meiosis, such as ascospores or basidiospores temperate: relating to or denoting a region or climate characterized by having a mild temperature ten-pin (shape): resembling a bott le or bowling pin tentacles: slender and somewhat flexible appendages terete: rounded like a broom handle terminal: pertaining to the end or extremity terminus: the end or extremity terrestrial: growing on the ground textura angularis: a short-celled tissue of polyhedral cells without intercellular spaces textura globulosa: a short-celled tissue of round cells with intercellular spaces thelephores: crust fungi and thin-fleshed bracket fungi with smooth or irregularly textured sporeproducing surface (no pores, gills, or spines) tomentose: covered with long, soft , densely matted hairs; wooly tomentum: covering of long, soft , matted hairs transecting: cutt ing or dividing crossways translucent: allowing light but not clear images to pass through transparent: allowing light to pass through so that objects can be distinctly seen transverse: running crosswise rather than longitudinally trilobate: having three lobes trooping: having large numbers of fruitbodies in a dense group truncate: ending abruptly, appearing chopped off; usually in reference to spores or the fruitbodies of some otideas tubercle: a small rounded projection

tuberculate: having tubercles tuberous: round and swollen turbinate: top-shaped, like an inverted cone turgor: rigidity caused by absorption of fluid ubiquitous: widely distributed in nature umber (color): dark yellowish brown umbilicate: having a small central depression, like a navel undulate: wavy undulate-rugose: wavy and wrinkled ungulates: warm-blooded animals with hooves unicellular: consisting of a single cell unigutt ulate: having a single oil drop uniseriate: arranged in a single row in the ascus; compare with biseriate or multiseriate unitunicate: a type of ascus that has two walls that function as a single unit urceolate: pitcher- or urn-shaped, rounded with the walls curving inward toward the opening at the top vacuolar: having small cavities (vacuoles) vacuolate: possessing vacuoles vacuole: a small cavity or space that is often bound by a membrane and fi lled with fluid var.: abbreviation for variety velutinous: covered with short, fi ne, soft hairs, like velvet vermiform: worm-shaped verrucose, verruculose: warty, fi nely warty villose: covered with long, thin, straight, noninterwoven hairs vinaceous (color): said to be that of red wine; in practice, not so deeply colored and with duller gray to brown tones yielding more of a dark salmon-pink violaceous: having purplish tones viscid: sticky viscid-gelatinous: sticky and having a jellylike consistency volatile: easily converted from a liquid or solid into a gas that can be inhaled xeric: pertaining to a lack of moisture, as in a dry habitat

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Photo Credits

Jacket/Cover Sclerotinia sulcata by Noah Siegel

Dedication Kit Scates Barnhart, by Harley Barnhart

Introduction Chorioactis geaster by Jim Murray Light Microscopy, pages 2–9 and 11–12 by Lawrence Leonard SEM Images page 10 by Rosanne Healy

Picture Key Sean Abbott: 8c12 Tuber cf. rufum, 17a5 Gyromitra ambigua, 19b4 Helvella unicolor, 66b4 Rhytisma salicinum Harley Barnhart: 15b7 Morchella cf. prava, 21b1 Cookeina sulcipes, 21b2 Cookeina tricholoma, 22b7 Xylaria hypoxylon, 22b8 Xylaria filiformis, 24b2 Cordyceps washingtonensis, 31c2 Microstoma floccosum, 33a3 Coryne atrovirens, 33c1 Cudonia circinans, 36a3 Geoglossum peckianum, 41b1 Tricharina gilva, 44a3 Plicaria endocarpoides, 66a5 Lasiosphaeria spermoides, 73a12 Melastiza chateri Kit Scates Barnhart: 3b1 Sarcosphaera coronaria, 11b1 Verpa bohemica, 11b2 Verpa bohemica yellow form, 14b1 Morchella snyderi, 14b9 Morchella cf. elata, 14b11 Morchella populiphila, 17a1 Gyromitra esculenta, 17a4 Gyromitra infula, 19a6 Helvella vespertina, 19b2 Helvella acetabulum, 21a4 Helvella fibrosa, 21a6 Tarzetta catinus, 33c3 Neolecta irregula-

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ris, 35b6 Spathularia flavida, 42a1 Geopora arenosa, 44a1 Gyromitra ancilis, 44b6 Peziza sublilacina, 50b1 Pseudosarcosoma latahense, 62a2 Otidea cf. concinna, 62a8 Wynnella silvicola, 64b2 Claviceps purpurea George Barron: 25b9 Roseodiscus subcarneus, 62b10 Geopora sepulta, 71b5 Belonidium sulphureum Alan and Arleen Bessette: 17b3 Gyromitra sphaerospora, 19a4 Helvella sulcata, 19b5 Helvella costifera, 25b6 Mitrula lunulatospora, 35b5 Spathularia flavida yellow form, 35b7 Spathulariopsis velutipes, 36a1 Geoglossum alveolatum, 36c1 Trichoglossum farlowii, 38a10 Hypomyces luteovirens, 38c3 Hypomyces chlorinigenus, 42a3 Rhodotarzetta rosea, 44b1 Peziza echinospora, 50b4 Camarops petersii, 50c6 Ascotremella faginea, 56a1 Diatrype stigma, 58a3 Annulohypoxylon cohaerens, 58b4 Hypoxylon ferrugineum, 58b5 Hypoxylon rubiginosum, 61a1 Sarcoscypha austriaca, 62b2 Peziza phyllogena, 62b3 Peziza badia, 62b7 Peziza arenaria, 66b2 Rhytisma cf. americanum, 68b2 Hydropisphaera peziza, 68b5 Hypocrea cf. chlorospora, 72c7 Chlorencoelia versiformis, 73a1 Acervus epispartius, 73a2 Neott iella rutilans, 73b9 Lamprospora crechqueraultii Michael Beug: 4b1 Geopora cooperi, 4b5 Hydnotrya variiformis, 8c11 Tuber cf. separans, 14b2–3 Morchella snyderi, 14b6 Morchella brunnea, 14b8 Morchella elata, 14b10 Morchella fr ustrata, 15b6 Morchella prava, 17b2 Gyromitra californica, 17c2 Gyromitra montana, 19a2 Helvella crispa, 19a5 Helvella dryophila, 19a7 Helvella maculata, 19b3 Helvella solitaria, 20c2 Helvella elastica, 20c6 Helvella compressa, 20c7 Helvella albella, 21a5 Pseudoplectania vogesiaca, 21c1 Sarcoscypha coccinea, 29c3 Pachycudonia monticola, 30b20 Chloro-

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ciboria aeruginosa, 34a2, 42a2 Geopyxis carbonaria, 34a3 Geopyxis vulcanalis, 34a4 Neournula pouchetii, 34a6 Helvella leucomelaena, 38a4 Hypomyces lateritius, 38d9 Hypomyces rosellus, 38e5 Helminthosphaeria clavariarum, 38e8 Hypomyces cervinigenus, 44b3 Peziza proteana f. sparassoides, 46b1 Peziza vesiculosa, 47a2 Pseudombrophila cervaria, 47a7 Cheilymenia theleboloides, 50b2 Urnula padeniana, 50b3 Plectania milleri, 50b5 Plectania melastoma, 60a1 Gyromitra olympiana, 60a2 Gyromitra ancilis, 60a3 Gyromitra mwb060911, 60a5 Gyromitra melaleucoides, 60a6 Disciotis venosa, 61b1, 3 Caloscypha fulgens, 62a3 Otidea leporina, 62a6 Otidea onotica, 62b5 Peziza varia, 64b1 Claviceps purpurea, 65b3 Taphrina occidentalis, 66b1 Coccomyces dentatus, 70a5 Scutellinia scutellata, 72b2 Pithya vulgaris, 72b6 Pithya cupressina, 72c11 Ciboria rufofusca Michael Castellano: 4a1 Gilkeya compacta, 6a4 Elaphomyces cf. anthracinus, 6c2 Elaphomyces cf. granulatus, 6c3 Elaphomyces cf. muricatus, 6c6 Elaphomyces subviscidus, 8a1 Balsamia nigrans, 8b1 Choiromyces alveolatus Adolf and Oluna Ceska: 21a1 Helvella corium, 30b1 Bisporella sulfurina, 30b2 Bisporella subpallida, 30b11 Lanzia luteovirescens, 31a2 Lachnellula cf. flavovirens, 31a3 Brunnipila cf. calyculiformis, 31a4 Velutarina rufo-olivacea, 31b2 Lachnellula gallica, 31c3 Dasyscyphella nivea, 31c6 Lachnum pudibundum, 36a5 Geoglossum umbratile, 36a7 Geoglossum simile + 36a6 asci and spores, 38d7 Protocrea farinosa, 43b7 Byssonectria fusispora, 47a3 Pseudombrophila theioleuca, 47a6 Lasiobolus cuniculi, 47b2 Ascobolus stercorarius, 49a3 Bulgariella pulla, 68b4 Neonectria fuckeliana, 68b7 Hypocrea cf. sinuosa, 71b4 Hyaloscypha britannica, 72a2 Orbilia xanthostigma, 72a3 Orbilia inflatula, 72c1 Mollisia cinerea, 72c4 Mollisia discolor, 72c5 Hyaloscypha albohyalina, 72c6 Patellariopsis clavispora, 73a4 Wilcoxina rehmii, 73a10 Belonopsis graminea, 73b5 Discinella boudieri Jules Cimon: 21a3 Helvella macropus, 25b11 Neocudoniella radicella, 29c2 Neocudoniella albiceps, 30b7 Crocicreas coronatum, 30b15 Rutstroemia bulgarioides, 30b16 Holwaya mucida and Crinula caliciiformis, 38b1 Hypomyces ochraceus (Cladobotryum verticillatum), 38c1 Hypomyces chrysospermus, 41b5

Trichophaea hemisphaerioides, 47a4 Pseudombrophila merdaria, 50c4 Neobulgaria pura, 57a3 Daldinia childiae, 63c2 Encoelia furfuracea, 72a1 Orbilia delicatula, 72c10 Catinella olivacea, 73a7 Podophacidium xanthomelum Dale Dunaway: 22b9 Xylaria oxyacanthae, 24b5 Cordyceps olivascens, 43a3 Plicaria carbonaria Todd Elliott: 5c3 Pachyphloeus virescens, 5c4 Pachyphloeus carneus, 6a1 Elaphomyces cf. loebae, 6a6 Elaphomyces macrosporus, 6c4 Elaphomyces americanum, 6c5 Elaphomyces verruculosus, 7a1 Imaia gigantea, 15a1 Morchella cf. diminutiva, 15b1 Morchella cf. americana, 17c5 Gyromitra brunnea, 20c1 Helvella latispora, 20c5 Helvella pezizoides, 22b11 Xylaria magnoliae, 22b12 Xylaria tentaculata, 14b1 Cordyceps militaris, 24b3 Cordyceps hesleri, 24b4 Elaphocordyceps subsessilis, 24b6 Akanthomyces aculeatus, 24b7 Cordyceps tuberculata, 24b9–11 Beauveria bassiana, 14b13 Gibellula leiopus, 24b17 Ophiocordyceps cf. gracilis, 24b18 Ophiocordyceps sphecocephala, 24b19 Ophiocordyceps ravenelii, 24b22 Ophiocordyceps superficialis, 24b23 Ophiocordyceps stylophora, 27a2 Dumontinia tuberosa, 38a3, 6 Hypomyces lactifluorum, 38e2 Hypocrea sulphurea, 41b8 Anthracobia melaloma, 50c1 Galiella rufa, 54a2 Hypocrea peltata, 54b2 Wolfina aurantiopsis, 56a2 Kretzschmaria deusta, 62a11 Wynnea americana Jonathan Frank: 4b3 Peziza ellipsospora, 5c6 Pachyphloeus austro-oregonensis, 8b3 Hydnobolites californicus, 8c9a&b Tuber quercicola Rosanne Healy: 4a4 Genea hispidula, 5c2 Pachyphloeus citrinus, 5c5 Pachyphloeus carneus Roger Heidt: 19a3 Underwoodia columnaris Bruce Horn: 8c10 Tuber lyonii, 25b7 Gelatinodiscus flavidus, 58a1 Annulohypoxylon thouarsianum Emily Johnson: 4b4 Hydnotrya cubispora, 15a2 Morchella cf. sceptriformis, 22b6 Xylaria cornudamae, 14b8 Hymenostilbe sphingum, 38a8 Nectriopsis tubariicola, 38a12 Hypomyces hyalinus, 38e1 Hypocrea latizonata, 50a2 Scorias spongiosa, 60a4 Gyromitra leucoxantha, 61a2 Sarcoscypha dudleyi, 62a9 Midotis irregularis, 66a1 Lasiosphaeria ovina, 71b2 Arachnopeziza aurelia, 72c9 Godronia urceolus Richard Kay: 17c6 Gyromitra caroliniana Sava Krstic: 31c5 Lachnum virgineum, 33c4 Pachycu-

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donia monticola, 43b8 Byssonectria fusispora, 47a9 Cheilymenia fimicola Jacques Landry: 41b4 Trichophaea abundans, 70a7 Scutellinia scutellata var. leucothecia Harold Larsen: 8c5 Tuber shearii, 30b6 Bryoscyphus marchantiae, 41b3 Trichophaea contradicta, 43a2 Peziza ostracoderma, 43b1 Pulvinula laeterubra, 43b5 Pyronema omphalodes, 47a5 Iodophanus carneus, 47b1 Ascobolus furfuraceus, 47b3 Byssonectria cartilaginea, 52a3 Miladina lecithina, 65a1 Chromelosporium fulvum, 73a9 Rhodoscypha ovilla, 73a11 Byssonectria tetraspora, 73b10 Pulvinula archeri, 73b12 Pulvinula laeterubra Renee Lebeuf: 3b4 Geopora pellita, 3b5 Geopora arenicola, 14b5 Morchella angusticeps, 14b7 Morchella elata, 14b12 Morchella punctipes, 15b2 Morchella americana, 15b3 Morchella americana var. lata, 15b4 Morchella ulmaria, 22b5 Xylaria longipes, 24b12 Isaria farinosa, 25b3 Cudoniella clavus, 29c1 Cudoniella acicularis, 30b4 Peziza waltersii, 30b10 Sarcoscypha occidentalis, 30b13 Chlorencoelia torta, 35b9 Microglossum olivaceum, 36a2 Geoglossum glabrum, 36a4 Geoglossum glutinosum, 36c3 Trichoglossum walteri, 38b2 Hypomyces tremellicola, 38d5 Protocrea pallida, 50c2 Sarcosoma globosum, 62b8 Peziza michelii, 62b14 Jafnea semitosta, 72b4 Phaeohelotium epiphyllum, 73a3 Neott iella vivida, 73a6 Sphaerosporella hinnulea, 73b6 Smardaea planchonis Lawrence Leonard: 65a3 Chromelosporium coerulescens hyphae, 72a5 Orbilia piloboloides David Lewis: 22b10 Xylaria liquidambaris, 24b15 Ophiocordyceps dipterigena, 38a2 Hypomyces macrosporus Hank Marshburn: 20c4 Helvella ephippium, 22b1 Xylaria polymorpha, 25b4 Mitrula paludosa, 25b8 Vibrissea truncorum, 26b1–2 Onygena equina, 30b3 Bisporella citrina, 30b5 Hymenoscyphus virgultorum, 30b12 Tatraea macrospora, 30b14 Chlorencoelia versiformis, 41b7 Anthracobia macrocystis, 43b4 Pyronema omphalodes, 43b4 Peziza proteana f. proteana, 43b5 Peziza praetervisa, 46b3 Peziza fimeti, 52a2 Pachyella clypeata, 58b2 Hypoxylon fragiforme, 62a10 Otidea rainierensis, 62b11 Humaria hemisphaerica, 62b15 Tarzetta bronca, 68b8 Hypocrea cf. strictipilosa, 68b13 Hypocrea cf. delicatula, 70a1 Scutellinia umbrorum, 70a3 Scutellinia erinaceus, 70a4

Scutellinia scutellata, 70a8 Scutellinia kerguelensis, 71b1 Arachnopeziza aurelia, 72a4 Orbilia luteorubella, 72b1 Chlorosplenium chlora Raymond McNeil: 6c1 Elaphomyces cf. asperulus, 17a3 Gyromitra esculenta var. alba, 19a1 Helvella lactea, 19b6 & 62b13 Jafnea fusicarpa, 21a2 Helvella cupuliformis, 21c3 Sowerbyella radiculata, 24a2 Elaphocordyceps longisegmentis, 3c2 Cudonia lutea, 36b3 Thuemenidium atropurpureum, 36c2 Trichoglossum velutipes, 43a1, 73b3 Peziza griseorosea, 43b3 Pulvinula carbonaria, 47a1 Pseudombrophila porcina, 55a3 Biscogniauxia mediterranea, 58b3 Thuemenella cubispora, 62b9 Peziza saccardoana, 68b6 Hypocrea cf. chromosperma, 68b9 Hypocrea cf. minutispora, 70a2 Scutellinia blumenaviensis, 70a6 Scutellinia setosa, 70b2 Scutellinia pennsylvanica, 73b2 Peziza azureoides, 73b8 Octospora humosa Ron Meyers: 17c4 Gyromitra brunnea Jim Murray: 3b3 Chorioactis geaster Andrew Parker: 21c3 Helvella atra, 21b3 Microstoma protractum, 43b2 Pulvinula convexella, 50c3 Entonaema liquescens, 62b1 Peziza arvernensis John Plischke III: 3b5 Geopora sepulta, 5c1 Pachyphloeus thysellii, 6a2 Elaphomyces cf. leveillei, 6a3 Elaphomyces cf. maculatus, 6a5 Elaphomyces cf. decipiens, 8b2 Choiromyces meandriformis, 8c7 Tuber lyonii, 14b4 Morchella angusticeps, 15b5 Morchella ulmaria, 17a6, 38e3 Sphaeronaemella helvellae, 22b2 Xylaria multiplex, 22b4 Xylaria cubensis (club) + Xylocoremium flabelliforme, 24b16 Ophiocordyceps variabilis, 24b20 Ophiocordyceps melolonthae, 24b21 Ophiocordyceps michiganensis, 30b9 Phaeohelotium epiphyllum, 33c6 Neolecta vitellina, 34a5 Tarzetta cupularis, 38a1 Hypomyces banningiae, 38a7 Hypomyces porphyreus, 38a9 Hypocrea avellanea, 38c2 Hypomyces microspermus, 38c4 Hypomyces melanocarpus, 38d1 Hypocrea americana, 38d2 Hypocrea lixii, 38d3 Hypocrea pulvinata, 38d4 Hypomyces aurantius, 38e4 Nectriopsis violacea, 41b2 Sphaerosporella brunnea, 41b6 Trichophaeopsis bicuspis, 41b9 Anthracobia maurilabra, 43b6 Pyronema domesticum, 43b9 Ascobolus carbonarius, 44b2 Peziza petersii, 49a2 Ascocoryne sarcoides, 52a1 Pachyella adnata, 54a1 Hypocrea peltata, 54b1 Wolfina aurantiopsis, 54a2 Biscogniauxia marginata, 57a1 Daldinia concentrica, 58a2 Annulohypoxylon multi-

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forme, 58b6 Hypoxylon howeanum, 62b4 Peziza domiciliana, 63c1 Diatrypella favaceae, 65a2 Chromelosporium coerulescens, 65b1 Taphrina robinsoniana, 66a3 Rosellinia quercina, 66a4 Lasiosphaeria lanuginosa, 68b3 Neonectria coccinea v. faginata, 68b10 Hypocrea pachybasioides, 70b1 Scutellinia crucipila, 70b3 Cheilymenia vitellina, 71b3 Arachnopeziza aurata, 72b5 Pithya cupressina, 72c3 Mollisia ventosa, 72c8 Ionomidotis fulvotingens, 73b1 Peziza gerardii Fred Rhoades: 22b3 Xylaria atropictor, 24b14 Ophiocordyceps myrmecophila, 25b10 Roseodiscus rhodoleucus, 30b18 Plectania nannfeldtii, 31c1 Lachnum bicolor, 31c4 Lachnum virgineum, 38a5 Hypomyces lateritius, 47a8 Cheilymenia stercorea, 56a3 Kretzschmaria deusta, 58b1 Hypoxylon fuscum, 61a3 Sarcoscypha coccinea, 62a1 Otidea alutacea, 66b3 Rhytisma punctatum, 72c2 Tapesia fusca William C. Roody: 11b3 Verpa conica, 17c3 Gyromitra korfii, 19b1 Helvella leucomelaena, 24a1 Elaphocordyceps capitata, 24a3 Elaphocordyceps ophioglossoides, 30b17 Urnula craterium, 30b19 Hymenoscyphus fr uctigenus, 33a1 Leotia lubrica, 33a2 Leotia viscosa, 44a2 Rhizina undulata, 50b6 Bulgaria inquinans, 55a5 Apiosporina morbosa, 62a12 Wynnea americana, 62b6 Peziza succosa, 62b12 Humaria hemisphaerica Gary Samuels: 38d8 Sporophagomyces chrysostomus, 68b11 Hypocrea schweinitzii, 68b12 Hypocrea viridescens Christian Schwartz: 27a3 Sclerotinia sulcata, 38d6 Hypomyces rosellus, 66a2 Rosellinia subiculata, 73b7 Smardaea planchonis Noah Siegel: 14a3 Morchella cf. anthracophila, 25b5 Mitrula elegans, 31a1 Lachnellula arida, 35b8 Microglossum fumosum, 38c5– 6 Hypomyces completus, 50c5 Camarops petersii, 55a4 Biscogniauxia atropunctata, 62a7 Otidea smithii Dianna Smith: 30b22 Chlorociboria aeruginascens, 35b1 Neolecta irregularis, 36a6 Geoglossum difforme, 38a11 Hypomyces luteovirens, 38e6 Hypomyces leotiicola, 46b2 Peziza vesiculosa, 49a1 Ascocoryne cylichnium, 62a4 Otidea unicisa Hugh and Sandi Smith: 14a2 Morchella exuberans, 15b9 Morchella rigida, 17b1 Gyromitra californica, 34a1 Sowerbyella rhenana, 35b2 Microglossum viride, 38e7 Hypomyces cervinigenus, 57a2 Daldinia grandis, 73a5 Chaetothiersia vernalis

Matt hew Smith: 4a3 Genea harknessii, 4a5 Genea arenaria, 4b2 Genabea cerebriformis, 8a2 Balsamia magnata, 8c3 Tuber canaliculatum, 8c8 Tuber candidum Walt Sturgeon: 29b2 Hypocrea alutacea, 50a1 Scorias spongiosa R. Greg Thorn: 3b2 Peziza ammophila, 63c3 Dermea cerasi James Trappe: 4a2 Gilkeya compacta Matt Trappe: 4b6 Hydnotrya cerebriformis, 7a2 Leucangium carthusianum, 7a3 Kalapuya brunnea, 8a3 Barssia oregonensis, 8c2 Tuber californicum, 8c4 Tuber sphaerosporum, 8c6 Tuber gibbosum Steve Trudell: 8c1 Tuber oregonense, 14a1 Morchella tomentosa, 15b8 Morchella rufobrunnea, 17a2 Gyromitra esculenta, 17c1 Gyromitra montana, 26b3 Onygena corvina, 29b1 Heyderia abietis, 31b3 Lachnellula suecica, 33c5 Hypocrea leucopus, 35b3 Microglossum rufum, 36b1 Geoglossum fallax, 61b2 Caloscypha fulgens, 61a7 Aleuria aurantia, 61a8 Pseudaleuria quinaultiana, 65b2 Taphrina occidentalis, 70a9 Scutellinia pilatii, 72b3 Pseudopithyella minuscula, 73b4 Pseudoplectania nigrella, 73b11 Pulvinula convexella Debbie Viess: 35b4 Spathularia cf. rufa, 62a5 Otidea cf. tuomikoskii Andrus Voitk: 25b1 Ascocoryne turficola, 25b2 Bryoglossum gracile, 36b2 Thuemenidium arenarium, 47b4 Byssonectria terrestris, 55a1 Biscogniauxia repanda Michael Wood: 21c2 Sowerbyella rhenana, 27a1 Sclerotinia veratri, 30b8 Kriegeria alutipes, 31b1 Lachnellula calyciformis, 68b1 Nectria cinnabarina, 73a8 Byssonectria terrestris

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Chapters 3 through 12 Harley Barnhart: Cookeina tricholoma, Galiella rufa, Helvella acetabulum, Otidea alutacea, Peziza domiciliana, Peziza praetervisa, Peziza proteana f. proteana, Urnula craterium, Claviceps purpurea, Lasiosphaeria spermoides, Bisporella citrina, Hymenoscyphus virgultorum, Geoglossum umbratile Kit Scates Barnhart: Geopora arenosa, Aleuria aurantia, Caloscypha fulgens, Gyromitra ancilis, Gyromitra infula, Gyromitra montana, Helvella latispora, Helvella maculata, Helvella solitaria, Peziza arvernensis, Peziza badia, Peziza echinospora, Plectania melastoma, Plectania nannfeldtii, Sarcosphaera

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coronaria, Scutellinia scutellata, Wynnella silvicola, Hypomyces lactifluorum George Barron: Beauveria bassiana + synnemata + mycelium with spores, Belonidium sulphureum, Roseodiscus subcarneus Alan and Arleen Bessette: Disciotis venosa, Gyromitra korfii, Helvella corium, Jafnea semitosta, Lamprospora crechqueraultii var. crechqueraultii, Microstoma floccosum, Morchella punctipes, Octospora humosa, Pachyella babingtonii, Peziza arenaria, Peziza varia, Peziza vesiculosa, Pseudoplectania nigrella, Rhodotarzetta rosea, Scutellinia erinaceus, Wynnea americana, Hypomyces chlorinigenus, Hypoxylon rubiginosum var. rubiginosum, Ophiocordyceps sphecocephala, Xylaria liquidambaris, Xylaria tentaculata, Ascocoryne cylichnium, Chlorosplenium chlora, Mollisia cinerea, Rhytisma americanum, Neolecta irregularis, Apiosporina morbosa Michael Beug: Cheilymenia fimicola, Gyromitra esculenta, Gyromitra melaleucoides, Helvella albella, Helvella crispa, Helvella fibrosa, Helvella vespertina, Morchella fr ustrata, Morchella elata, Morchella snyderi, Otidea onotica, Peziza proteana f. sparassoides, Plectania milleri, Pseudombrophila cervaria, Pseudoplectania vogesiaca, Rhizina undulata, Sarcoscypha coccinea, Trichophaea hemisphaerioides, Urnula padeniana, Verpa bohemica var. bohemica, Verpa conica, Cordyceps militaris, Helminthosphaeria clavariarum, Hypomyces aurantius, Coccomyces dentatus, Pachycudonia monticola, Trichoglossum hirsutum, Taphrina occidentalis Adolf and Oluna Ceska: Ciboria rufofusca Jules Cimon: Hypomyces ochraceus, Nectriopsis violacea, Crocicreas coronatum, Neobulgaria pura var. pura, Podophacidium xanthomelum Todd Elliott: Elaphomyces americanum, Elaphomyces fallax, Elaphomyces verruculosus, Imaia gigantea, Leucangium carthusianum, Pachyphloeus carneus, Tuber canaliculatum, Tuber shearii, Anthracobia melaloma, Morchella cf. americana, Neournula pouchetii, Plicaria endocarpoides, Wolfina aurantiopsis, Akanthomyces aculeatus, Cordyceps hesleri, Daldinia concentrica, Elaphocordyceps subsessilis, Hypomyces chrysospermus, Hypomyces lateritius, Ophiocordyceps melolonthae, Ophiocordyceps ravenelii, Ophiocordyceps variabilis, Gibellula leiopus, Dumontinia tuberosa

Jonathan Frank: Hydnobolites californicus Rosanne Healy: Genea hispidula, Pachyphloeus citrinus Bruce Horn: Annulohypoxylon thouarsianum var. thouarsianum, Entonaema liquescens, Gelatinodiscus flavidus Emily Johnson: Acervus epispartius, Cheilymenia theleboloides, Gyromitra brunnea, Helvella elastica, Helvella macropus, Helvella pezizoides, Sarcoscypha dudleyi, Annulohypoxylon cohaerens, Hypomyces cervinigenus, Hypomyces hyalinus, Hypomyces luteovirens, Hypomyces tremellicola, Nectriopsis tubariicola, Xylaria polymorpha, Bulgaria inquinans, Chlorociboria aeruginascens ssp. aeruginascens, Godronia urceolus, Rutstroemia bulgarioides Jacques Landry: Gyromitra sphaerospora Harold Larsen: Morchella cf. prava, Rhodoscypha ovilla Renee Lebeuf: Midotis irregularis, Neott iella vivida, Peziza waltersii, Pseudombrophila merdaria, Sarcoscypha occidentalis, Sarcosoma globosum, Hypocrea alutacea, Mitrula lunulatospora, Geoglossum difforme, Microglossum olivaceum David Lewis: Ophiocordyceps dipterigena Hank Marshburn: Humaria hemisphaerica, Peziza fimeti, Sowerbyella rhenana, Hypocrea pulvinata, Cudoniella acicularis, Cudoniella clavus, Holwaya mucida, Spathularia flavida, Orbilia luteorubella Raymond McNeil: Genea arenaria, Peziza ostracoderma, Pulvinula carbonaria, Sowerbyella radiculata, Biscogniauxia mediterranea, Daldinia childiae, Encoelia furfuracea, Geoglossum simile, Thuemenidium atropurpureum Ron Meyers: Gyromitra caroliniana Jim Murray: Chorioactis geaster Andrew Parker: Microstoma protractum, Pulvinula convexella John Plischke III: Choiromyces meandriformis, Anthracobia maurilabra, Ascobolus carbonarius, Morchella diminutiva, Peziza gerardii, Peziza petersii, Peziza succosa, Pithya cupressina, Plicaria carbonaria, Scutellinia crucipila, Sphaerosporella brunnea, Wynnea sparassoides, Hypocrea latizonata, Hypocrea pachybasioides, Hypocrea peltata, Hypomyces porphyreus, Hypoxylon fuscum, Ophiocordyceps superficialis, Thuemenella cubispora, Xylaria cubensis with Xylocoremium flabelliforme, Arachnopeziza aurelia, Ascotremella faginea, Catinella olivacea, Ionomidotis fulvotingens, Lachnum virgineum, Leotia viscosa

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Fred Rhoades: Nectria cinnabarina William C. Roody: Helvella atra, Morchella angusticeps, Peziza phyllogena, Peziza sublilacina, Sarcoscypha austriaca, Tarzetta catinus, Hypocrea sulphurea, Kretzschmaria deusta, Xylaria hypoxylon, Chlorencoelia versiformis, Mitrula elegans, Vibrissea truncorum, Microglossum rufum Gary Samuels: Hypocrea schweinitzii, Hypocrea strictipilosa, Hypocrea viridescens Christian Schwartz: Camarops petersii, Rosellinia subiculata Noah Siegel: Otidea smithii, Biscogniauxia atropunctata, Elaphocordyceps capitata, Elaphocordyceps ophioglossoides, Sclerotinia sulcata Dianna Smith: Diatrype stigma Hugh and Sandi Smith: Geopyxis carbonaria, Melastiza chateri, Morchella rufobrunnea, Daldinia grandis Matt hew Smith: Balsamia magnata, Tuber lyonii Roger Smith: Dermea cerasi, Neocudoniella radicella Walt Sturgeon: Scorias spongiosa

R. Greg Thorn: Peziza ammophila, Underwoodia columnaris James Trappe: Gilkeya compacta, Hydnotrya cubispora Matt Trappe: Balsamia nigrans, Genabea cerebriformis, Genea harknessii, Hydnotrya tulasnei, Tuber californicum, Tuber gibbosum, Helvella cupuliformis Steve Trudell: Geopora cooperi, Hydnotrya variiformis, Gyromitra californica, Helvella compressa, Helvella leucomelaena, Morchella tomentosa, Pseudaleuria quinaultiana, Tricharina gilva, Cudonia circinans, Dasyscyphella nivea, Heyderia abietis, Hymenoscyphus fr uctigenus, Lachnellula agassizii, Lachnellula arida, Lachnum bicolor, Leotia lubrica, Phaeohelotium epiphyllum var. epiphyllum, Tatraea macrospora, Onygena corvina Debbie Viess: Chaetothiersia vernalis Andrus Voitk: Byssonectria terrestris, Ascocoryne turficola, Bryoglossum gracile Michael Wood: Ascobolus stercorarius, Pyronema omphalodes

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Index to Common Names

acorn cup fungus, 385 adder’s tongue, 298 alder tongues, 442 amanita mold, 320 anemone cup, 378 beech aphid poop-eater, 439 beech hypoxylon, 330 beefsteak morel, 149 beetle cordyceps, 338 berry truffle, 107 black elfi n saddle, 170 black foot morel, 193 black helvella, 157 black jelly drops, 364 black knot of cherry, 438 black morel, 120, 121 black scurfy fairy cup, 159 bladder cup, 223 blueberry cup, 378 blue-stain fungus, 367 blue-staining cup, 133 bolete eater, 319 bolete mold, 319 bonfi re cauliflower, 218 brain mushroom, 149 brown-haired white cup, 172 bull-nose false morel, 151 burn site ochre cup, 128 burn site shield cup, 129 candle snuff fungus, 349 carbon balls, 293 carbon cushion, 332 Carolina false morel, 148 catkin cup, 370 coal fungus, 293

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common alder tongue, 442 common black earth tongue, 422 common brown cup, 215 common earth tongue, 420 common eastern black morel, 181 common elfi n saddle, 162 common morel, 179 common western black morel, 191 compressed elfi n saddle, 158 cone fl ickers, 350 conifer false morel, 149 coral spot fungus, 277, 334 cramp balls, 291, 293 crown fungus, 250 crustlike cup, 242 dark elfi n saddle, 157 dead man’s fi ngers, 351 deer truffle, 82 devil’s cigar, 139 devil’s urn, 264 dog’s nose fungus, 287 domicile cup fungus, 208 donkey ears, 200 early morel, 267 eastern half-free morel, 189 elephant ear, 146 elf cup, 258 elfi n cup, 155 ergot, 275, 288 eyelash cup, 253 eyelash dung cup, 135 eyelash fungus, 253 fairy sparklers, 352 fairy tub, 206

fall Oregon white truffle, 111–112 false morel, 149 false orange peel, 256 false sparassis, 218 fluted black helvella, 170 fluted brown elfi n saddle, 167 fluted-stalked fungus, 263 fluted white helvella, 160 fuzzy foot, 193 fuzzy-foot morel, 193 fuzzy truffle, 91 gabled false morel, 146 ghosts, 193 giant gel cup, 265 golden hypomyces, 319 golden-thread elaphocordyceps, 298 gray morel, 193 green-capped jelly baby, 396 green earth tongue, 425 green-headed jelly club, 396 green lobster, 324 greens, 194 green-stain fungus, 367 hairy black cup, 236 hairy earth tongue, 429 hairy rubber cup, 142 half-free morel, 189 hare’s ear, 200 head-like elaphocordyceps, 296 irregular earth tongue, 432 jelly babies, 394 jellylike black urn, 227

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King Alfred’s cakes, 291, 293 landscape morel, 184 lemon drops, 362 lobster fungus, 276, 321 lobster mushroom, 321 long-stalked gray cup, 166 lorchel, 149 magnolia cone xylaria, 350 mini peach pig’s ears, 120, 145 moose antlers, 270 mountain blond morel, 185 mulch morel, 184 natural black morel, 192, 194 ochre jelly club, 394 olive earth tongue, 425 Olympic pig’s ears, 145 orange earth tongue, 427 orange eyelash cup, 252 orange otidea, 200 orange peel, 126 orange pig’s ears, 145 Oregon black truffle, 104 Oregon brown truffle, 104 peat mould, 212 pecan truffle, 113 pig’s ears, 144, 207 pig truffle, 78 pine fi re cushion, 242 pink burn cup, 244 pink crown, 250 pinks, 194

pixie cup, 143 purple fairy cup, 216 purple jelly drops, 357 rabbit ears, 270 recurved cup, 221 red-brown blushing morel, 190 red cushion hypoxylon, 330 red false morel, 148 rhinoceros-beetle cordyceps, 338 ribbed elfi n saucer, 169 ribbed-stalked cup, 155 rooster’s comb, 146 rose goblet, 196 round-headed elaphocordyceps, 296 round-spored false morel, 154 round-spored gyromitra, 154 Russula mold, 324 saddle-shaped false morel, 150 sand-loving cup, 204 sand morel, 188 scarlet cup, 245, 246 scarlet elf cup, 246 scurfy alder cup, 379 shaggy scarlet cup, 176 smooth-stalked helvella, 162 smooth thimble-cap, 268 snowbank orange peel fungus, 133 snow mushroom, 153 split skin carbon cushion, 287 spreading brown cup fungus, 221 spring Oregon white truffle, 107 spruce needle beacon, 383 stalked cauliflower fungus, 271 stalked hairy fairy cup, 393

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stalked orange peel fungus, 256 stalked scarlet cup, 248 starving man’s licorice, 265 swamp beacon, 397 tar spot of maple, 406 toothed tar spot, 371 trooping cordyceps, 290 tulip cup, 250 tulip morel, 182 umbrella false morel, 147 veined cup, 141 velvet stalked fairy fan, 412 velvety black earth tongue, 429 velvety fairy fan, 412 violet fairy cup, 412 viscid black earth tongue, 420 walnut, 153 water club, 414 western black morel, 191, 192 western blond, 185, 186, 194 western blond morel, 185, 186, 194 western urnula, 196 white elfi n saddle, 160 white morel, 120, 182 witches’-brooms, 441, 442 wrinkled thimble cap, 267 yellow cudonia, 373 yellow fairy cups, 362 yellow morel, 120, 121, 179 yellow pig’s ears, 145 yellow sap cup, 220

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Index to Scientific Names

Note: Boldface type indicates the pages where descriptions appear. Acervus epispartius, 68, 123, 125, 126 Acetabula sulcata, 155 Aconitum, 410 Acremonium fingicola, 336 tulasnei, 323 Agaricus, 328 Akanthomyces aculeatus, 31, 279–280 aranearum, 280 sphingum, 280 Aleuria abietina, 200 amplissima, 206 aurantia, xii, 7, 11, 57, 123, 125, 126, 232, 256 proteana, 217 proteana var. sparassoides, 218 rhenana, 256 sylvestris, 206 umbrina, 209 violacea, 219 Alnus incana, 442 rubra, 442 Amanita, 276, 320, 321 rubescens, 320 Anemone, 355 nemorosa, 378 ranunculoides, 378 trifolia, 378 Annulata, 278

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Annulohypoxylon, 278, 282, 330 annulatum, 281, 282 cohaerens, 55, 281 multiforme var. multiforme, 55, 281 thouarsianum var. macrosporum, 282 thouarsianum var. thouarsianum, 55, 282–283 truncatum, 281, 282 Anthopeziza floccosa, 176 Anthracobia, 122, 240 macrocystis, 46, 128 maurilabra, 46, 127 melaloma, 46, 128, 238 Apiocrea hyalina, 320 Apiosporina morbosa, 54, 437, 438 Arachnopeziza arctostaphyli, 356 aurata, 65, 356 aurelia, 65, 356, 360 delicatula, 356 Arthrobotrys, 3, 434 Ascobolaceae, 118 Ascobolus, 4, 118 atrofuscus, 129 carbonarius, 48, 129, 231 crenulatus, 130 furfuraceus, 50, 130 geophilus, 129 immersus, 6 lignatilis, 130

michaudii, 130 minutus, 2 stercorarius, 50, 130–131, 136, 219 Ascocoryne, 355 cylichnium, 50, 357 sarcoides, 50, 357 turficola, 32, 358 Ascocorynium irregulare, 432 Ascophanus, 240 Ascotremella faginea, 52, 355, 359 Aspergillus, 416 Balsamia, 71, 73, 119 alba, 75 magnata, 19, 75– 76 nigrans, 19, 77 nigrens, 77 vulgaris, 75 Barlaea constellatio, 239 Barssia, 71, 73, 119 oregonensis, 19, 76 Beauveria, 277 bassiana, 31, 284 Belonidium, 355, 362 sulphureum, 65, 355, 360–361 Belonopsis graminea, 68, 400 Berberis, 277 Betula nana, 442 pubescens, 442

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Bionectria, 277 Bionectriaceae, 274, 277 Biscogniauxia, 278, 286 atropunctata, 54, 285 marginata, 53, 286 mediterranea, 53, 286 repanda, 53, 286 Bisporella, 225, 355, 386, 404 citrina, 35, 362 claroflava, 362 subpallida, 35, 362 sulphurina, 35, 362 Boletus, 324 pallidus, 318 Botrytis, 212, 410 Brunnipila calyculiformis, 37, 390 Bryoglossum, 355 gracile, 32, 355, 363 rehmii, 363 Bryoscyphus marchantiae, 35, 386 Bulgaria, 354 globosa, 249 inquinans, 51, 354, 364 melastoma, 227 mexicana, 266 pura, 401 rufa, 142 Bulgariaceae, 354 Bulgariella pulla, 50, 355, 364 Byssonectria, 123 cartilaginea, 50 fusispora, 48, 132 luteovirens, 324 seaveri, 132 terrestris, 11, 50, 68, 132 tetraspora, 68, 197 Caloscypha fulgens, vi, 57, 118, 133 Caloscyphaceae, 118 Calycella alba, 372 Calycina, 408 Camarops petersii, 51, 52, 287

Capnodiaceae, 437 Capnodiales, 437, 439 Carex, 410 Caroliniana, 119, 146 Catinella olivacea, 67, 365, 437 Cazia, 71, 73 flexiascus, 94 quercicola, 94 Cenangium, 382 furfuraceum, 379 Cercophora, 8 Chaetothiersia vernalis, 68, 122, 134 Cheilymenia, 123, 130, 234, 240, 252, 259 calvescens, 251 coprinaria, 135 crucipila, 251 fimicola, 49, 135–136, 138 raripila, 135 stercorea, 49, 135, 138, 251 theleboloides, 49, 135, 137–138 vitellina, 64, 138 Chlorencoelia, 355 torta, 36, 355, 366 versiformis, 36, 67, 178, 355, 366 Chlorociboria, 355 aeruginascens, 36, 367 aeruginascens ssp. aeruginascens, 367 aeruginosa, 36, 367 bulgarioides, 409 versiformis, 366 Chloroscypha, 355 Chlorosplenium aeruginascens, 367 chlora, 66, 354, 368 olivaceum, 368 versiforme, 366 Choiromyces, 71, 73 alveolatus, 20, 78 meandriformis, 20, 78 venosus, 78 Choriactidaceae, 118, 266 Chorioactis geaster, 1, 16, 118, 139

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Chromelosporium coerulescens, 60, 212 fulvum, 60, 212 ollare, 212 Chromocreopsis cubispora, 346 Ciboria, 370, 413 amentacea, 370 bulgarioides, 409 caucus, 370 fr uctigena, 385 peckiana, 413 rufofusca, 67, 355, 369 Ciliaria brunnea, 257 Cistella, 376 Cladobotryum varium, 315–316 verticillatum, 325 Clavaria, 277, 424 Claviceps purpurea, 60, 275–276, 288 Clavicipitaceae, 274–275 Clavicipitales, 274 Clavulina, 301 cinerea, 301 coralloides, 301 Coccomyces, 354, 371 coronatus, 371 dentatus, 8, 62, 371 Coleoptera, 341, 344 Cookeina, 124 sulcipes, 28, 140 tetraspora, 140 tricholoma, 28, 140 Coprobia, 239 Coprotus, 239 Cordyceps, 277 acicularis, 341 alutacea, 302 arachnophila, 347 bassiana, 284 canadensis, 290 capitata, 296 capitata var. canadensis, 296 cardinalis, 290 dipteragena, 337

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facis, 299 hesleri, 30, 289 ithacanensis, 344 melolonthae, 338 militaris, 30, 275, 290, 338 muscicola, 337 olivascens, 30, 290 ophioglossoides, 298 pseudomilitaris, 275 ravenelii, 339 sphecocephala, 340 sphingum, 280 superficialis, 341 thaxteri, 280 tuberculata, 31, 279–280 viperina, 344 washingtonensis, 30, 290 Cordycipitaceae, 274–275 Coremium gracile, 280 Coryne atrovirens, 38, 394 dubia, 355, 357 turficola, 358 Costapeda crispa, 160 Crepidotus, 322, 327 Crinula caliciiformis, 36, 384 Crocicreas, 372 coronatum, 35, 372 Cudonia, 354 circinans, 39, 373, 394, 403 grisea, 373, 403 lutea, 39, 373 monticola, 403 Cudoniaceae, 354 Cudoniella, 355, 375 acicularis, 34, 374, 375 aquatica, 375 borealis, 374 clavus, 32, 363, 374, 375, 402 clavus var. grandis, 375 Cyathicula coronata, 372 cyathoidea, 386 Cyathipodia macropus, 166

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Cyathus striatus, 276, 303 Cytarriales, 354 Daldinia, 278, 282, 330 childiae, 54, 291–292, 294 clavata, 292 concentrica, 54, 287, 292, 293 concentrica f. californica, 294 concentrica f. confluens, 292 concentrica f. intermedia, 292 concentrica var. minuta, 292 gelatinosa, 292, 294 grandis, 54, 292, 293, 294 lloydii, 292 loculata, 292 malleola, 282 tuberosa, 293 Daleomyces phillipsii, 257 Dasyscypha bicolor, 392 Dasyscyphella montana, 376 nivea, 38, 376, 392, 393 Dasyscyphus, 390 agassizii, 389 arida, 390 niveus, 376 pini, 390 sulphureus, 360 virgineus, 393 Delphinium, 410 Dermataceae, 354 Dermea cerasi, 60, 354, 377 Diatrypaceae, 278 Diatrypales, 278 Diatrype, 278 atropunctata, 285 stigma, 54, 295 undulata, 295 Diatrypella, 278 favacea, 60, 295 verruciformis, 295 Dibotryon morbosum, 438 Dichobotrys, 262

Diptera, 337, 343 Discina, 119, 151 abietina, 200 ancilis, 144 apiculata, 145 larryi, 145 leucoxantha, 145 olympiana, 145 perlata, 145 Discinaceae, 71, 118 Discinella boudieri, 69, 215, 355 Disciotis, 120 venosa, 56, 120, 141, 145, 474 Discocainea, 382 Donadinia, 229 nigrella, 229 Dothidiomycetes, 437 Dumontinia tuberosa, 34, 355, 378, 410 Durandiomyces phillipsii, 257 Elaphocordyceps, 9, 296 capitata, 30, 296–297, 298 fracta, 296 japonica, 296 longisegmentis, 30, 296 ophioglossoides, 30, 275, 296, 298, 339 paradoxa, 298 subsessilis, 30, 299 tenuispora, 296 Elaphomyces, 72, 73, 275, 296, 416 americanum, 19, 79, 82 anthracinus, 18, 81 appalachiensis, 81, 296 asperulus, 19, 82 cyanosporus, 81 decipiens, 18, 80–81 fallax, 80–81 granulatus, 19, 72, 82, 296 leveillei, 18, 81 loebae, 18, 81 macrosporus, 18, 81 maculatus, 18, 81 muricatus, 19, 79, 416 reticulatus, 79

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Elaphomyces (continued) subviscidus, 19, 82 verruculosus, 19, 82–83 Elaphomycetaceae, 72, 416 Elata, 120 Encoelia, 382 furfuracea, 60, 355, 379, 382 Entoloma, 322 strigosissimum, 326 Entonaema, 278 liquescens, 52, 300 Equisetum, 386, 408 Erinellina nylanderi, 360 Erisyphales, 354 Erysiphe, 2 Esculenta, 120, 188 Eurotiales, 72, 416 Eurotiomycetes, 416 Exidia, 312 nucleata, 276 Fischerula, 71, 73 subcaulis, 71 Fomitopsis pinicola, 316 Fuligo septica, 277, 336 Galactinia badia, 207 praetervisa, 216 proteana, 217 sarrazini, 213 succosa, 220 vesiculosa, 223 violacea, 219 Galiella, 124 rufa, 52, 124, 142, 269 Geastrum, 139 Gelatinodiscus flavidus, 33, 226, 355, 380 Genabea, 71, 74 cerebriformis, 17, 84, 91, 93, 100 Genea, 71, 74, 84, 93 anthracina, 86 arenaria, 17, 85, 88 balsleyi, 86

bihymeniata, 86 brachytheca, 86, 88 cerebriformis, 84 compacta, 85, 93 echinospora, 85 gardneri, 86 harknessii, 17, 86–87 hispidula, 17, 88 intermedia, 93 macrosiphon, 85 thaxteri, 85 Geoglossaceae, 355, 419 Geoglossum, 9, 355, 419, 429 affine, 420 alveolatum, 40, 423 arenarium, 428 atropurpureum, 428 cohaerens, 423 cookeanum, 419 difforme, 41, 419, 420–421, 424 fallax, 41, 419, 423 glabrum, 40, 419, 423, 424 glabrum var. simile, 423 glutinosum, 41, 419, 420 intermedium, 419, 423 irregulare, 432 littorale, 423 luteum, 427 nigritum, 424 olivaceum, 425 peckianum, 40, 420 pumilum, 423 pygmaeum, 423, 424 simile, 41, 419, 422–423, 424 umbratile, 41, 419, 424 viride, 426 Geopora, 72, 74, 91, 134 annulata, 91 arenicola, 16, 89 arenosa, 47, 89– 90, 91 clausa, 89– 90 cooperi, 17, 91– 92, 100 harknessii, 91 magnifica, 91 pellita, 16, 89 sepulta, 16, 59, 89, 172, 259 Geopyxis, 122, 258 ammophila, 204

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Beug-final.indb 480

carbonaria, 39, 47, 143 craterium, 264 floccosa, 176 occidentalis, 248 vulcanalis, 39, 143, 258 Gibellula leiopus, 31, 347 pleiopus, 347 pulchra, 347 Gilkeya, 72, 74 compacta, 17, 84, 93 Godronia, 381–382 cassandrae, 381 urceolus, 67, 355, 381 Gremmeniella, 382 Guepiniopsis, 380 Gymnomitrula abietis, 383 gracilis, 363 Gyromitra, 119, 149 ambigua, viii, 25, 119, 149, 150 ancilis, 6, 48, 56, 119, 120, 141, 144–145, 152 brunnea, 25, 119, 146, 148 californica, 25, 147, 154 caroliniana, 25, 119, 146, 148 esculenta, 24, 119, 149 esculenta var. alba, 24, 149 fastigiata, 146 fluctuans, 144 gigas, 119, 151, 153 infula, 25, 119, 149, 150 korfii, 25, 119, 151, 153 larryi, 145 leucoxantha, 56, 119, 145, 152 macrospora, 144 melaleucoides, 56, 119, 145, 152 montana, 25, 119, 151, 153 mwb060911, 56, 120, 145 olympiana, 56, 119, 145, 152 parma, 119 recurva, 152 sphaerospora, 25, 147, 154 warnei, 144 Helminthosphaeria clavariarum, 45, 277, 301 Helminthosphariaceae, 277

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Helotiaceae, 354, 380 Helotiales, 354, 355 Helotium chlora, 368 citrinum, 362 clavus, 375 destructor, 407 fr uctigenum, 385 Helvella, 119, 277, 317 acetabulum, 27, 155 aestivalis, 165 albella, 27, 156, 157, 158, 162, 164 ambigua, 150 arctica, 159 atra, 27, 156, 157, 159, 168 brevis, 166 californica, 147 caroliniana, 148 chinensis, 163 cochleata, 207 columnaris, 263 compressa, 27, 158, 164 confusa, 165 connivens, 164 corium, 28, 159, 229 costifera, 27, 155, 169 crassitunicata, 165 crispa, 26, 160, 167 cupuliformis, 28, 161, 163, 166, 169 dissingii, 163 dryophila, 26, 159, 170–171 elastica, 27, 156, 158, 162, 164 ephippium, 27, 156, 168 esculenta, 149 fibrosa, 28, 161, 163, 166, 169 griseoalba, 155 hyperborea, 155, 169 infula, 150 lactea, 26, 160 lacunosa, 170, 171 latispora, 27, 156, 158, 164 leporina, 200 leucomelaena, 27, 39, 165, 169 lubrica, 394 macropus, 28, 163, 166 macropus var. brevis, 166 maculata, 26, 160, 167, 169 murina, 156

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nigrella, 229 nigricans var. atra, 157 pallidula, 163 palustris, 171 pedunculata, 165 pezizoides, 27, 156, 157, 159, 168 queletii, 169 rivularis, 161 robusta, 155 silvicola, 272 solitaria, 27, 155, 159, 161, 163, 169 sphaerospora, 154 stevensii, 164 subglabra, 157 sulcata, 26, 171 underwoodii, 146 unicolor, 27, 169 verruculosa, 159 vespertina, 26, 159, 167, 170–171 villosa, 163 Helvellaceae, 71, 118, 119 Hemiphacidiaceae, 355 Hemisphaeria concentrica, 293 tuberosa, 293 Heyderia, 355 abietis, 34, 355, 383 Hirsutella stylophora, 344 Holwaya gigantea, 384 leptosperma, 384 mucida, 36, 354, 384 Humaria, 123, 134 fimeti, 210 gerardii, 211 hemisphaerica, 59, 90, 172, 261 hemisphaerioides, 261 humosa, 197 maurilabra, 127 miniata, 175 Humarina aggregata, 132 gerardii, 211 Hyalorbilia, 434 inflatula, 435 Hyaloscypha, 355, 360, 362 albohyalina, 67, 360

albohyalina var. albohyalina, 360 albohyalina var. spiralis, 360 albohyalina var. tigillari, 360 aureliella, 360 britannica, 65, 355, 360 candida, 360 Hyaloscyphaceae, 354, 355 Hydnobolites, 71, 73, 74 californicus, 20, 94– 95 tulasnei, 98 Hydnocystis, 72 compacta, 93 Hydnotrya, 71, 74, 91, 100, 119 carnea, 98 cerebriformis, 17, 98, 100 cubispora, 17, 96– 97 ellipsospora, 98 inordinata, 96 michaelis, 96 subnix, 98 tulasnei, 96, 98– 99 variiformis, 17, 100–101 variiformis var. pallida, 100 variiformis var. variiformis, 100 Hydnotryopsis, 71, 74, 100 Hydropisphaera, 277 peziza, 62, 277, 334 Hymenoscyphus, 225, 355, 362, 370, 375, 386, 404, 408, 413 albidus, 386 calyculus, 386 coronatus, 372 epiphyllus, 226, 404 fr uctigenus, 36, 385, 386 rhodoleucus, 386 subcarneus, 407, 408 tuberosus, 378 vernus, 386 virgultorum, 35, 386–387 Hymenostilbe dipterigena, 337 sphecocephala, 340 sphingum, 30, 280 Hypocrea, 276, 306, 316, 328 alutacea, 34, 276, 302 americana, 44, 316 atroviridis, 314 aurantius, 316

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Hypocrea (continued) aureoviridis f. macrospora, 311 avellanea, 43, 276, 327–328 ceracea, 311 chlorospora, 63, 311 chromosperma, 63, 311 citrina, 325 citrina f. fungicola, 307 delicatula, 63, 316, 328 farinosa, 312 fungicola, 307 gelatinosa, 311 jecorina, 308 latizonata, 45, 276, 303 leucopus, 39, 302 lixii, 44, 308 luteovirens, 324 minutispora, 63, 314 pachybasioides, 63, 304 pallida, 316 peltata, 53, 305–306 petersenii, 314 pulvinata, 44, 307 rogersonii, 314 rufa, 314 schweinitzii, 63, 304, 308–309 sinuosa, 63, 311 strictipilosa, 63, 310–311 stromatica, 328 sulphurea, 45, 276, 312, 325 thelephoricola, 311 viridescens, 63, 313–314 Hypocreaceae, 274, 276 Hypocreales, 274–276 Hypomyces, 276, 316, 322, 328, 336 armeniacus, 325 aurantius, 44, 307, 312, 315–316 banningiae, 42, 322 banningii, 322 boletiphagus, 318 cervinigenus, 45, 276–277, 317, 326 chlorinigenus, 43, 318 chrysospermus, 43, 319 chrysostomus, 326 completus, 44, 319 hyalinus, 42, 320 insignis, 321 lactifluorum, 42, 276, 277, 321–322

lateritius, 42, 323 leotiicola, 45, 394 lithuanicus, 323 luteovirens, 43, 324 macrosporus, 42, 322 melanocarpus, 44, 319 microspermus, 43, 319 ochraceus, 42, 277, 325 porphyreus, 43, 326 purpureus, 321 rosellus, 44, 316 spadiceus, 323 stephanomatis, 172 subaurantius, 316 thiryanus, 323 torminosus, 323 tremellicola, 42, 327–328 tulasneanus, 319, 324 viridis, 324 Hypoxylon, 278, 286 amaniense, 282 annulatum, 282 atropunctatum, 285 coccineum, 330 cohaerens, 281 cohaerens var. microsporum, 281 ferrugineum, 55, 331 fragiforme, 55, 330, 331 fuscum, 55, 329–330 howeanum, 55, 330 howeianum, 330 multiforme var. australe, 330 occidentale, 282 petersii, 287 rubiginosum, 55, 331 rubiginosum var. ferrugineum, 331 rubiginosum var. rubiginosum, 331 thouarsianum var. gilletianum, 282 Imaia, 71, 73 gigantea, 19, 102–103 Inermisia fusispora, 132 Iodophanus, 121, 130 carneus, 49, 121, 234, 240

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Beug-final.indb 482

Ionomidotis fulvotingens, 67, 355, 388 irregularis, 178 Isaria, 277 farinosa, 31, 280 shiotae, 284 sphingum, 280 Jafnea, 123 fusicarpa, 27, 59, 173 semitosta, 59, 173 Kalapuya, 71, 73 brunnea, 19, 104 Kretzschmaria, 278 deusta, 54, 332 Kriegeria alutipes, 35, 380 Lachnea ascoboloides, 138 hemisphaerica, 172 Lachnella, 390 agassizii, 389 arida, 390 bicolor, 392 nivea, 376 rufo-olivacea, 390 sulphurea, 360 virginea, 393 Lachnellula, 355, 362, 376, 389, 392 agassizii, 389 arida, 37, 390–391 calyciformis, 37, 389 flavovirens, 37, 390 gallica, 37, 389 occidentalis, 389 suecica, 37, 389 willkommii, 389 Lachnum, 12, 355, 362, 376, 392 bicolor, 38, 392 calyculiforme, 390 niveum, 376 pudibundum, 38, 393 virgineum, 38, 393 Lactarius, 276, 277, 322 camphoratus, 323 piperatus, 322

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pubescens, 323 torminosus, 323 Laetiporus sulphureus, 307 Lamprospora, 123 carbonaria, 238 crechqueraultii, 69, 174 crechqueraultii var. crechqueraultii, 174 crechqueraultii var. macrantha, 174 macrantha, 174 pyrophila, 239 Lanzia, 355 luteovirescens, 35, 355, 409 Lasiobolus, 123, 130 cuniculi, 49, 130 intermedius, 12 Lasiosphaeria, 277 lanuginosa, 61, 333 ovina, 61, 333 spermoides, 61, 333 Lasiosphaeriaceae, 277 Lecanicillium, 277 Leotia, 355, 375 albiceps, 402 circinans, 373 elegans, 397 lubrica, 38, 355, 373, 394–395, 403 truncorum, 414 viscosa, 38, 355, 394, 396 Leotiaceae, 355, 375 Leotiomycetes, 354–355, 419, 428, 437 Lepidoptera, 337, 338, 342 Leptoglossum alabamense, 426 fumosum, 427 luteum, 427 Leptonia strigosissima, 326 Leptopodia albella, 156 elastica, 162 Leucangium, 71, 73 carthusianum, 19, 102, 104 Leucomelaenae, 159

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Leucoscypha ovilla, 243 vivida, 195 Liquidambar styraciflua, 350 Litorella, 423 Marasmius subnudus, 276, 327 Marcelleina atroviolacea, 211, 236 persoonii, 211 Melaleucoides, 119, 152 Melastiza, 123 charteri, 175 chateri, 68, 175 cornubiensis, 175 miniata, 175 Mertensia, 410 Microglossum, 355, 419, 428 contortum, 425 elegans, 397 fumosum, 40, 425, 427 longisporum, 427 olivaceisquamulosum, 426 olivaceum, 40, 425–426 rufum, 40, 412, 427 viride, 40, 394, 425–426 Microhilum, 277 Microsphaera, 5 Microstoma, 124 floccosum, 38, 124, 176, 248 floccosa, 176 protracta, 177 protractum, 28, 124, 177, 248 Midotis, 119, 178 irregularis, 58, 178, 200 versiformis, 366 Miladina, 122 lecithina, 52, 203 Mitophora semilibera, 189 Mitrula, 355, 363 abietis, 383 borealis, 363, 397 elegans, 33, 397 gracilis, 363 irregularis, 432

lunulatospora, 33, 398 paludosa, 33, 397 Mollisia, 355, 399 cinerea, 67, 355, 399–400 discolor, 67, 400 ventosa, 67, 400 Monilinia, 370, 413 vaccinii-corymbosi, 378 Moniliophthora perniciosa, 328 Morchella, 120 americana, 23, 179–180, 182, 188, 190 americana var. lata, 23, 180 angusticeps, 22, 181, 192 anthracophila, 22, 194 brunnea, 22, 192 californica, 180 capitata, 194 carbonaria, 194 conica, 192 costata, 184 crassipes, 180 cryptica, 180 deliciosa, 182, 188 diminutiva, 23, 182–183 dunensis, 188 elata, 22, 181, 184, 186, 192 esculenta, 180, 190 esculenta var. crassipes, 180 esculentoides, 180 eximia, 194 exuberans, 22, 194 fr ustrata, 23, 185–186 guatemalensis, 190 herediana, 190 hotsonii, 184 importuna, 184 Mel-8, 192 populina, 180 populina var. lata, 180 populiphila, 23, 189, 267, 268 prava, 24, 180, 187–188 pulchella, 181 punctipes, 23, 189, 267, 268 rigida, 24, 180 rufobrunnea, 24, 120, 180, 190 sceptriformis, 23, 182

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Morchella (continued) semilibera, 189 septentrionalis, 181 septimelata, 194 sextelata, 194 snyderi, 22, 186, 191–192 tomentosa, 22, 193–194 ulmaria, 24, 180, 188 virginiana, 182 vulgaris, 188 Morchellaceae, 71, 120 Mycogone cervina, 317 Mycolachnea hemisphaerica, 172 Mycosphaerella, 437 Myrmecocystis candida, 84 cerebriformis, 84 Myxotrichaceae, 354 Nectria, 277, 327, 335 cinnabarina, 62, 277, 334 coccinea, 334 peziza, 334 purpurea, 334 Nectriaceae, 274, 277 Nectriopsis tremellicola, 327 tubariicola, 43, 335 violacea, 45, 277, 336 Neobulgaria, 355 pura var. pura, 52, 401 Neocudoniella albiceps, 34, 375, 402 radicella, 33, 374, 402 Neolecta, 431 irregularis, 39, 40, 431, 432 vitellina, 39, 432 Neolectomycetes, 431 Neonectria coccinea var. faginata, 63, 277, 334 fuckeliana, 63, 334 Neott iella, 122, 195, 240 ascoboloides, 138 hetieri, 195 ovilla, 243 rutilans, 68, 195 vivida, 68, 195

Neournula, 266 nordmanensis, 196 pouchetii, 39, 118, 196 Niessliaceae, 274, 277 Nodulisporium, 300, 330, 331 Numariola petersii, 287 Nummularia cinerea, 285 Octospora, 123, 195, 197, 240 axillaris, 6 crechqueraultii, 174 humosa, 69, 197 leucoloma, 197 leucolomoides, 197 tetraspora, 197 Ombrophila clavus, 375 pura, 401 turficola, 358 Onygena, 416 corvina, 33, 417 equina, 33, 417 Onygenales, 416 Ophiocordyceps, 296 acicularis, 341 crinalis, 342 dipterigena, 31, 296, 337 gracilis, 32, 337 macularis, 344 melolonthae, 32, 338, 344 michiganensis, 32, 341–342 myrmecophila, 31, 296, 337, 340 paludosa, 342 ravenelii, 32, 339 sinensis, 275 sobolifera, 289 sphecocephala, 32, 340 stylophora, 32, 344 superficialis, 32, 341–342 unilateralis, 275, 344 variabilis, 31, 275, 343–344 Ophiocordycipitaceae, 274–275 Orbilia, 434 delicatula, 65, 435 inflatula, 65, 435 leucostigma, 435

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Beug-final.indb 484

luteorubella, 66, 435 piloboloides, 66, 435 xanthostigma, 65, 435 Orbiliaceae, 434 Orbiliales, 434 Orbiliomycetes, 434 Otidea, 72, 122, 178 abietina, 200 alutacea, 57, 198–199 alutacea var. microspora, 198 cantharella var. minor, 198 concinna, x, 57, 198 doratophora, 178 grandis, 198, 200 leporina, 57, 198, 200, 444 leporina var. minor, 198 microspora, 198 onotica, 58, 200–201 rainierensis, 58, 202 silvicola, 272 smithii, 58, 202, 272 subterranea, 73, 91– 92 tuomikoskii, 58, 200 unicisa, 58, 198, 200 Pachycudonia monticola, 34, 39, 403 Pachyella, 121 adnata, 52, 203 babingtonii, 203 clypeata, 52, 203 depressa, 203 punctispora, 203 Pachyphloeus, 71, 73 austro-oregonensis, 18, 106, 107–108 carneus, 10, 18, 76, 105–106 citrinus, 18, 105, 107–108 conglomeratus, 105 marroninus, 105–106 melanoxanthus, 106 thysellii, 18, 106, 108 virescens, 18, 106, 108 Paecilomyces farinosus, 280 Paludella squarrosa, 363 Patella contradicta, 262

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coprinaria, 135 setosa, 252 Patellaria leptosperma, 384 Patellariopsis clavispora, 67, 400 Paxina acetabulum, 155 nigrella, 229 semitosta, 173 subclavipes, 166 Peckiella luteovirens, 324 Pedicularis, 410 Penicillium, 3, 416 alboaurantium, 280 Peridoxylon petersii, 287 Peziza, 71, 74, 121–122, 206 aeruginascens, 367 aggregata, 132 alaskana, 207 alcis, 210 ammophila, 16, 204, 257 ampelina, 205, 216 ampliata, 210, 222 anthracophila, 209 arenaria, 59, 205, 216 arvernensis, 58, 206 atroviolaceae, 211 aurelia, 356 azureoides, 69, 211, 219 babingtonii, 230 badia, 58, 207, 215 badioconfusa, 215 bovina, 210 brunneoatra, 215 carbonaria, 143 cerea, 221–222 chlora, 368 coccinea, 246 coprinaria, 135 craterium, 264 domiciliana, 11, 59, 206, 208, 210 dudleyi, 247 echinispora, 209 echinospora, 48, 209, 213 ellipsospora, 17, 98 epispartia, 125

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fimeti, 49, 210 fulgens, 133 furfuracea, 379 fusispora var. aggregata, 132 gemmea, 239 gerardi, 211 gerardii, 69, 211, 219 griseorosea, 47, 69, 219 hemisphaerica, 172 howsei, 205, 216 humosa var. humosa, 197 inquinans, 364 ionella, 211 irregularis, 178 limbricalis, 213 lobulata, 205, 216 maurilabra, 127 melastoma, 227 michelii, 59, 220 micropus, 221–222 nivalis, 210, 213–214 olivacea, 365 ostracoderma, 47, 212, 214 petersii, 48, 209, 213–214, 217, 231 phyllogena, 7, 58, 207, 215 praetervisa, 48, 205, 209, 211, 213, 216, 217, 219 proteana f. campbellii, 218 proteana f. proteana, 48, 214, 217 proteana f. sparassoides, 48, 217, 218 pseudoviolacea, 219 pura, 401 repanda, 222 saccardiana, 219 saccardoana, 59, 219 scutellata, 253 semitosta, 173 sepiatra, 214 sepulta, 90 silvestris, 206 stuntzii, 74 subisabellina, 214 sublilacina, 48, 205, 216, 219 subviolacea, 216 succosa, 59, 220, 223 sulcata, 155 varia, 59, 206, 208, 221–222 venosa, 141

versiformis, 366 vesiculosa, 49, 172, 220, 223 violacea, 205, 211, 219 virginea, 393 waltersii, 35, 224 Pezizaceae, 71, 121 Pezizales, 118–124 Pezizomycetes, 118–124 Phaeobulgaria inquinans, 364 Phaeohelotium, 362, 386, 404 carneum, 408 epiphyllum var. epiphyllum, 35, 66, 355, 404 Phaeosphaeria nodorum, 437 Phialea subcarnea, 407 Pholiota, 322 Phyllophaga, 339 Phymatotrichopis, 119 Phymatotrichum, 212 Picoa carthusiana, 104 Piptoporus betulinus, 316 Pithya, 124 cupressina, 66, 225–226, 362 vulgaris, 66, 225–226 Plectania, 124 floccosa, 176 latahensis, 266 melaena, 228, 237 melastoma, 51, 227, 228, 237 milleri, 51, 228, 237 nannfeldtii, 36, 228, 229, 237 nigrella, 236 occidentalis, 248 Pleosporales, 437 Pleurotus, 327 Plicaria, 121, 122, 209, 213 anthracina, 230 badia, 207 carbonaria, 47, 230, 231 endocarpoides, 48, 231 leiocarpa, 231 trachycarpa, 230, 231 trachycarpa var. muricata, 230 Pneumocystidomycetes, 441 Pneumocystis, 441

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Podophacidium xanthomelum, 68, 405 Podostroma, 276 alutaceum, 302 Polyporus, 327 squamosus, 334 Protocrea, 276, 316, 328 farinosa, 44, 312, 316 pallida, 44, 316 Prunus, 377, 381 Pseudaleuria quinaultiana, 57, 122, 232 Pseudeurotiaceae, 354 Pseudobalsamia magnata, 75 nigrans, 77 nigrens, 77 Pseudocollema cartilagineum, 132 Pseudombrophila, 123, 136 cervaria, 49, 233 deerrata, 234 hepatica, 233 merdaria, 49, 136, 233, 234– 235 porcina, 49, 136, 234 theioleuca, 49, 233 Pseudopithyella minuscula, 66, 226 Pseudoplectania, 124, 227 fulgens, 133 melaena, 237 nigrella, 69, 228, 236, 237 vogesiaca, 28, 237 Pseudorhizina, 119, 147 californica, 147 sphaerospora, 154 Pseudosarcosoma latahense, 51, 118, 228, 266 Pseudotis abietina, 200 Psilopezia, 119, 240 Ptychoverpa, 120 bohemica, 120, 267 Pulvinula, 123, 240 archeri, 69, 239 carbonaria, 47, 238 constellatio, 239

convexella, 47, 69, 239 laeterubra, 47, 69, 239 Pustularia catinus, 258 rosea, 244 Pyrenomycetaceae, 71– 72 pyrenomycetes, 274 Pyronema, 122, 240 confluens, 240 domesticum, 240 omphalodes, 47, 128, 238, 240–241 Pyronemalla, 240 Pyronemataceae, 122 Ramsbottomia crechqueraultii, 174 Rhizina inflata, 242 undulata, 48, 119, 242 zonata, 242 Rhizinaceae, 118 Rhizopogon meandriformis, 78 Rhodoscypha, 123 ovilla, 68, 243 Rhodotarzetta, 122 rosea, 47, 244 Rhytisma, 354, 371 acerinum, 406 americanum, 62, 406 punctatum, 62, 406 salicinum, 62, 406 Rhystimatales, 354, 371 Rhytismataceae, 354, 371, 382 Ribes, 277 Rosellinia, 278 quercina, 61, 345 subiculata, 61, 345 Roseodiscus, 408 equisetinus, 408 rhodoleucus, 33, 355, 408 subcarneus, 33, 355, 402, 407–408 Russula, 276, 277, 324 brevipes, 276, 321 Russulaceae, 325 Rutstroemia, 355, 370, 409, 413 bulgarioides, 36, 355, 409 macrospora, 413

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Rutstroemiaceae, 355 Ruzenia spermoides, 333 Sarcoleotia, 419 globosa, 419 turficola, 358 Sarcoscypha, 124 austriaca, 56, 245, 246, 247 coccinea, 28, 56, 232, 245, 246, 247 craterium, 264 dudleyi, 56, 245, 246, 247 occidentalis, 35, 245, 248 Sarcoscyphaceae, 124 Sarcosoma, 124 globosum, 52, 124, 249, 266 latahense, 228, 266 mexicana, 266 mexicanum, 266 Sarcosomataceae, 124 Sarcosphaera, 121 ammophila, 204 coronaria, 16, 121, 250 crassa, 250 eximia, 250 Scabropeziza flavovirens, 106 Scleroderma, 79, 276 Sclerotinia, 355, 378, 410, 413 coloradensis, 410 sclerotiorum, 410 sulcata, 34, 355, 410–411 tuberosa, 378 veratri, 34, 355, 410 Sclerotiniaceae, 355 Scodellina badia, 207 leporina, 200 onotica, 200 Scorias spongiosa, 51, 437, 439 Scutellinia, 122–123, 259 blumenaviensis, 64, 251 crucipila, 64, 136, 138, 251 erinaceus, 64, 138, 252, 254 kerguelensis, 64, 254 nigrohirtula, 252 pennsylvanica, 64, 253

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pilatii, 64, 254 scutellata, 64, 252, 253–254 scutellata var. leucothecia, 64, 253 setosa, 64, 252 theleboloides, 138 umbrorum, 64, 253–254 verrucipolaris, 253 Sepedonium, 276 brunneum, 319 chlorinum, 318 chrysospermum, 318 microspermum, 318 tulasneanum, 318 Sepultaria arenosa, 89 semitosta, 173 sepulta, 90 Sequoia gigantea, 134 Simplicillium, 277 Smardaea, 123–124 planchonis, 69, 211, 236 Sordaria, 3, 4 superba, 2 Sordariales, 277 Sordariomycetes, 274–278 Sowerbyella, 124 radiculata, 28, 255 rhenana, 28, 39, 126, 256 unicisa, 200 Spadicoides clavariarum, 301 Sparassis, 218 Spathularia, 354 flavida, 40, 412, 427 rufa, 40, 412 Spathulariopsis, 354 velutipes, 40, 412 Sphaeria hyalina, 320 sphecocephala, 340 stigma, 295 Sphaeronaemella helvellae, 25, 45, 150 Sphaerosporella, 122 brunnea, 46, 257 hinnulea, 68, 257 Sphagnum, 358, 402

I n de x to Sci en t i fic Na m e s

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Sporophagomyces, 276 chrysostomus, 44, 326 Spragueola irregularis, 432 Stamnaria, 408 Steccherinum ochraceum, 311 Stephensia, 74 shanorii, 98 Stereum, 276 Stromatinia rapulum, 378 Strumella coryneoidea, 264 Syngiocladium tetanopsis, 343–344 Tapesia, 355 fusca, 67, 400 Taphrina, 441 alni, 442 amentorum, 442 betulina, 442 epiphylla, 442 nana, 442 occidentalis, 61, 442–443 populina, 442 robinsoniana, 61, 442 Taphrinaceae, 441 Taphrinales, 441 Taphrinomycetes, 441 Taphrinomycotina, 441 Tarzetta, 124, 143, 206 ammophila, 204 bronca, 59, 258 catinus, 28, 258 cupularis, 39, 258 rosea, 244 Tatraea, 370 macrospora, 35, 355, 413 Terfezia, 71, 102 gigantea, 102 oligosperma, 115 Thecotheus, 130 Thelebolales, 354 Thelebolus stercoreus, 354 terrestris, 132

Thuemenella, 278 cubispora, 55, 346 Thuemenidium, 419 arenarium, 41, 419, 428 atropurpureum, 41, 355, 419, 428 Tolypocladium inflatum, 299 Torrubia capitata, 296 melolonthae, 338 ophioglossoides, 298 sphecocephala, 340 Torrubiella arachnophila var. leiopus, 347 arachnophila var. pulchra, 347 gibellulae, 347 Trametes versicolor, 334 Tricharina, 122, 124, 134, 259 gilva, 46, 90, 138, 172, 252, 259 Trichoderma, 276, 306, 328 aggressivum, 328 alutaceum, 302 atroviride, 314 citrinoviride, 308 hazarianum, 308 polysporum, 304 strictipili, 310–311 viride, 314 viridescens, 313–314 Trichoderma section Hypocreanum, 312 Trichoglossum, 419, 420 farlowii, 42, 429 hirsutum, 429 variabile, 429 velutipes, 42, 429 walteri, 42, 429 Trichophaea, 122, 134, 259 abundans, 46, 134, 262 brunnea, 257 contradicta, 46, 262 gilva, 259 gregaria, 262 hemisphaerioides, 46, 90, 172, 261 woolhopeia, 134, 261–262 Trichophaeopsis, 134 bicuspis, 46, 262

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Tritarchium shiotae, 284 Tubaria furfuracea, 335 Tuber, 71, 73, 110 album, 78 anniae, 115–116 beyerlei, 116 bisporum, 110 californicum, 20, 109 canaliculatum, 20, 110 candidum, 20, 113–114 castilloi, 116 gardneri, 116 gibbosum, 20, 111–112 giganteum, 111 guevarai, 116 lauryi, 116 levissimum, 116 lyoniae, 113 lyonii, 20, 21, 113–114 mexiusanum, 116 miquihuanense, 116 monticola, 116 oligosperma, 115 oregonense, 7, 20, 111–112 quercicola, 20, 114 rufum, 21, 114 separans, 21, 116 shearii, 20, 115–116 sphaerosporum, 20, 109 texense, 113 walkeri, 116 whetstonense, 116 Tuberaceae, 71

Tubercularia vulgaris, 334 Underwoodia, 119 campbellii, 218 columnaris, 26, 263 Urnula, 124 craterium, 36, 264 geaster, 139 mexicana, 266 padeniana, 51, 227, 228, 265–266 pouchetii, 196 Ustulina deusta, 332 Vaccinum, 381 Velutarina rufo-olivacea, 37, 390 Venturia inaequalis, 437 Venturiaceae, 437 Verpa, 120 bohemica var. bohemica, 21, 120, 189, 267, 268, 456 conica, 21, 120, 189, 267, 268 Verticillium, 298, 327 Vibrissea foliorum, 414 truncorum, 33, 355, 414 Vigariella, 329 Whetzelinia tuberosa, 378 Wilcoxina, 123, 134 rehmii, 68, 134

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sequoia, 134 Wolfina aurantiopsis, 53, 118, 142, 266, 269 Wynnea, 124, 178 americana, 58, 124, 270, 468 atrofusca, 272 sparassoides, 124, 270, 271 Wynnella, 119 auricula, 200 silvicola, 58, 272 Xylaria, 278, 332, 351 atropictor, 29, 348 corniformis, 351 cornu-damae, 29, 349 cubensis, 29, 348 curta, 351 filiformis, 29, 350 hypoxylon, 29, 349 liquidambar, 350 liquidambaris, 29, 350 longiana, 349 longipes, 29, 351 magnoliae, 29, 350 multiplex, 29, 349 oxyacanthae, 29, 349 persicaria, 349 poitei, 348 polymorpha, 29, 351 tentaculata, 29, 352 Xylariaceae, 278 Xylariales, 278 Xylocoremium, 348 flabelliforme, 29, 348

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