'~--------.

--

l
I

GASTEROMVCETES
Morphological and Development
Features
with
Keys to the Orders, Families, and Genera

by Orson K. Miller, Ir., and Hope H. Miller

illustrations by Cynthia Clem

GASTEROMYCETES
Morphological and Developmental
Features
with
Keys to the Orders, Families, and Genera

by
Orson K. Miller, Ir.
and

Hope H. Miller
Illustrations by Cynthia Clem

1988 Mad River Press

Published by Mad River Press, Inc.

141 Carter Lane
Eureka, CA 95501-9528
Printed by

Eureka Printing Co., Inc.

106 T Street
Eureka, CA 95501

ISBN 0-916422-74-7

TABLE OF CONTENTS

... ............ .... .. . .. .. ... . . .................. .. v

list of Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
legend for Abbreviations on the Figures . . . . . . . . . . . . . . . . . .. . . . . . . VII
Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . VIII
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
List of Plates

General Morphology
Spores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Basidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Hyphal Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Peridium ...... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

. .

..

.. .......... . .... . .. . ..... . ............. . ..

Key to the Orders of Gasteromycetes . . . . . . . . . . . . . . . . . . . . . . . . . .

Chemical Reagents

lycoperdales

25
29

31
Tulostomatales ............................................. . 48
Sclerodermatales 61
N idu lariales 69
Phallales . . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Gautieriales 96
Melanogastrales 98
leucogastrales . . . . . . . . , 101

.. .

102

122

123

loculate, Secotioid, and Agaricoid Gasteromycetes

Excluded Taxa
Glossary

Bibliography
Index

134

144

"'----------

-------------

>

LIST OF PLATES
Plate I.

A. Ca/vatia booniana. B. Langermannia gigantea.

C. Mycenastrum corium. D. Calbovista subsculpta. .. x

Plate
Plate
Plate
Plate
Plate
Plate
Plate

Basidiospores .................................... 6
Basidiospores .................................... 7
Basidia and Basid iospores ......................... 8
Peridium and Capillitium ........................ 14
Capillitium and Spores ........................... 15
Capillitium ..................................... 16
Capillitium ..................................... 17

II.
III.
IV.
V.
VI.
VII.
VIII.

LIST OF FIGURES
Figure 1
Figure 2
Figure 3
Figure 4
Figure 5
Figure 6
Figure 7
Figure 8
Figure 9
Figure 10
Figure 11
Figure 12
Figure 13
Figure 14
Figure 15
Figure 16
Figure 17
Figure 18
Figure 19
Figure 20
Figure 21
Figure 22
Figure 23

Development ...................................
Fruiting Body Types in the Lycoperdales ...........
Fruiting Body Types in the Lycoperdales ...........
Exoperidial Fibrils and Warts .....................
Lycoperdon perlatum ...........................
Lycoperdon marginatum .........................
Lycoperdon pulcherrimum .......................
Lycoperdon mammaeforme ......................
Bovista pila .....................................
Bovistella radicata ........... . ...................
VascellufT! pratense . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..
Gastrosporium simplex ............... ~ . . . . . . . . ..
Disciseda candida ...............................
Castoreum cretaceum ...........................
Mesophellia ....................................
Radiigera fuscog/eba ............................
Geastrum fornicatum ............................
Geastrum bryantii ...............................
Geastrum pectinatum ...........................
Geastrum saccatum .............................
Geastrum triplex ................................
Tulostoma verrucosum ..........................
Tulostoma simulans .............................

21
24
24
39
40
41
41
41
42
42

43
43
44
44
44
45
45
46
46
47
47
53
54

VI

Figure 24
Figure 25
Figure 26
Figure 27
Figure 28
Figure 29
Figure 30
Figure 31
Figure 32
Figure 33
Figure 34
Figure 35
Figure 36
Figure 37
Figure 38
Figure 39
Figure 40
Figure 41
Figure 42
Figure 43
Figure 44
Figure 45
Figure 46
Figure 47(
Figure 48
Figure 49
Figure 50
Figure 51
Figure 52
Figure 53
Figure 54
Figure 55
Figure 56
Figure 57
Figure 58
Figure 59
Figure 60
Figure 61
Figure 62
Figure 63
Figure 64
Figure 65
Figure 66

Chlamydopus meyenaianus ...................... 55

Schizostoma laceratum .......................... 55
Que/etia mirabilis ............................... 56
Phellorinia inquinans ............................ 56
Battarreoides digueti ............................ 57
Battarrea laciniata ............................... 57
Dictyocepha/os attenuatus ....................... 58
Ca/ostoma cinnabarina .......................... 59
Ca/ostoma lutescens ............................. 59
Ca/ostoma cinnabarina .......................... 59
Ca/ostoma insignis .............................. 60
Scleroderma po/yrhizon ......................... 65
Astraeus hygrometricus .......................... 66
Pisolithus tinctorius ............................. 67
Myriostoma coliforme ............... ;........... 68
Broomeia congregata ............................ 68
Cyathus striatus ................................. 73
Cyathus striatus ................................. 73
Sphaerobo/us stellatus ........................... 74
Dictyophora duplicata ........................... 86
Dictyophora duplicata ........................... 86
/tajahya ga/ericu/ata ............................. 87
Staheliomyces cinctus ........................... 87
Mutinus ravenelii ............................... 88
Phallogaster saccatus ............................ 88
Protubera jamaciensis ........................... 89
Kobayasia nipponica ............................ 89
Simblum sphaerocepha/um ...................... 90
Lysurus borealis ................................. 90
Aseroe rubra ........................ ,....' . . . . . .. 91
Kalchbrennera corallocepha/a .................... 91
Pseudoco/us fusiform is .......................... 92
Coltis hirudinosus ............................... 92
Laternea pusilla ................................. 93
Laternea triscapa ................................ 93
Linderia co/umnata .............................. 94
Anthurus archeri ................................ 94
lIeodictyon gracilis .............................. 95
Clathrus ruber ................................... 95
Gautieria graveolens ........................... 97
Me/anogaster variegatus ......................... 97
Podaxis pistillaris ............................... 116
Montagnea arenaria ............................ 116

..

VII

Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure
Figure

67
68
69
70
71
72
73
74
75

Truncocolumella citrina ....................... .

Thaxterogaster pingue ......................... .
Ga/eropsis desertorum ......................... .
Weraroa cucullata ............ . ................ .
Nivatogastrium nubigenum .................... .
Gyrophragmium defilei ............... . ........ .
Endoptychum agaricoides ...................... .
LonguJa texensis .............................. .
Brauniellu/a albipes

117
117
118
118
119
119
120
120
121

LEGEND FOR THE ABBREVIATIONS

ON THE FIGURES
ap
b
ca

Clr
co
col
dia
en

epJ
f
fun
g
gl
gu
h
ha

J
la
10

me
mp
0

p
pe

apophysis
basidia
calyptra
circumscissile dehiscence
collar
columella
diaphragm
endoperidium
epiphragm
fimbriate
funicular cord
gleba
glebifer

gussets
hollow center
hapteron
indusium
lantern
lacerate ostiole
mesoperidium
middle piece
ostiole
peridium (exoperidium)
pedicel

pr
ps
pc
pu
r
re
rh

n
s
sc
se
sh
sp
sr
st
stc
str
sub
sui
tr
tu
v
ve

peridiole
peristome
pseudocolumella
purse
ray
receptacle
rhizomorph
ridges
stoma
sand case
secotioid
sheath
spores
saccate ray
stalk (stipe)
sti pe-col umella
striate (striae)
subgleba
sulcate
trabeculae
tunica
volva
veil

VIII

PREFACE
There are many macrofungi which are classified under the ambiguous category of Gasteromycetes, but very little has been published
concerning their ecology, morphology, and development. In fact, the
gulf between the technical writings on Gasteromycetes and popular
field guides is very wide. The lack of common knowledge of the
anatomy and morphology of the group has by itself hindered most
interested mycologists who are not students of higher fungi from
actively seeking information on species or even generic determinations
of taxa. The first objective of this initial volume is to provide information concerning the anatomy of the Gasteromycetes, the definition of
the terms used to describe the taxa involved, and references to the
pertinent works which have been published. The second objective is to
establish concepts of the most widely accepted genera and to provide
keys to enable one to identify the Gasteromycetes to genus. It is neither
possible nor practical to include only genera from North America or
the Northern Hemisphere, since many genera have species which are
widely distributed decomposers or have species which have been
introduced to non-native habitats in the past 50 to 100 years. Indeed,
the potential for this to occur is ever present, and the likelihood of
discovering newly introduced taxa seems to be greater than with other
higher fungi. It was decided, for these reasons, to include under one
cover all the major or well-documented genera now known. It is hoped
that this work will stimulate the production of regional mycofloras of
species, subspecies, and varieties, which will provide us with a l\Iuch
clearer appreciation of this fascinating group of fungi. A third objective
is to focus attention on generic concepts which need critical investigation.
We have received much cooperation, many good suggestions, and
invaluable help from both amateur and professional mycologists. We
are especially indebted to Dr. William Burk, Department of Botany,
Univ. of North Carolina; Dr. Roy Watling, Senior Scientific Officer and
Curator of Fungi, Royal Botanic Garden, Edinburgh, Scotland; and Dr.
Curry Marr, Biology Dept. SUNY, Oneonta, NY for their critical reviews
and suggestions during the preparation of the manuscript. The excellent illustrations by Cyndi Clem have truly made the fungi come alive.
We also appreciate the information, reprints, and loan of specimens by

IX

Dr. David Farr and Dr. Gerald Bills, Mycological Collections, BARCWest, USDA, Beltsville, Maryland, Dr. Steven Miller, Forestry Sciences
Laboratory, Corvallis, Oregon, and Dr. D. Jean Lodge, University of
Puerto Rico, San Juan, Puerto Rico. We greatly appreciate the translation of many passages in the "Gasteromycetes" by Dr. Pilat from
Czechoslovakian to German by the late Dr. Ruzina Hilber. We acknowledge the cooperation and support of the Biology Department at Virginia Polytechnic Institute and State University and the University of Montana Biological Station. Lastly, our graduate students have offered many
insights and much encouragement during the preparation of this book.
Their enthusiasm, stimulation, and help are greatly appreciated.

plate I. A. Ca/vatia booniana has discrete, medallion-like scales over

the surface of the exoperidium. B. Langermannia gigantea has a
smooth exoperidium. C. Mycenastrum corium develops a characteristic checkered exoperidium in age. D. Calbovista subsculpta has an
exoperidium of truncated warts over the upper surface and a thick
endoperidium.

INTRODUCTION
Traditionally all taxa which are unable to carry out forcible spore
discharge and have spores mature within the fruiting body were
automatically placed within the group of orders which make up the
Gasteromycetes. This series of orders of Basidiomycetes includes the
Lycoperdales (puffballs, earth stars, etc.), Tulostomatales (stalked puffballs), Sclerodermatales (earth balls), Phallales (stinkhorns), and Nidulariales (birds' nests). Along with these, three lesser-known orders
including the Gautieriales, Melanogastrales, and Leucogastrales are
also included in this book. In addition, a series of secotioid and
agaricoid fungi considered to be Gasteromycetes but related to the
lamellate or gilled fungi and to the boletes or pored fungi are also
included, in the section "Loculate, Secotioid, and Agaricoid Gasteromycetes".
The word "gasteromycete" is derived from the Greek "gaster",
which means "stomach", and "mycetes", which means "fungus". The
prefix "gastero-" refers to the developmental process in which the
spores mature within the fruiting body and are protected by the sterile
outer layers known as the peridium. After the developmental process,
the mature basidiospores are disseminated by various means. The chief
methods include wind, rain, insects, and mammals, but forcible discharge of the spore from the basidium does not exist. A basidium which
cannot forcibly discharge its spores is referred to as statismosporic.
Contemporary study of many Gasteromycetes by the use of transmission electl'on mkroscopy (TEM) and scanning electron microscopy
(SEM) has been augmented by more sophisticated studies of fungal
ecology. The relationships of these fungi to those that forcibly discharge their spores is now often known. Most are either decomposers
of organic matter such as wood or needles or are mycorrhizal with
higher plants including trees and shrubs. As a result of these studies,
many mycologists have become convinced that some Basidiomycetes
which now function as Gasteromycetes are in reality very closely
related to the Russulales, Boletales, and Agaricales, which forcibly
discharge their spores. Their anatomical characteristics and ecological
roles are the same as those of their counterparts which forcibly
discharge their spores, and they are now placed within their respective
families in the three orders referred to above. These are referred to as

secotioid and agaricoid taxa. They are usually epigeous at maturity and
are prominent in many habitats, including the desert biome. We have
also included some taxa in the Hymenogastrales. These fungi have
lamellae, pores, and/or other features of taxa which forcibly discharge
their spores but have no obvious relationships to the Russulales,
Boletales, and Agaricales. Their position is usually referred to as
unknown, and the Hymenogastrales is the order in which they are
placed.
One of the objectives of this work is to present the diagnostic
features of the Gasteromycetes. Each of the orders is described briefly.
This is done to illustrate how the construction of the fruiting body has
embodied adaptive features to aid it in both protecting the developing
hymenium and discharging the mature spores. The structures which
have evolved to accomplish passive spore dispersal depart markedly
from those associated with forcible spore discharge. The distinctive life
forms of Gasteromycetes have led to the designation of many genera
which are either monotypic or contain only a few species. This means
that the macroscopic appearance of many species makes them recognizable at once. The distinctive appearance is particularly true in the
Phallales, for example. On the other hand, the majority of hypogeous
taxa in such genera as Alpova and Rhizopogon must be identified by
the use of microscopic features. It follows that more macroscopic
features will be shown in orders where the emphasis is on identification
by this means. Conversely, relatively few hypogeous taxa will be
illustrated because they cannot be identified to either genus or species
by macroscopic means.

GENERAL MORPHOLOGY
Spores
Spores of the Gasteromycetes are orthotropic in development and
statismosporic, as illustrated in Plate IV A & B. The attachment of the
spore to the apiculus is symmetrical, usually by a short or long pedicel,
a portion of which remains attached to the spore. The area of
attachment often forms a collar and is most frequently thickened just at
the spore (PI. IV F). In some pedicellate species the spore breaks off
close to the basidium, leaving a long pedicel (PI. II D & E) attached to
the spore. Some species, such as Lycoperdon pedicel/a tum, are characterized by pedicellate spores. In the Phallales and Nidulariales a hilar
scar (hilum) is evident after the spores are detached from the basidium.
Spores of the Gasteromycetes related to agaric orders, may be heterotrophic in development but have lost the ballistosporic ability and can
no longer forcibly discharge their spores. The spores, however, are
canted to one side and are not oriented directly over the sterigmata
(see Largent et aI., 1977).
There are as many as five identifiable wall layers in some spores.
When spore development occurs, the walls synthesized by the basidium may be composed of one or two wall layers. If there is one layer it
is referred to as a perisporium (PI. II B), but if there are two layers, the
outer layer is an ectosporium and the inner layer is a perisporium. The
spore itself may synthesize three wall layers beneath these two,
including an outer exosporium, an episporium which is often thick and
well developed, and an endosporium which is next to the plasma
membrane. The ultrastructure of most Gasteromycete spores has not as
.
yet been determined.
There is a strong tendency for the spore shape and morphology to be
quite similar within a given order of Gasteromycetes. However, the
shape and morphology vary considerably among the orders. The
Phallales have uniformly cylindric spores with a gelatinous or mucilaginous exosporium, while the Lycoperdales have ovoid to globose
warted spores (PI. II A-F). In general, the warts are synthesized from the
exo- or episporium. Many species within the Tulostomatales have a
spore morphology (PI. VI D) which is very similar to that of the
Lycoperdales. The Sclerodermatales are typified by reticulate-warted
(PI. IV D) or warted spores (PI. IV C) which are ovoid to globose. They
are constructed somewhat differently from those of the orders described above. The Nidulariales have short elliptical, ovoid, to globose

spores which are smooth with a thick-walled to thin-walled exospor

lum.

The spore morphology in these orders has evolved to accommodate

spore dissemination. The mucilaginous covering of the insect-disseminated spores of the Phallales causes them to sti,ck to the feet or legs
of insects. The spores of the Lycoperdales, Tulostomatales, and Sclerodermatales are all wind-disseminated and are ovoid to globose with
warts or spines. The shape not only accommodates wind dissemination
but the warts and spines enable the spores to stick readily to various
kinds of surfaces. The Nidulariaceae (Nidulariales), on the other hand,
have peridiole dissemination, which usually takes place in open areas
by the splash-cup system. Splashing raindrops tend to knock the
peridioles out from a cluster of fruiting bodies in all directions, so that
they form a circle surrounding the cluster. New colonization of woody
substrates often occurs, therefore, in a circular area surrounding the
old fruiting bodies. On slopes, new colonization may be observed
along natural drainage patterns where the peridioles have been washed
by rain water. The Sphaerobolaceae (Nidulariales) have similar fruiting
bodies, but they contain only one peridiole. At maturity, the entire
peridiole is forcibly discharged up to 2 or 3 meters from the fruiting
body. The actual discharge is a light-dependent response. The peridiole
eventually breaks down, releasing the spores which germinate and
colonize more woody substrates.
The Order Hymenogastrales is also considered to be in the series
Gasteromycetes. Both hypogeous and epigeous taxa are included in
this order. In contrast to the orders discussed above, the spores vary
widely within this order, as illustrated in Plates IV B, E, and F, and many
of the genera are not closely related to each other. The mechanism of
forcible spore discharge appears to have been lost by some fungi, while
in others it apparently never evolved. The largest group of genera is
composed of hypogeous taxa, which form fruiting bodies within the
duff or soil. Spore dissemination takes place by decomposition of the
fruiting bodies and percolation of the spores through the soil. In many
taxa this event is preceded by rodent or invertebrate herbivory. The
thick-walled, often ornamented spores found in Hymenogaster (PI. IV
B), for example, are capable of withstanding passage through digestive
tracts of animals and severe soil conditions to reproduce the species.
Characteristic odors produced by volatile compounds signal the maturity of the spores and attract mammals and invertebrates to the cryptic
locations of the fruiting bodies.
A smaller but widely distributed group of epigeous, secotioid genera
also exists within the Hymenogastrales and within families of the agarics
(Agaricales and Russulales). The striking similarities of many of the taxa
"

".

to agarics with forcible spore discharge have resulted in their placement within agaric families. Other taxa, with less certain affinities, are
retained within the Hymenogastrales or placed in the' Podaxales. In
general, the epigeous secotioid taxa such as Montagnea and Podaxis
(Figs. 66, 65) are well adapted to xeric micro-habitats and are typically
found in deserts throughout the world. Secondary modifications of the
fruiting bodies have usually involved the development of a capillitial
hyphal system, thickened peridial layers, and a modified pileus which
no longer opens, as in Nivatogastrium (Fig. 71) or Brauniellu/a (Fig. 75).
Spore dissemination usually takes place when the peridium is weathered away and the spores are blown from the disintegrating gleba or
are eaten by rodents. In some cases, as illustrated by Montagnea (Fig.
66), the trama expands, recurves somewhat, and becomes hard or
indurate creating a series of upraised plates called gussets. The gussets
stand exposed to the desert winds, which erode away small pieces of
the gussets covered with a layer of thick-walled spores. The spores of
most of the taxa associated with xeric or desert habitats have become
thick-walled and pigmented, as in Podaxis (PI. III D), adaptations which
make them more resistant to desiccation and lethal UV radiation .

Plate II. Basidiospores. A. Mature basidiospores of Ca/vatia with warts

and short pedicel, on left. B. Basidiospore of Calvatia . with pieces of
perisporium and warts of exosporial origin. C. Basidiospore showing
the hollow pedicel through which the nucleus passed into the develop- .
ing spore. D. Basidiospore of Bovista plumbea with fine exosporial
warts and a long pedicel. E. Basidiospore of Bovista showing the
exosporial warts and the pedicel, below right. F. Orthotropic basidiospore of Calostoma with echinulations.

c.

D."'_......

E.

,I

i F.
Plate III. Basidiospores. A. Pitted spores of Ca/ostoma. B. Low warts
an d persistent pedicel of Gastrosporium spore. C. Orthotropic spores
with a collar, Me/anogaster. D. Orthotropic, statismosporic spores of
Podaxis with apical germ pore (see I), very thick wall, and central lipid
body. E. Spores of Montagnea with thickened wall and subapical germ
pore (see I). F. Endoptychum spores with an open-pored hilum and a
th ick wall, but without an apical pore.

c.

D.

Plate IV. Basidia and Basidiospores. A. Plectobasidia of Podaxis pistillaris; note the orthotropic spores with an apical germ pore, erect on the
basidia. B. Orthotropic spores on hypha-like basidia of Hymenogaster.
C. Warted spores of Scleroderma cepa. D. Reticulate-warted ornamentation of Scleroderma citrina. E. Angular spores of Nigropogon
asterosporus with a very prominent apiculus. F. Orthotropic spores of
Hysterangium with a well-formed collar formed by the separation of
the spore from the pedicel.

Basidia
The Gasteromycetes which are unrelated to agaric families have
highly variable basidia which arise irregularly from glebal hyphal tissue.
This type ,of basidium is called a plectobasidium (Pl.-IV A), and it is not
oriented to project into an air space from a hymenium organized to
accommodate forcible spore discharge. Plectobasidia often form fertile
layers or areas referred to as a pseudohymenium. These fertile areas
can, for example, surround smalilocuies in the gleba, be concentrated
within peridioles, or occur as layers on the receptaculum of a stinkhorn. However it is organized, the plectobasidium facilitates some
non-forcible type of dissemination. The lack of uniformity which
typifies the ballistosporic basidium arises for several reasons. First, the
basidium is statismosporic (PI. III D) and not oriented for forcible spore
discharge. This means that a clavate, or narrowly clavate, form with
apical sterigmata is not necessary. Secondly, other shapes and sterigmatic placement are undoubtedly selected to facilitate other types of
spore dissemination. In some cases, reasons for the form are strongly
suggested by the observed function. In other cases there is no reason
which has as yet become apparent to link form and function. In some of
the plectobasidial fungi several generations of basidia may develop
within one fruiting body (Dring, 1973). Their function varies. It is
theorized that some act as nurse-hyphae, providing nutrients to spores
shed prematurely from the plectobasidium.
The gastroid basidium is basically a single cell, or holobasidium.
Spore attachment is either orthotropic or less commonly heterotropic
but, by definition, is always statismosporic. The taxa with this type of
basidium are included within the series Gasteromycetes excluded from
the subclass Holobasidiomycetidae (Alexopoulos and Mims, 1979).
Passive spore detachment is a unifying characteristic of the Gasteromycetes. Moreover, this cbaracteristic is usually accompanied by dramatic alterations in the morphology of the fruiting body to accommodate spore discharge. The basidium, as indicated above, is very
polymorphic, ranging from an undifferentiated, hyphal-like structure
(PI. IV B) to an ovoid or subglobose cell (PI. III D). Adaptations to insect
dissemination in the Phallales have led to the development of a very
well formed, homogenous hymenium with a strict apical orientation of
the sterigmata, which are whorled about the apex. Eight spores per
basidium are common in Mutinus and Phallus. Conversely, there is no
selection for a strict orientation of the sterigmata within the developing
peridiole of the Sclerodermatales. In this order the placement of the
sterigmata is nearly random and spores may develop at any point over
the whole upper surface of the basidium, as in Astraeus.

10

In the Lycoperdales, locules form as the hymenium matures. The

basidia develop in clusters surrounding the locules. The sterigmata are
more or less apically oriented, and a delicate tube, the pedicel (PI. II 0),
extends from the basidium into the locule. Spores develop on the
apices of the pedicels (PI. II E) and at maturity drop into the locule.
Since there is no forcible spore discharge, this process has the effect of
placing the spores within the center of the locule where they are not
entrapped within the collapsing basidial layer. In some species the
pedicel breaks at or near the basidium (PI. II 0), leaving a long pedicel,
while in other species the break is closer to the spore, leaving a short
pedicel (PI. II A).
There is a strong tendency in the Nidulariales toward narrowly
clavate to hypha-like basidia. Although no pedicels form, the basidia
often have long hypha-like stalks and extend out into the locule, where
the spores are deposited at maturity (Fig. 41). The thin stalks may
elongate up to 45 jLm, as in Cyathus stercoreus. The same function is
accomplished by the elongated pedicel in the Lycoperdales (PI. II 0).
It is only in the Hymenogastrales and secotioid agarics that heterotropic, statismosporic basidia are observed. It is assumed by many mycologists that the secotioid agarics, through adaptations, have . lost the
forcible spore discharge mechanism (Miller & Watling, 1987). The
basidium, in some cases, still has a hilar appendix body which is present
in the ballistosporic basidium but it is no longer functional (5. Miller,
1985) . In most cases the apical orientation of the sterigmata has not as
yet been lost. In some desert-adapted taxa, such as Podaxis, the
basidium has become thick-walled the gleba has lost all evidence of
tramal plates, and plectobasidia are present (PI. IV A).

11

Hyphal Systems
The hypha I systems which have evolved in the Gasteromycetes do
not differ materially from those of the Basidiomycetes in general. A
thorough discussion of these systems is presented in "How to Identify
Mushrooms to Genus III: Microscopic Features" (Largent et aI., 1977).
Therefore, this section will deal with hyphal organization and differentiation peculiar to or repeatedly encountered within the Gasteromycetes.
In the mitic system or "hyphal analysis" system described by Largent
et al. (1977) as applied to the Gasteromycetes, one would recognize
only two hyphal types, namely generative and skeletal hyphae. The
hyphal systems, therefore, may be either monomitic or dimitic. If
dimitic, the system would contain both generative and skeletal hyphae,
since binding hyphae (see Largent et aI. , 1977, Fig. 8) are not reported
within the Gasteromycetes. Skeletal hyphae in the gleba of the Gasteromycetes are commonly referred to as capillitium, which is of two types:
true
(PI. V E & VIII A), which is usually cyanophilic, and
paracapillitium (PI. VIII C), which is acyanophilic.
True capillitium is composed of hyphae 4-35 Jlm wide with secondarily
thickened walls and a lumen which may be large or small (PI. VII C & D),
depending upon the wall thickness. The walls are usually pigmented
brown, septation may be common to rare, and branching occurs
frequently to rarely. Some capillitial elements contain pores (PI. VII C),
others have distinctive slits (PI. VI A), while others have neither (PI. V E).
Capillitial elements may also be equal or extremely variable in width
and are often very flexuous (PI. VII D). In some cases a tapering system
is differentiated, with ultimate tapering ends only 1-2 Jlm wide (PI. VIII
D). True capillitium is characteristically confined to the Lycoperdales
and Tulostomatales and is either positively stained with aniline blue in
lactic acid and phenol (cotton blue in lactophenol) or has brown, thick
walls. The positive reaction is commonly known as the cyanophilous
reaction initially described by Kotlaba (1964). Paracapillitium and generative hyphae are not cyanophilous; that is, their walls do not absorb
the stain (PI. VIII C). These are referred to as acyanophilic walls. A
dimitic system arises when either paracapillitium or true capillitium or
both develop within the gleba of one species. Vascellum is dimitic, with
abundant paracapillitium (PI. VIII C) and scattered, usually peripheral,
capillitium. In Bovista paracapillitium is regularly formed during development, but it may be scarce or absent at maturity.
Paracapillitial cells are made up of generative hyphae which are
unpigmented, regularly septate, and thin-walled sometimes becoming
thick-walled in age (PI. VIII B), as described by Kreisel (1962). Most

12
paracapillitium is infrequently to frequently branched. Paracapillitium
is the only type which has developed in the Sclerodermatales. At times
the paracapillitial cells may have very slightly thickened to thick walls
when compared directly with generative hyphae, but no pores or slits
are present. They may also have a lightly pigmented context, and clamp
connections are sometimes present, as shown in Astraeus (PI. VII B).
Clamp connections are absent in true capillitium.
Special reduced capillitial elements in the gleba of Battarrea and
Battarreoides are called elaters. These free or solitary units are usually
from 30 to 130 p,m long (PI. VI C & D), taper at both ends, and have thick
spiral bands at maturity. They are not branched and are known only in
the above genera, where they comprise the only capillitial type. It is
hypothesized that as they dry or moisten under desert weather conditions they rapidly coil or straighten out, freeing the highly warted
spores which are then wind-disseminated. Similar capillitial units, but
without spiral bands, are differentiated in the mature gleba of Chlamydopus (PI. VIII B).
The peridium (detailed in the next chapter) is quite variable throughout the Gasteromycetes and ranges in general from one to three layers.
A typical three-layered peridium would have an exoperidium, mesoperidium, and endoperidium. The tissue types which make up these
layers are sometimes characterized by the system described for the
Ascomycetes by Korf (1958) and adapted by Miller (1971) for the
Basidiomycetes in culture. These types are as follows:
Textura globosa-composed of round to ovoid cells, with thin or

slightly thickened walls, often pigmented, with intercellular spaces

(often characterized as pseudoparenchymatous).
Textura angularis-composed of polyhedral cells, thin- or 'thick-walled,

pigmented or not, without intercellular spaces (often characterized as

pseudoparenchymatous).
Textura intricata-composed of interwoven, separate hyphae, without

common walls, with usually thin or slightly thickened walls, rarely

pigmented; clamp connections sometimes present.
Textura epidermoidea-usually forms a firm to tough layer of cells with

what appear to be common walls or walls which are tightly bound

together, leaving few intercellular spaces; commonly pigmented.
Textura oblita-composed of hyphae of interwoven to parallel, thick-

walled, usually pigmented cells; clamp connections sometimes present.

13
Textura prismatica-composed of small, angular, sometimes nearly

isodiametric cells, thin or slightly thick-walled, usually with what

appear to be common walls.
In general, clamp connections are rare in peridial layers. However,
they are occasionally found, as in Mycocalia in the Nidulariales. In
these cases one observes an exoperidium of interwoven hyphae with
clamp connections at each septum, a textura intricata .

14

~
I

.~

Plate V. Peridium and Capillitium. A. Setae in the cuticle of Lycoperdon me/anesicum. B. Close-up of several thick-walled setae. C.
Cuticle of exoperidium is a textura prismatica in Calvatia fragilis.
D. Connivent fibrils composed of a polycystoderm in Calvatia cretacea. E. Spiny capillitium in the gleba of Mycenastrum.

15

D.

Plate VI. Capillitium and Spores. A. Capillitial element with slits

(see I ) surrounded by globose, warted spores in Calvaria. B. Exoperidial hyphae which form the unique, powdery tomentum of
Nidu/aria. C & D. Spirally thickened capillitial elements of the gleba of
Battarrea, known as elaters, surrounded by the globose, warted
spores.

16

'",

'

D.

Plate VII. Capillitium. A. Capillitium of Mycenastrum. B. Paracapillitium with clamp connection (see / ) of Astraeus. C. Capillitial element with pores (see / ). D. Capillitium of Lycoperdon showing
thick-walled, branched, flexuous hyphae.

17

t.

c.

Plate VIII. Capillitium. A. Capillitium with swollen septa (see / ) of

Tu/ostoma finkii. B. Elongate, thick-walled capillitial units of Chlamydopus meyenianus. C. Paracapillitium of Vascellum showing thick but
unstained walls in cotton blue in lactic acid (acyanophilic). D. Tapering
capillitial units of Bovista plumbea; note the central, thick-walled cell
with branches tapering to a very small diameter.

18

Peridium
"Peridium", from the Greek "peri" meaning around, refers to the
sterile outer layers of the Gasteromycete fruiting body. There are
usually from one to three walls which surround the fruiting body (Fig.
16). The term peridium is used for species with one wall; exoperidium
and endoperidium when two walls are present; and exoperidium,
mesoperidium, and endoperidium for those with three wall layers. The
outer wall or walls may remain persistent over the base and serve as a
cup or volva from which the mature sporocarp expands to full maturity.
This is generally the case in the Phallales, as shown in Figures 43 and 44,
and is often found in the Tulostomatales (Figs. 28 & 29) where the young
embryo is borne underground and is protected during development.
Members of the secotioid Hymenogastrales, such as Montagnea (Fig.
66) and Gyrophragmium (Fig. 72), which develop hypogeously, also
have a volva. In the Nidulariaceae the peridiallayers act as a splash cup
(Figs. 40 & 41), which serves in the dissemination of the peridioles. The
peridioles in turn have a series of wall layers (Fig. 41) which protect the
developing hymenium. The name given to these wall layers is the
tunica, which may have at least two morphologically distinct walls. In
the Sphaerobolaceae the peridium (Fig. 42) is also persistent, four walls
thick, and serves to hold the endoperidium when it rapidly evaginates
in the process of violent peridiole discharge. The single peridiole
surrounds the hymenium and is composed of a single, thick wall.
The exoperidium in the Lycoperdales serves to protect the developing fruiting body. In mid- to late maturity, the exoperidium begins to
crack or slough off, or it simply ceases to grow (Figs. 6 & 9). Meanwhile,
the often tougher endoperidium undergoes morphogenesis, in some
genera, to form a stoma often with a characteristic type of fibril
surrounding it (e. g. fimbriate peristome, Fig. 21). Sometimes the
exoperidium remains, as in Lycoperdon mammaeforme (Fig. 8), in the
form of patches or some other recognizable characteristic tissue: The
drastic changes during the entire developmental process, compounded
by changes brought about by the vicissitudes of often severe weather
patterns, can greatly alter sporocarp appearance. Exoperidial layers are
often composed of fibrils, connivent fibrils, and pyramid-shaped warts
(Fig. 4), which are characteristic of a given species. Some species of
Lycoperdon produce thick-walled, irregular exoperidial cells called
setae (PI. V A & 8) by Demoulin (1976). The endoperidium is often
coriaceous or papyraceous, an adaptation which enables it to survive
over weeks or months during which the spores are gradually disch.uged. Spore discharge usually occurs when droplets of rain or dew
strike the surface of the endoperidium, causing small clouds of spores

,
19

to be discharged from the stoma in Lycoperdon, Disciseda, and

Tulostoma, for example (Figs. 7, 13, 22). In Bovista the entire sporocarp
becomes detached, and dissemination of spores occurs as the windblown sporocarps tumble over the landscape. The tough, exposed
endoperidium (Fig. 9) eventually becomes lacerate as it tumbles about,
slowly releasing the contents. In Gastrosporium the apex of the partially
buried sporocarp is weathered away (Fig. 12), exposing the gleba.
In the Sclerodermatales there are one to several peridial layers. At
maturity, however, the entire peridium sloughs off to expose the naked
gleba (Fig. 35). The peridium may weather away as in Pisolithus (Fig. 37),
or split and become recurved as in Scleroderma polyrhizon (Fig. 35).
The gleba is either blown away or washed away, usually leaving the
remains of the tough peridium. It is in Astraeus that an endoperidium is
persistent and serves to protect the gleba (Fig. 36), as in Geastrum in the
Lycoperdales. The exoperidium in both genera splits and recurves (Figs.
36 & 18) in a stellate pattern (resembling a star), giving rise to the
common name "earth star". In Astraeus and some species of Geastrum,
the exoperidium is hygroscopic. The exoperidium recurves as it dries
and covers the endoperidium but gradually expands again as it becomes wet and exposes the spore sac. This mechanism obviously
enhances spore discharge under the most favorable conditions, but it is
present in only a few species. Several species of Geastrum, including G.
fornicatum (Fig. 17), have a persistent cup-like exoperidium which is
attached to the soil by rhizomorphs. The mesoperidium evaginat,e s but
remains attached to the margin of the cup. This effectively raises the
gleba (enclosed by the endoperidium) into the air and holds it in place,
facilitating spore discharge. The species in Radiigera are closely related
to Geastrum, as described by Askew and Miller (1977), but no stoma
develops in Radiigera, and the exoperidium is persistent (Fig. 16A).
Dissemination of the spores is carried out by invertebrates or rodents,
such as squirrels, which eat the gleba and carry the sporocarps about in
the process of creating caches of sporocarps.
In the Tulostomatales, Tulostoma is similar in morphology and
function to the Lycoperdales. However, in other genera considerable
differences exist. Schizostoma (Fig. 25) differs in the extreme splitting
and longitudinal dehiscence of the endoperidium to expose the gleba.
Chlamydopus develops hypogeously and is enclosed by a persistent
volva. The endoperidium is exposed as growth and expansion of the
stipe takes place (Fig. 24) . A small irregular stoma develops to accommodate spore dissemination, similar to Tulostoma. A series of tough
exoperidial layers protects the developing Battarrea buttons as they
mature in the soil (Fig. 29). The young expanding fruiting body pushes
through the tough exoperidium and grows to full maturity. The gleba is

20
protected by the endoperidium during the growth of the stalk. In age
the endoperidium dehisces in a circumscissile manner, exposing the
gleba on the convex lower part, as shown in the two sporocarps of
Figure 29. Under desert conditions with limited periods of moisture,
the complete exposure of the gleba to the wind provides a rapid means
of spore dispersal. In Battarraeoides the development is similar, but the
spores escape at maturity through several ostioles which develop over
the endoperidium, as shown in Figure 28. The spores are discharged in
quantity when rain drops strike the endoperidium, insuring moisture
during periods of spore release.
Under the more humid conditions of the southern temperate deciduous forest, the exoperidium of the species in Calostoma develops a
thick, gelatinous (Fig. 31) to a somewhat thinner, viscid layer (Fig. 32).
This layer is most extreme in C. cinnabarina Desv., best seen in color in
Miller, 1972, pI. 369. As the viscid, lacunose, gelatinous stalk expands,
the exoperidium sloughs off, exposing the endoperidium and the
raised, delimited peristome (Figs. 33 & 34) (Castro-Mendosa et aI., 1983).
Since these taxa may fruit in late fall or winter, the exoperidium may
serve to protect the maturing gleba from the vicissitudes of weather as
well as insect attack and herbivory.
The peridium in the Hymenogastrales is quite variable and depends
to a large extent on the nature of the sporocarp. Genera such as
Endoptychum (Fig. 73), Nivatogastrium (Fig. 71), Montagnea (Fig. 66),
Podaxis (Fig. 65), and Longu/a (Fig. 74) have peridial layers similar to
those in the agaric families to which they are related. These agaric
tissues are described in detail by Largent et al. (1977) in Chapter V "The
Pellis or Cortex". The arrangement of the peridial tissues in a derm or
cutis parallels in every way the spore characteristics, cystidia. and other
features of the agaric families. Specific affinities of secotioid genera, as
we understand them at the present time, are discussed under the
individual generic descriptions.

21

A.

B.

"

'.

'

..

.-

. . . . . ..
.' ".
I

. '

c.

E.

D.

F.

~h

+-h

Fig. 1. Developmental types in Gasteromycetes. A. Homogeneous. B.

Lacunar. C. Forate. D. Aulaeate. E. Pileate. F. Multipileate. (h) hymenium. Arrows indicate direction of growth of primordial glebal
tissue.

22

Development
Lohwag (1926) first detailed the process of development in the
Gasteromycetes and recognized the important differences which occur
at the generic, family, and ordinal levels. The differentiation of fungal
structures in the fruiting body has evolved to facilitate spore dissemination. However, the same result has often been achieved by rather
diverse means. Since large taxa with world-wide distribution have
essentially the same type of development, it would appear that these
types have considerable evolutionary significance. Development, then,
should not be viewed solely as an exercise in comparative anatomy.
Relationships among visually or superficially unrelated taxa have been
found to be very close upon detailed examination of development, and
the taxa often share the same biological role (Miller, 1982). The process
of development is one of the characteristics which must be an integral
part of the full or complete study of any taxon or series of taxa.
There are six methods of development which have be~_n described
and refined by various mycologists (Lohwag, 1926; Gaumann, 1926;
l,3essey, 1950; Kreisel, 1969; and Dring, 1973). The methods include
homogeneous, lacunar, forate, aulaeate, pileate, and multipileate development. Pilch (1958) was the only one who recognized the homogeneous type. Each of these developmental types is described below
and illustrated in Figure 1.
1. Homogeneous (Homogen) (Fig. 1 A): A simple type of development
in which the basidia are evenly distributed throughout the gleba and
are maintained in this pattern during the developmental process.
Tu/ostoma is an example given by Pilat (1958).

2. Lacunar (Kammerartig) (Fig. 1 B): A homogeneous glebal tissue is

differentiated into a series of regular chambers which are lined by a
well-developed hymenium (Fig. 1 B). Typical representatives of this
type are the members of the Nidulariales and Melanogastrales (Fig. 64).
In the Sclerodermatales it is best observed in Piso/ithus (Fig. 37) and
Astraeus (Fig. 36), which show initially well-developed peridioles lined
by the hymenium.
3. Forate (Koralloid) (Fig. 1 C): Differentiation of plectobasidia occurs
among preformed primordial tissue within minute locules arising near
the peridium and maturing inward (centripetal). Development occurs
from the periphery of the gleba near the peridium or at a centrum in
the region of the sterile base, as in Lycoperdon (Fig. 5), and proceeds
inward. This process is typical of the Gautieriales (Fig. 63) and is widespread throughout the Lycoperdales. Kreisel (1967) has delineated several variations of the forate process, which are described below.

23

I\

!II

q
I.

I!
J,
I

I,

4. Aulaeate (Invers koralloid) (Fig. 1 0): This type occurs when plectobasidia differentiate on the surface of down-growing primordial tissue
which usually grows from the upper surface of the peridium. As maturation develops, small areas of tissue may break away to form locules.
Hymenial development could be characterized as centrifugal, since the
hymenium forms on the inner surfaces of the primordial tissue. Gleba
developing in this manner is the aulaeothecium of Kreisel (1969), and it
occurs in the Hymenogastrales and in Gastrosporium (Fig. 12) in the
Lycoperdales.
5. Pileate (Monopileat) (Fig. 1 E): A single stalk with the gleba arranged,
as in an embryonic agaric button, with gill-like tramal plates (Podaxales,
Podaxis, Fig. 65; Secotiaceae, Nivatogastrium, Fig. 71), or with the gleba
in the form of a cylinder of hymenial tissue, which differentiates around
the sterile central stalk, as in Dictyophora (Fig. 43), Mutinus (Fig. 47),
and in other members of the Phallaceae (Phallales).
6. Multipileate (Polypileat) (Fig. 1 F): A series of 3 or more individual
secondary outgrowths of sterile tissue which arise from a central stalk.
Each secondary stalk differentiates its own hymenial tissues, either in a
thin layer or as a concentrated glebal mass. This is a common type in the
C1athraceae of the Phallales, and it is well illustrated by Clathrus (Fig.
62), Linderia (Fig. 59), Anthurus (Fig. 60), an'd Aseroe (Fig. 53), to
mention a few.
Kreisel (1967) introduced several terms which describe various types
of forate development usually encountered within the Lycoperdales.
They are based primarily on macroscopic differences. The five types are
listed and described below and illustrated in Figures 2 and 3.

i'
"

,,'
,

,I
!

:I
,,

,,

1. Lycoperdoid (Fig. 2 A): With a sterile base and a gleba with forate
development and a continuous capillitium, which is nearly unbranched
to sparingly so or frequently branched. Calvatia and Lycoperdon are
examples.
2. Vascelloid (Fig. 2 B): A condition in which the gleba is separated from
the sterile subgleba by a well-developed diaphragm (Fig. 11). Vascellum
is an example.

"

'~

'~

, ,I
.,
,
I

3. Globaroid (Fig. 2 C): With sterile base reduced or absent, gleba with
some continuous capillitium , which is usually near the endoperidium.
The rest of the capillitium consists of discrete, tapering units (PI. VIII 0).
Some species of Bovista represent this type.
4. Bovistelloid (Fig. 3 A): With a sterile base and a gleba with forate
development and a capillitium of discrete, tapering units (PI. VIII 0).
This type is common in the genus Bovistella .

24

5. Bovistoid (Fig. 3 B): Sterile base is absent or reduced, gleba without

continuous capillitium near the endoperidium. Instead, the capillitium
is composed of discrete tapering units. Examples are also species in the
genus Bovista.

A.

B.

c.

Fig. 2. A. Lycoperdoid. B. Vascelloid. C. Globaroid.

Fig. 3. A. Bovistelloid . B. Bovistoid.

25

CHEMICAL REAGENTS
A complete discussion of chemical reagents is presented by Largent et
al. (1977), who cover the higher fungi and the agarics in particular. In this
section those reagents specifically used for Gasteromycetes are presented and discussed. Be aware that some of these chemicals are
dangerous and should be handled with care. They range from corrosive
to highly toxic if ingested. The formulae for the preparation of each
reagent and several of the most notable uses for them are presented.
More information can be obtained from the reference cited above.

Alcohol (ETOH)
Formula: 70% or 95% ethyl alcohol (ethanol) in water.
Use: The tissue of many of the Gasteromycetes is so constructed that it is
often resistant to wetting by water alone. One of the cheapest and
quickest wetting agents is ethyl alcohol. A portion of the gleba or a piece
of tissue is first soaked in alcohol. Only seconds are usually required with
the capillitium of Lycoperdon, for example, but the peridium of a
Crucibulum may take 2 or 3 minutes to be completely wetted. This
procedure is usually followed by adding a drop of water, and then the
desired mounting medium or micro-chemical reagent is used. I have
found it to be desirable to use a 70% ETOH solution with fresh or dried
immature gleba, wait several minutes, then add water followed by a very
small amount of cotton bhle or congo red. The choice of stllins depends
upon the tissue being examined and the purpose of the examination,
which are discussed under each reagent. In either case, the revival of
thin-walled basidia, generative hyphae, and immature spores is enhanced by this approach. It is also desirable to have one mount in water
in order to observe the color and characteristics of the unstained tissue.
The use of phase microscopy with a water mount will often reveal
basidia, generative hyphae, and much more detail than standard light

microscopy.

Congo Red: 1% aqueous solution

Formula: Dissolve 1 gm of congo red in 99 cc of water.
Use: Stains the walls of basidia and generative hyphae of immature
gleba, which are thin-walled and otherwise very hard to see. A small
amount of 1% congo red reagent can be added to sections mounted in
3% KOH (potassium hydroxide). A soft paper tissue is used as a wick on

26
the opposite side of the cover slip to draw the stain across the material
under the cover slip. The process of staining can then be observed as it
takes place. This works well for the basidia of Lycoperdon, Ca/vatia, and
Astraeus, for example.

Cotton Blue: 1% cotton blue in lactic acid/phenol

Formula: Dissolve 50 ml of 1% aqueous solution of aniline blue (cotton
blue) in 100 gm lactic acid, 100 gm phenol, 150 ml glycerine
(optional), and 50 ml of water. (See Largent et aI., 1977, for an
alternate formulation.)
Use: In Gasteromycetes true capillitium (PI. VII C & D) often has walls
which are clearly and strongly cyanophilic (blue-stained). One must be
certain that mature capillitium is examined. The gleba should be the dark
color typical of the older fruiting bodies to insure maturity. The walls of
paracapillitium do not stain or show very irregular staining in which a
septum or short length of the wall at the very most occasionally stains, as
in Plate VIII C. In both types of capillitium, casual blue staining of
incrusted material on or near the hyphal wall or the staining of hyphal
contents is not considered a positive cyanophilic reaction. The cell wall
must be uniformly blue-stained before it is considered to show a positive
reaction. The positive reaction is usually seen immediately through the
10-power objective when the slide is scanned, and it is easily confirmed
by observing the scattered unstained thin-walled generative hyphae. It is
desirable to select a fresh Bovista, Lycoperdon, or Ca/vatia, genera which
produce true capillitium, and to observe the immature white gleba with
subsequent observations as the gleba matures.
The spores of many taxa, such as Lycoperdon and Ca/vatia, are
cyanophilic. The reaction causes an intense or dark blue staining of the
exosporium in contrast to the inner wall layers.

Melzer's Reagent
Formula: 0.5 gm of iodine, 1.5 gm of potassium iodide, 20 cc of water, and
20 gm of chloral hydrate. The components dissolve better in
warm water.
Use: The uses here are much less than in other orders of Homobasidiomycetes such as the Agaricales and Aphyllophorales. Some secotioid
taxa within the Hymenogastrales (placed in the Russulales by some
authors) have amyloid or blue spore walls. In addition, the peridiallayers
of some species of Ca/vatia are dextrinoid, turning red to deep cherry
red. However, the reaction of many tissues has not been systematically

27

recorded. It is possible that additional dextrinoid or amyloid reactions

will be recorded in the future.

Potassium Hydroxide (KOH)

Formula: 3% aqueous solution.
Use: In Gasteromycetes no reactions of taxonomic significance have
been recorded. It is generally used as a mounting agent and helps,
somewhat, in restoring dried tissues to their full size. In most other
Basidiomycetes, a color change to yellow or red has proved to be
taxonomically significant.

Gum guaiac
Formula: 500 mg in 30 cc of 70% or 95% ETOH.
Use: Gum guaiac is a substrate for the laccase enzyme system possessed
by all fungi which degrade lignin. It is used as a spot test on the fresh
fruiting-body tissue and on cultures (anamorphs) of fungi. A positive
reaction is a change of the reagent to blue or blue-green. The reaction
usually takes place in less than one to several minutes. It is desirable to
allow at least an hour for a final reading when testing in culture dishes.
Since specific taxonomic groups of fungi, known as white rots, are able
to delignify wood, while others, known as brown rots, can only degrade
cellulose and hemicellulose, wider use of this reagent will undoubtedly
take place in the future. It must be realized that in a small percentage of
organisms, tyrosinase can give a false positive reaction, which is a weak to
strong blue to bluish-green color. It is, therefore, desirable to combine
this test with the use of syringaldazine, described below, which is specific
for the laccase enzyme complex. Gum guaiac must be made up fresh
every 4 to 5 weeks, and it is desirable to use a known positive-reacting
species as a control. Pure cultures of species of Lentinellus or Agrocybe
in the Agaricales yield strong positive reactions within 3 minutes.

Syringaldazine
Formula: 0.1% aqueous solution.
Use: Most commonly used jointly with gum guaiac as a spot test on fresh
tissue of fruiting bodies or on culture mats or mycelium in situ.
Syringaldazine yields a pink to bright pink positive test for the laccase
enzyme system within one to 20 minutes but most often in the first 3
minutes. The use, in conjunction with gum guaiac, allows one in most
cases to determine whether a given taxon has a tyrosinase system or the

28
important laccase enzyme system possessed by all fungi which delignify
wood (see gum guaiac above). Major groups of fungi, such as the
Sclerodermatales, have evolved as ectomycorrhizal symbionts (Miller,
1983) and are seldom capable of delignifying wood. Other orders, such
as the lycoperdales, have evolved as wood-rotters, and at least some taxa
cause white rots, while others cause brown rots; consequently, this test
has become important (Harkin et aI., 1974) in distinguishing one from the
other.

Cresyl Blue (= Brilliant Cresyl Blue)

Formula: 0.5-1.0% in water.
Use: This reagent is not commonly used in Gasteromycetes, and when it
is used it is applied to genera included in the Agaricoid Gasteromycetes.
The inner spore wall turns violet to reddish when placed in cresyl blue.
Such a reaction is called metachromatic and usually occurs in thickwalled spores. Secotium has metachromatic spores.

Schaeffer Reaction
Formula: Pure aniline and concentrated nitric acid. (POISONOUS)
Use: The Schaeffer reaction is used for specific Sections in the genus
Agaricus (Agaricales). A thin line of aniline is applied to the stipe or pileal
cuticle and is crossed with a thin line of nitric acid. A positive reaction
results in a bright orange to orange-red reaction at the point where the
two lines intersect. A positive Schaeffer's reaction occurs in the genus
Longula (page 110), which is an Agaricoid Gasteromycete. The two
chemicals aniline and nitric acid are poisonous to humans and should
not be touched or ingested and the fumes should not be breathed.
Aniline is light-sensitive and must be kept in a covered or opaque
bottle.

29

KEY TO THE ORDERS OF GASTEROMYCETES

1. Small (1-10 mm broad), cup- or urn-shaped, epigeous fruiting bodies
containing one to numerous oval to more commonly disc-shaped
peridioles, which are loose or embedded in a gelatinous matrix;
commonly gregarious or single, on wood or grass and plant material
in dung (Figs. 40-42) .............................. Nidulariales
1. Larger or differently constructed fruiting bodies, peridioles when
present with common walls (Fig. 37) or appearing like sand grains
(see Arachnion); on wood, ground, or various substrates ....... 2
2. Fruiting body hypogeous, contained in (see Protophallaceae) or
emerging from (becoming epigeous) a tough, flaccid, oval to
globose button, with stout rhizomorphs attached to the base;
hymenium slimy, olive, olive-gray, to olive-brown, often fetidsmelling with rod-shaped to narrowly elliptical spores and
located most commonly on an erect receptaculum which is single or has arms, arches, or is clathroid (Figs. 43-62) .. Phallales
2. Fruiting body agaricoid, secotioid, earthstar-like, or with an oval,
pear-shaped to irregular peridium, stalked or unstalked; gleba at
maturity with hymenium and spores in gel-filled locules, lining
tramal plates in empty locules, in a powdery mass, or on gussets
(Fig. 66) ............................................ . : ... 3

3. Fruiting body hypogeous or epigeous at maturity, oval with a lacunose or chambered, gel-filled gleba usually with a wet or glistening
appearance; rubbery and tough when fresh, very hard when dry
..........................................................

3. Fruiting body shaped differently or with a differently constructed

gleba or a sterile subgleba ................................... 5
4. Gleba gel-filled, gel latex-like, white to transparent; spores
thick-walled, spinose to reticulate ............ Leucogastrales
4. Gleba gel-filled, gel orange, changing to muddy red when
bruised and cut, or black and unchanging; spores thick-walled,
smooth .................................. Melanogastrales
5. Mature gleba powdery, with subglobose to globose, warted to reticulate spores surrounded by capillitium and/or paracapillitium or
without a capillitium; fruiting body stalked (Fig. 22) with a sterile
subgleba (Fig. 5) or completely composed of gleba ............ 6
5. Mature gleba brittle, tough but not powdery; spores variable, capillirlum absent ................................................ 8

30

6. Fruiting body with a well-developed stalk composed of longitudinally arranged hyphae, woody at maturity; capillitium and/or
paracapillitium present (Figs. 22-34) .......... Tulostomatales
6. Fruiting body may have a sterile, usually chambered, base but
lacks a stalk ............................................ 7

7. Gleba white at first, with capillitium or paracapillitium; peridioles

absent; spores globose, with blunt warts (Figs. 1-21) ............ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Lycoperdales

7. Gleba purple, purple-brown, to brown when immature, with obscure to clearly visible peridioles or chambers (Figs. 35 & 37); spores
globose, with spines, warts, or reticulate ornamentation (Figs. 35-39)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Sclerodermatales
8. Fruiting body hypogeous, lacunose, ovoid, 1-6 cm broad, with a
finely branched columella when present, and thin or thick peridium (Fig. 63); spores orthotropic, longitudinally ridged, yellowbrown, greenish-brown, to rusty brown ......... Gautieriales
8. Fruiting body epigeous and hypogeous, lacunose to secotioid or
boletinoid; spores variable but not longitudinally ridged ... 9
9. Spores globose with amyloid warts or reticulation; some sporocarps
have heteromerous trama, and some have latex in the trama; fruiting
body lacunose, columellate to secotioid ....................... .
.................................... Russulales (not included)
9. Spores variously shaped, smooth to ornamented, non-amyloid;
heteromerous trama and latex absent; fruiting body lacunose,
columellate to secotioid or agaricoid ........................ 10
10. Fruiting body usually hypogeous, and in some cas~s epigeous at
maturity; gleba loculate, with or without a columella; spores
statismosporic, ovoid to elliptical, smooth, echinulate, to warted
(PI. IV B), light to dark brown; predominantly ectomycorrhizal
. . . . . . . . . . . . . . . . . .. Hymenogastrales (only 1 genus included)
10. Fruiting body usually epigeous, sometimes hypogeous, largely
agaricoid to boletinoid; with a stipe-columella and with contorted lamellae or irregularly lacunose tramal plates; spores and
other features characteristically like those of the related agaric
families, statismosporic and/or orthotropic; decomposers or
mycorrhizal (Figs. 65-75) .......... Agaricoid Gasteromycetes

31

LYCOPERDALES Clements
General: The true puffballs and earthstars have an epigeous or rarely a

partially hypogeous habit, are most commonly oval to pear-shaped, and

have a several-layered peridium. A two-layered peridium composed of
exo- and endoperidium is typical, but both three and four peridiallayers
are typical of some species or genera. The exoperidium is typically
lacerate and caducous, or splits and is shed, as the fruiting body matures.
The immature gleba is white, homogeneous, and composed of minute
but not clearly delineated locules. As the maturing process takes place,
the young white gleba changes color as the evanescent, hyaline
plectobasidia collapse and give way to the developing capillitium,
paracapillitium, or a combination of both. At maturity the gleba is
powdery and characteristically colored as a result of the pigmentation
acquired by the mature capillitium and spores. A chambered subgleba
(Fig. 5) or a columella as shown in Geastrum (Fig. 16B) may be present,
or the gleba may occupy the entire interior of the fruiting body. The
subgleba is often chambered, either white or characteristically colored,
but does not change color as the fruiting body matures. A subgleba is not
present in Ca/vatia gigantea, and for this reason it is placed by some
authors in the genus Langermannia Rostkovius. The spores are globose
to subglobose or occasionally broadly elliptical, becoming warted to
echinulate at maturity. They are predominantly wind-disseminated. The
thick-walled, ornamented, exosporium contains the pigmentation,
which along with that of the mature thick-walled capillitium gives the
gleba its color. The species are distributed among four families which are
distinguished primarily by the mode of development of the fruiting
body. Special emphasis is placed on the development of the peridial
layers, the presence of a stoma, and the morphology of the capillitium.
Ecology: The order is distributed worldwide. A small number of taxa is

found in the tundra and taiga biomes. Larger numbers occur in the
boreal or conifer biome, and the order is well represented in temperate,
tropical, savanna, and desert biomes. The vast majority of species are
decomposers of plant material such as wood, leaf mulch, moss, and
unincorporated organic matter. There are several genera, including
Radiigera and Geastrum, which often appear in specific higher-plant
associations. Their specific role has not been determined, nor have they
been pure-cultured and examined in vitro as yet.

32

Key to the Families of the Lycoperdales

1. Fruiting body small (0.5-2 cm broad), ovoid; gleba with minute peridioles which resemble sand grains .............. 4. Arachniaceae
1. Fruiting body larger; gleba without peridioles, powdery at maturity
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2
2. Exoperidium splitting to form an earthstar or remaining
unopened (Figs. 18 & 19), having a prominent, central columella
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. Geastraceae

2. Not forming an earthstar and not with a prominent, central

columella .............................................. 3

3. Fruiting body 1-3 cm high, hypogeous at first, exoperidium with

stellate calcium oxalate crystals, endoperidium a thick gelatinous
layer (Fig. 12) ............................. 3. Gastrosporiaceae
3. Fruiting body larger or with differently constructed exo- and endoperidium ..................................... . ............ 4

4. Fruiting body with trabeculae and a central core (Fig. 15), or the
exoperidium splitting at maturity to form erect lobes (Fig. 14);
spores elliptical, verrucose to warted with a basal collar, pedicel
absent ................................ 5. Mesophelliaceae
4. Fruiting body without trabeculae, nor is there the distinctively
splitting exoperidium; spores subglobose to globose with a short
or long pedicel (PI. II A-F); capillitium or paracapillitium usually
present and prominent ................... 1. Lycoperdaceae

1. Lycoperdaceae Smarda
Fruiting body epigeous or becoming so at maturity, ovoid, elliptical,
pyriform, 0.5-140 cm broad, exoperidium sloughing to reveal a wellformed ostiole or a lacerate opening for spore discharge, or lacking a
discrete opening and the peridium simply wearing away at maturity;
gleba of capillitium, paracapillitium, or both, not in locules or peridioles,
with or occasionally without a chambered, sterile subgleba; spores 3-17
J-Lm, globose to subglobose, with minute to large, blunt warts (PI. II),
sometimes with a persistent, short or long pedicel. Rhizomorphs are
commonly present and often extensive. Cosmopolitan decomposers of
plant debris.

33

Key to the Genera of the Lycoperdaceae

I. Subgleba (sterile base) absent ............................... 2
1. Subgleba usually well developed, often chambered ............ 6
2. Very large fruiting body (10-140 cm broad), exoperidium smooth,
gleba olive-brown at maturity. . . . . . . . . . . . . . .. Langermannia
2. Usually much smaller (if large, peridium is not smooth) ..... 3
3. Fruiting body small, 5-40 mm broad, covered with a sand case over
the lower half (Fig. 13) ...................................... 4
3. Fruiting body larger, (1.5-) 3-30 cm broad, sand case absent ... " 5
4. Endoperidium at maturity with a well-formed pore .. Disciseda
4. Endoperidium at maturity splitting irregularly but developing no
pore ............................................ Abstoma
5. Fruiting body 1-8 cm broad, detached from the substrate at maturity;
capillitium of discrete units (PI. VIII 0) ................... Bovista
5. Fruiting body 3-30 cm broad, often persistent at maturity; capillitium
of infrequently branched to unbranched units (PI. VII 0) .. Calvatia
6. Only paracapillitium present, of thin-walled, hyaline units with
walls unstained in cotton blue (contents may stain) ......... 7
6. Capillitium present, of thick-walled units some with pores or
slits, others without, deep blue walls in cotton blue ........ 8
7. Well-developed membrane separates the chambered, sterile base
from the gleba ..................................... Vascellum
7. Membrane absent ................................ Morganella
8. Exoperidium sloughs off to reveal a smooth endoperidium with a
well-developed pore, or the capillitium contains discrete units
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 9
8. Exoperidium mayor may not slough off, usually lacerate, pore
absent, endoperidium splits at maturity; capillitium spiny (PI. VII
A) to branched, or unbranched, or with discrete units ..... 10
9. A well-developed pore is formed at maturity; capillitium of
branched, interwoven hyphae, without tapered tips .. Lycoperdon
9. Pore present or absent; capillitium of discrete units culminating in
narrow, tapered tips (PI. VIIIO) ...................... Bovistella

34
10. Fruiting body 6-24 cm broad, with a checkered peridium at
maturity; capillitium composed of intricate branches which form
long spines (PI. V E & VII A) .................. Mycenastrum
10. Fruiting body not as above; capillitium not spinose ....... 11
11. Peridium thin or thick, variously colored, smooth or with fibrils,
connivent fibrils, or pyramidal warts (Fig. 4); capillitium infrequently
to frequently branched (see also choice 5) .............. Calvatia
11. Peridium thick, dull white with low, truncate warts; capillitium of
usually tapered, separate units; western North America ......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Calbovista
Abstoma Cunn.

Fruiting body up to 4 cm broad, depressed-ovoid to ovoid, exoperidium a thick, firm sand case, dull purple, breaking away to reveal a thin
endoperidium which is membranous, purple to purple-black, without a
pore but splitting irregularly at the apex. Subgleba absent: Capillitium
smooth, deep brown. Spores globose, 8-17 J,Lm, deeply reticulate (1.5-2
J,Lm high), dark brown.
Bovista Pers.

PI. II D & E; VIII D; Fig. 9

Fruiting body oval to globose, 1.2-8 cm broad, exoperidium white or

light colored, fragile, sloughing off when mature to reveal a tough,
smooth endoperidium with an apical pore. Sterile base absent. Capillitium of discrete, tapering, regularly branched units, ending in fine,
narrow tips. Spores brown to purple-brown, globose to subglobose,
3.5-7 J,Lm with a short or long pedicel.
Bovistella Morgan

Fig. 10

Fruiting body subglobose, 0.5-8 cm broad, exoperidium of soft, often

connivent fibrils (Fig. 4), usually white, irregularly sloughing off .to
reveal a thin, tough, metallic or silvery endoperidium with an apical
pore or irregular, often lacerate slit. Subgleba well developed, cellular,
with a zone which delimits the subgleba from the gleba. Capillitium of
discrete units (PI. VIII D) with regular, tapering branches culminating in
blunt, narrow tips. Spores brown, globose, 3.5-5.5 J,Lm with persistent
pedicels.

Calvatia Fr.

PI. I A; II A, B, C; VI A; Fig. 1 B

Fruiting body ovoid to pear-shaped, 3-30 cm broad, without a stoma,

exoperidium sloughing off at maturity, endoperidium becoming irre-

35

gularly lacerate at maturity exposing the gleba, with or without a

persistent sterile base. Capillitium infrequently to frequently branched,
not tapering. Spores 4-5.5 (-7.5) p,m, globose or broadly elliptical, rough
or smooth.

Calboll;sta Morse

PI. I D

Fruiting body ovoid, exoperidium of blunt, truncate warts (Fig. 4 D),

6-15 cm broad, no stoma, endoperidium irregularly lacerate at maturity
with a well-developed sterile base. Capillitium of separate units, usually
tapered. Spores 3-5 p,m, globose, minutely warted with a short pedicel,
brown.

D;sciseda Czern.

Fig. 13

Fruiting body ovoid to ovoid-depressed, 0.5-3.5 cm broad, exoperidium

a thick, brittle, sand case, breaking away over the top half to reveal a
membranous endoperidium with a well-formed pore. Sterile base
(subgleba) absent. Capillitium of short-branched, non-pitted elements.
Spores dark brown, globose, 4-5.3 p,m, warted, a short pedicel sometimes present.

Langermann;a Rostk.

PI. I B

Fruiting body oval to slightly depressed, 10-50 (-140) cm broad, exoperidium white, smooth, dry, of one layer, not a pseudoparenchyma,
becoming water-soaked, dingy, and irregularly sloughing off to reveal
the endoperidium which is also white to gray-brown and dingy in age,
cracking irregularly to expose the gleba at maturity. Gleba white to
yellow, powdery and olive-brown at maturity. Subgleba absent. Capillitium thick-walled, branched with pores and septa, yellow-brown.
Spores globose, 4-6 p,m, with low, small warts, olive-brown at maturity.
(Calvatia of many authors.) Monotypic, L. gigantea (Batsch ex Pers.)
Rostk.

Lycoperdon Tourn. ex Pers.

PI. V A & B; VII D; Figs. 5-8

Fruiting body 0.5-5 (-7) cm broad, globose to pear-shaped, exoperidium smooth or with fibrils to long spines, white to variously colored,
endoperidium membranous with a pore and well-developed stoma at
maturity. Subgleba mostly well developed, no diaphragm, chambered,
usually with a rudimentary to well-developed pseudocolumella. Capillitium branched, interwoven, well developed. Spores 3-5.5 (-6) p,m,
globose or broadly elliptical, minutely warted to warted, with or
without pedicel.

36

Morganella Zeller
Fruiting body 1-2 cm broad, exoperidium granular or with colored
fibrils; endoperidium very thin with a pore at maturity. Subgleba
chambered or interwoven with a delimiting membrane. Capillitium
absent, paracapillitium present. Spores 3.8-5 J.'m, globose, thick-walled,
warted, brown.

Mycenastrum Desv.

PI. I C, V E & VII A

Fruiting body depressed-ovoid to ovoid, 6-24 cm broad, exoperidium

floccose, thin, soon darkening with a typically checkered pattern;
endoperidium dark brown, membranous, splitting irregularly in large
flaps and pieces. Subgleba absent. Capillitium composed o( intricate
branches with multiple terminal spines. Spores brown, globose, 8-13
J.'m, reticulate with irregular, coarse warts.

Vascellum Smarda

PI. VII I C, Fig. 11

Fruiting body 1-6 cm broad, exoperidium of fibrils, sloughing at

maturity, pore develops in endoperidium at maturity, sometimes surrounded by mycosclerids (Smith, 1974). Membrane separates chambered subgleba from gleba (diaphragm). Capillitium absent, paracapillitium present. Spores 3-4.5 (-5) J.'m, globose, with minute to well-formed
warts, light red-brown.

2. Geastraceae Corda
Fruiting body 1-5 (-12) cm broad, ovoid, bluntly pointed, to acuminate,
smooth to roughened, with three peridial layers; exoperidium splitting
from apex to form 4-12 rays which open in a characteristic manner for
each species but usually in a stellate pattern; mesoperidium usually
inseparable from the exoperidium and both layers forming the d'laracteristic rays, which following their maturity reveal the papery thin,
usually tough, glabrous to roughened endoperidium, which develops
an ostiole and a characteristic delimited peristome or disc and is sessile
or pedicellate (in Geastrum, Fig. 18), or the endoperidium does not split
but just erodes away to expose the gleba (in Radiigera, Fig. 16 A). Gleba
develops radially from a central, ovoid or knob-like, white pseudo
columella (Fig. 16 A) and is brown to olive-brown at maturity. Capillitium abundant, pigmented, mostly unbranched, cylindric to irregular,
thick-walled. Spores 2.5-10 J.'m, globose to subglobose, thick-walled,
with individual warts or blunt spines, and a very short or inconspicuous
pedicel. Cosmopolitan decomposers.

37

Geastrum Pers.

Figs. 16 B-21

(Characters of the family)

Radiigera Zeller

Fig. 16 A

Fruiting body (morphologically a Geastrum without a stoma) hypogeous or partially epigeous when mature, 2-6.5 cm broad, oval to
oval-depressed, exoperidium indehiscent, brown, roughened with
impregnated sand grains and debris, mesoperidium thick, fleshy, with a
thin endoperidium surrounding the gleba which is formed radially
from a central, ovoid or knob-like, white pseudocolumella (Fig. 16 A).
m abundant, pigmented, largely unbranched, thick-walled.
Spores (4.5-) 5-5.5 ILm, globose to subglobose, thick-walled, with individual blunt warts and a short, inconspicuous pedicel.

3. Gastrosporiaceae Pilat
Fruiting body 1-3 cm high, 0.8-1.7 cm wide, ovoid to elliptical, chalky
white, mealy, hypogeous to emergent at maturity, spore release occurs
through an eroded apex (Fig. 12), exoperidium of interwoven, clamped
hyphae which are interspersed with calcium deposits in the form of
stellate crystals; endoperidium a thick, gelatinized layer of interwoven
hyphae with clamp connections. Paracapillitium of hyaline, branched
hyphae with numerous clamp connections. Spores 3.5-6 x 3-5 ILm,
globose to subglobose, with scattered to dense warts and an occasional
short pedicel. Known from grasslands in Europe and North America.

Gastrosporium Mattir.

PI. III B; Fig. 12

(Characters of the family)

4. Arachniaceae Moravec
Fruiting body 0.5-2 cm broad, globose to ovoid, often lumpy and
irregular in shape, epigeous, gregarious, dry, dull white, sometimes
buff at maturity, with stout rhizomorphs at the base; peridium thin,
single, breaking up at maturity. Sterile base absent. Gleba develops as a
regular hymenium lining the inside of minute chambers or peridioles,
which separate at maturity forming loose, ash-gray particles resembling
small grains of sand. Capillitium absent. Spores oval to short elliptical,
smooth, 4.2-6 (-10) x 3.5-5 (-6) ILm, thick-walled with a pedicel, 0.5-1.5
ILm long. Known from the southeastern United States.

38

Arachnion Schw.
(Characters of the family)

5. Mesophelliaceae (Cunn.) Julich

Fruiting body epigeous or becoming so at maturity, 1-5 cm broad,
peridium 3-layered, thick, cemented together with sand particles,
tough to brittle at maturity and either splitting or rupturing at maturity.
Gleba olivaceous to yellow-brown, occupying the entire center of the
fruiting body or located between the peridium and a sterile, central
core and held in place by trabeculae (Fig. 15). Capillitium hyaline,
septate, sometimes in parallel bundles. Spores elliptical to narrowly
elliptical, 7-18 x 3-10 Ilm, thick- to thin-walled, warted, yellow-brown to
brown with a basal collar with or without a gelatinous exosporium.
Probably mycorrhizal, known only from Australia (Cunningham, 1944).

Mesophellia Berk.

Fig. 15

Fruiting body 1-5 cm broad, oval to oval-depressed; exoperidium 1-3

cm thick, brittle, cemented together with fine sand and debris, mesoand endoperidium present and well developed. All three layers splitting at maturity by irregular weathering. Gleba olivaceous to avellanous, located between the endoperidium and a solid, firm central core
(Fig. 15) held in position by trabeculae. Capillitium hyaline, composed
of septate hyphae in parallel bundles. Spores elliptical to narrowly
elliptical, 7-14 x 3-6 Ilm, minutely to coarsely warted, base with a
conspicuous collar, yellow-brown to brown, or olivaceous-brown.

Castoreum Cooke & Massee

Fig. 14

Fruiting body subglobose, 1-3.5 cm broad, exoperidium up to 2 mm

thick, composed of coarsely interwoven hyphae with loosely adhering
sand particles, brown; endoperidium 1-2 mm thick, tough, leathery.
Both layers rupturing at apex into several to many unequal lobes. Gleba
pale olivaceous to yellow-brown. Subgleba absent. Capillitium hyaline,
septate. Spores elliptical, 7-18 x 4.5-10 Ilm, thick-walled, surface verrucose with irregular warts, a gelatinous exosporium, and a basal
collar.

39

A.

B.

'.. '

c.

D.

E.

F.

Fig. 4. Exoperidial fibrils and warts. A. Tapering fibrils. B. Connivent

fibrils . C. Double-connivent fibrils. D. Truncate pyramidal warts.
E. Pyramid-shaped warts. F. Truncate warts with apical tufts of fibrils.

40
A
~-B

......-:

o
en

....--- g

pc
B

sub

. ... . ... . .
.
. . . . .,
.' "

.-

...

...

Fig. 5. Lycoperdon per/atum. Middle-aged fruiting body with exoperidium (p) composed of conical warts (B) which are deciduous and leave
shallow pits (A) when dislodged. The gleba (g) has irregular chambers, a
pseudocolumella (pc) in the center, and a chambered subgleba (sub).
When the exoperidium (p) sloughs away in age a preformed ostiole (0)
will be visible on the apex of the endoperidium (en).

41

,....-- 0

Fig. 6. Lycoperdon marginatum

-en

with a characteristically loose

exoperidium (p), ostiole (0),
and tough endoperidium (en).

. -. ....

'::

..... : . .
.....~~.",
.' .::"':~,.,
..
....
,..
.. . . .
'.
. '...
.
..
.' ' ...

'

--en

p
-p

Fig. 7. Lycoperdon pulcherrimum

with the distinctive fringe of
strigose hairs the exoperidium (p)
just above the sterile base. The
ostiole (0) is seen along with the
endoperidium (en).

0-

en

Fig. 8. Lycoperdon mammaeforme with the

unique cottony patches
formed as the exoperidlum (p) breaks up, revealing the ostiole (0)
and endoperidium (en).
Rhizomorphs (rh) proliferate from the steri Ie
base.

it:? P

')---rh

42

en

.. ,

. .'

. ..'. . .
."
.
. " ..

. ..

..

p
. ..

en -

---'-0-

..
!. :

p
>-----rh
Fig. 9. Bovista pi/a. Progressive stages of maturation from young stage in
lower left, sloughing of exoperidium (p) in center and the progressive
splitting of the endoperidium (en), top, to reveal the gleba (g). Fine
rhizomorphs (rh) are present at the base.

.: . , ..
. . .
'.

10

en

.
. '. .:, . .
. .. .. .'. '
.
"oj
,.:',.!;.
......,'; .....'.: ......:r".....
... " ,_,
.

"

.'

.'

~.~.,

:.

..

'.

...

'"

."!'}..

..

.~.

:
"
..;-"; ,
...::":;.~
:.~:,
,. . "~'.
'~""~:'"
..........
:

i;.

j
",.::.~', ':,,!.";~:
..,/
.., ....
. '.'''''
. Ar..... ,' -,
. . _,'
'"
~r.."\.: ,".:'
.
~-:':.,;.to.,\ ..:. / .... . . ~-,,'
.\.~
;

" ')' . ". . ',

., I
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I
.

.....
.
.
.. ,.'.:........

.. . . . . . . ,~..j,:....'

.. ' ";' ,

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. :.~:

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,-

:. ,....

"

.... .' ".,.

'" "~.::"'.

or,
"
'.",
',_'
..\..
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:
.....
':; :: -'."
. ~'.:
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-....
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.

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.,r ..
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:::.;.~~"

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........

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..... ,';!'..... ."

,.:...~

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....:,..

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...'r',
:.:.... ,::
... . ." .
.

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...... ,..

A',
.." ':'"
..........
'

. .(".

.~'

".:"

',; '.'- .'.:." :.:.....": .. :.'

. .i{~ ,; . .;,.:.:..' ,:'. ,.:!-.: " ::. :.~:.' ,
''',~
~ ....' "lo: ....... ,> . . . . . . .
'
~:.~.~.>!.,:-::~;.\: t~::i:.:::;"':::::' " .
:;.~/,~"'.;"

::.....
'.:>...... ......., .... ......... .....
.. '..
. ... .
. .. ...

. .

'

'

'" "'.' .

..
. ".
' , ; r ...
,;
..
....'. -. .'..... . .. .
.
'.

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..,

.'

"

A,

'.'

"

,.

'

'. "

':'"
,

.'

, .

' .
,.

. .
:.

g
Fig. 10. Bovistella radicata, showing
(lower right) the subgleba (sub), experidium (p), endoperidium (en),
and gleba (g). A lacerate ostiole (10) is
formed at maturity after the thin, fragile, white ' peridium (p) sloughs off.

'r--

H--en

'--sub

43

10
. .... '. ..

'
... ".

. '? . . .

.....:

.'..

. '~"

,"

.~

"',
"

. ..,

.
"

,.. -, ,
...... : ....,,:;.
. '... . .
".~.. ,'~'

. ".
..Iot"'' ' \"'".
.
. (ilP:' ... "
. " ~ -:~,... ~:~~,~,":' .'.:' . .
....... ,.. -.....
:,

",

,-".,,,,,,~;,.

.' .

. :

.
.

..

"

.~.,:.~
":'1...
,~

.'

. . ..

'.
i!.

'.

~:',

A.",;;.

.;2'

.'.

.
.'

. .'",",

.'

. ,:,i-;iif.t'

, ,.',
.

g--l

."

.::

~;.

. '.

..

'".;,
~.
"

.. ,;:

..+

" f.l'

.'~'.

..',' .- ' ......

'"#<'

' .

..'

.
..
.

,"

.."

ft-g

...,.
;:'

dia

'1--- P

t--sub

Fig. 11. Vascellum pratense. Section at right shows the diaphragm (dia),
unique to the genus, which separates the subgleba (sub) from the gleba
(g), which is surrounded by the peridium (p). A lacerate ostiole (10)
allows the spores to escape.

Fig. 12. Gastrosporium simplex.

Young fruiting body on left and
mature stages on right, showing
appearance following abrasion
and weathering of the apex of
the exoperidium (p) to
reveal the gleba
(g) at the surface of
the soil. Note the
stout rhizomorphs
(rh) which wither
away at maturity.

rh

44
..;;;......",....--- 0

en Fig. 13. Disciseda candida. At maturity the

exoperidium, which is
a sand case (sc), re- .
mains persistent over
the base, and endoperidium (en) and ostiole (0) at apex are exposed.
"

sc

Fig. 14. Castoreum cretaceum has a distinctive

coronate splitting of
the exoperidium (p) at
maturity to reveal the
gleba (g).

..
... .

. :. . ..:
.. .
. .. :.. \ ..., . .. ::....
.
.

.'

'

,"

,-

p
tr

g
core
en

.'. . . '
,

.' :
:. .

.. .

~----p

"

:;i~

"
,".

-"'J.

..

.'
;

".~.,.

~~

.,

.,''/i;~~:;~:~
./
... ;:~i... :
.
," .

,',

",. '.

.o'

,",'"
' .'.:1-'"
,

' ,'

...
,.

.:.~

~'f..'

. '. :.

....,

, ',

'

,
,

f\

"
"}o,'

'.'

'

' - - core

Fig. 15. Mesophellia in cross section, with a young section, lower right,
showing the trabeculae (tr) which connect the proteinaceous core
(core) to the tough exoperidium (p). The upper specimen has lost most
of the gleba (g).

A.

~---pe

Fig. 16 A. Radiigera fuscogleba, left, sectioned to show

r ----'
the three-layered peridium
me---~:
composed of an exoperidium
P _ _ _---i
(p), mesoperidium (me), and
endoperidium (en). The
B.
prominent pseudocolumella
(pc) is surrounded by radiating tramal plates that form
the gleba (g); however, no ostiole is formed. Geastrum, B. right, is
sectioned to show the ostiole (0), peristome (ps), and endoperidium (en).
The ray (r) is most often composed of the exoperidium (p) and mesoperidium (me). A pedicel (pe) is sometimes present and supports the pseudocolumella (pc) and radiating gleba (g) characteristically formed in the
Geastraceae. Compare Astraeus (Fig. 36), which has no columella.
0----

en--

Fig. 17. Geastrum fornicatum.

co ---....;:
pe - - - - : : :

At maturity, the mesoperidium (me) evaginates from the

exoperidium (p), which is a
membranous sand case (sc),
and is held tightly by the tips
of the arched rays (r), called
fornicate rays. This elevates
the endoperidium (en), ostiole (0), and pedicel (pe).

me ------,

... . .
sc

.'

. .r

-.;...,.,;4,
. ,....

..

'

'-

-. - ~

\.... - - .

....

" ..

' .

46
f
ps

en
::....-----_- ap
'-------co
-l.'.~,',~. .t:.:,::::-:~_-;.._----- pe
. ..
.

'

. .- .

::..,...---- r

.. ... ...
.

Fig. 18. Geastrum bryantii. The endoperidium (en) has a fimbriate (f)
peristome (ps) and a ventral apophysis (ap) with a narrow, hanging collar
(co), supported on a pedicel (pe) and seated on recurved rays (r).
P---- sul /

---ps
";':".,..-- en

sui
~ps

ap ------=
pe --""'--.:;::;;:r;;
..
.

Fig. 19. Geastrum pectinatum. The peristome (ps) is sulcate (sui). The
striate-wrinkled apophysis (ap) is supported by a narrow pedicel (pe).
The exoperidium and mesoperidium form the recurved rays (r).

47
ps
~---en

~--~

sr

me

----,.<..,...".~

..
",

...

" ;,,'

. ': :
",

Fig. 20. Geastrum saccatum. The saccate rays (sr) become revolute (r),
cracking to form a shallow bowl in which the endoperidium (en) is
seated directly on the mesoperidium (me). The peristome (ps) is
irregularly wrinkled.
o

-~......"..--

ps

en

--me
sr

.,-o-f---

me

Fig. 21. Geastrum triplex. The saccate rays (sr) become revolute, with
the mesoperidium (me) breaking completely apart, leaving the endoperidium (en) in a bowl. A delimited fimbriate (f) peristome (ps)
surrounds the ostiole (0). The rays (r) have long acuminate tips which
resulted when they split from the characteristically acuminately formed
button.

48

TULOSTOMATALES Demoulin
General: The stalked puffballs are usually epigeous at maturity, although they may spend the early developmental period in hypogeous
situations. The oval or oval-depressed gleba develops at the apex ora
well-developed stalk (Fig. 22), which may develop from a persistent
volva (Fig. 24L a mycelial pad (Fig. 22), or a cluster of rhizomorphs (Fig.
23). The stalk is most often composed of longitudinally arranged,
thick-walled, filamentous, cuticular hyphae with a central core of
interwoven hyphae, or in Ca/ostoma the stalk is glutinous (Fig. 31). (In
contrast, the Lycoperdales have a chambered, sterile subgleba or a
small columella or are sessile.) The young, white gleba is composed of
plectobasidia, which collapse following meiosis and spore formation.
The development of capillitium (PI. VIII A), paracapillitium, a combination of both, or the development of elaters (PI. VI C & D) commences as
the basidiospores mature and change to a characteristic glebal color for
a species, as in the Lycoperdales. The mature gleba is powdery, and the
basidiospores are warted and globose to broadly elliptical and are
wind-disseminated.
Ecology: The species are rare in the tundra and taiga and more frequent
in boreat temperate, and tropical biomes. The taxa are placed in two
families, the Tulostomataceae Fischer and the Calostomataceae Pilat.
The Tulostomataceae reach their full development in xeric microhabitats, savannas, and deserts, where their hypogeous early development
provides them with an adaptive advantage. The Calostomataceae are
found in warm-temperate and tropical ~iomes. The species all appear
to be decomposers of roots, buried wood, and other organic matter of
plant origin.

Key to the Families of the Tulostomatales

1. Fruiting body with a cottony, viscid to gelatinous exoperidium, a
white inner spore sac suspended from the peristome, and with buff
spores at maturity; stalk fluted to pitted, dry, or covered with a
viscid to a gelatinous layer, capillitium absent, paracapillitium
absent at maturity (Figs. 31-34) .............. 2. Calostomataceae
1. Fruiting body dry, opening by a pore, slit, lacerate peridium, or
circumscissile dehiscence; capillitium, paracapillitium, or elaters
present (Figs. 22-30) ....................... 1. Tulostomataceae

49

1. Tulostomataceae Fischer
Fruiting body epigeous at maturity, oval to oval-depressed, dull white
to brown, supported by a well-developed, non-gelatinous stalk, volva
often present and persistent. Peridium of two to three layers, mostly
non-gelatinous. Gleba white at first, with plectobasidia which collapse
as the capillitium and/or paracapillitium develops, elaters present in
two genera. Spores 2-9 1Lm, globose to subglobose, nearly smooth to
warted, spiny, or reticulate, yellow-brown to brown. Clamp connections present. Cosmopolitan, decomposers of plant litter.

Key to the Genera of the Tulostomataceae

1. Volva present and usually well developed; capillitium or paracapillitium present, sometimes elaters present also (see also Schizostoma)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 5
1. Volva absent; capillitium present or absent, paracapillitium someti mes present, elaters absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2
2. Stipe nearly equal, narrow (2-5 mm thick), inserted in oval to
oval-depressed spore case (Figs. 22,23,25); at maturity either an
ostiole is formed or the entire peridium splits and recurves . 3
2. Stipe rather thick, often shaggy, if inserted 15-20 mm thick, splitting at apex but peridium not recurving, pore and percurrent
columella absent ....................................... 4

3. Ostiole present and sometimes a peristome formed at maturity

(Figs. 22 & 23) ..................................... Tulostoma
3. Peridium splitting and recurving at maturity (Fig. 25) ............ .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Schizostoma
4. Peridium scaly or with pyramidal warts, fruiting body superficially resembles Podaxis but without a percurrent columella;
in deserts and sand dunes; disagreeable odor when fresh (Fig.
27) ........................................... Phellorinia
4. Peridium smooth or not scaly; stipe with shaggy, white fibrils,
not resembling Podaxis; very rare, fruiting among spent tan
bark; no odor (Fig. 26) ........................... Queletia
5. Fruiting body woody, long-stalked; capillitium composed of elaters
(Figs. 28 & 29) .............................................. 6
5. Fruiting body long- or short-stalked; elaters absent, capillitium or
paracapillitium present ..................................... 7

50

6. Fruiting body oval to depressed; peridium splitting by circumscissile dehiscence (Fig. 29) ....................... Battarrea
6. Fruiting body oval to depressed; peridium persistent, developing multiple ostioles at maturity (Fig. 28) ........ Battarreoides
7. Fruiting body large (head 5-13 cm broad); volva well developed,
persistent, 3-11 cm tall; stalk smooth to lacerate and appearing
twisted; gleba cellular, pore absent; odor of decaying fish (Fig. 30)

. . . . . . . . . . ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Dictyocephalos
7. Fruiting body small (head 2-3.4 cm broad); volva very small (4-15
mm tall) and inconspicuous; stalk longitudinally ridged; gleba not
cellular, pore present; no distinctive odor (Fig. 24) .. Chlamydopus

Battarrea Per-so

PI. VI C & D; Fig. 29

Fruiting body epigeous at maturity, 10-45 (-55) cm tall, with an ovaldepressed, disc-like, smooth, white head, 2-8 em broad by 1.5-4 cm
thick, with remains of endoperidium over the apex, dehiscence circumscissile at maturity (Fig. 29). Originating 5-20 cm below ground
from a persistent, woody, several-layered volva, 3-16 cm broad. Stalk
8-40 cm tall, 2-4 cm thick, woody, equal or tapering toward base, rough,
scaly at maturity. Gleba dark brown, powdery at maturity.
m of
elaters varying in length (usually under 80 p,m long) with the species.
Spores 5-7 x 4.5-6.5 p,m, globose to subglobose, thick-walled, warted or
partially reticulate, with a very short pedicel, brown. Decomposers of
the roots of desert plants (e. g. Atriplex, Artemisia, and Purshia).

Battarreoides Herrera

Fig. 28

Fruiting body epigeous at maturity, 11-21 cm tall, with an ovaldepressed, smooth, white head, 18-35 mm broad by 9-15 mm thick,
exoperidium absent, dehiscence by many pores over the surface (Fig.
28). Originating 5-10 cm below ground from a persistent, woody volva,
11-18 mm wide and 13-20 mm high. Stalk 10-18 cm tall by 4-9 mm thick,
woody, tapering toward base, rough, scaly at maturity. Gleba dark
brown, powdery at maturity. Capillitium of elaters (62-) 80-130 x 4-6 p,m,
occasionally branched. Spores 4-5 (-8) p,m, globose to ovoid, densely
warted, with a very short pedicel, dark brown. Decomposers of the
dead roots of Atriplex and other desert shrubs. Monotypic: R digueti
(Pat. & Har.) Herr.

51

Chlamydopus Speg.

PI. VIII B; Fig. 24

Fruiting body epigeous at maturity, 2-3.4 cm broad, depressed-oval to

nearly globose, exoperidium a thin, hypha I sand case which breaks up
and falls away; endoperidium smooth, tough, ochraceous to pallid
cream-colored, apical pore formed with undifferentiated peristome
(Fig. 24). Stalk 5-35 cm long, 10-35 mm broad at apex, tapering gradually
from the apex to a narrow base, woody, dull white with distinct
longitudinal grooves, flat at apex and inserted below peridium, arising
at base from a small volva, 4-15 mm tall. Gleba yellow-brown, capillitium of long, distinctive cells (PI. VIII B). Spores 5-7 (-9) p,m, globose,
warted, thick-walled, yellow-brown. Clamp connections present. Decomposer of desert-plant remains, such as Atriplex roots. Monotypic:
C. meyenianus (Kotzsch) Lloyd.

D;ctyocephalos Underw.

Fig. 30

Fruiting body epigeous at maturity, 7-56 cm tall, with a globose to ovoid

head, 5-13 cm broad, 2-6 cm high, seated on the convex apex of a stalk
which arises from a well-developed, persistent volva, 3-11 cm tall, and
usually 2-8 cm below the soil surface. Mesoperidium with scales or
pyramidal warts at maturity, fleshy to gelatinous when young (Long et
aI., 1940); endoperidium 1-2 mm thick on the side, brittle and thin at
apex, breaking up exposing gleba when mature. Gleba cellular, brown,
with odor of decaying fish, texture chalky. Capillitium absent. Spores
6-9.2 x 6-6.7 p,m, subglobose to globose, minutely warted, light brown.
Decomposer of desert-plant roots and organic matter, noted on
Atriplex, rare, locally abundant. Monotypic: D. attenuatus (Pk.) Long &
Plunkett.

Phellor;n;a Berk.

Fig. 27

Fruiting body epigeous, 5-15 cm tall, club-shaped with the apex 2-5 cm
broad, scaly or with pyramidal warts, dry, white to cinnamon in age,
dehiscing at apex to expose the gleba. Stalk equal or nearly so, not
inserted into the gleba, woody, dry, solid, with surface fibrils; base of
stalk with a large ball of mycelium and sand, often with stout, radicating
rhizomorphs, not volvate (Fig. 27). Gleba bright mustard-yellow to
cinnamon, powdery to sticky at maturity. Capillitium absent. Spores
5-8.5 p,m, globose, thick-walled, echinulate, no pedicel, yellow-brown,
many remain attached to the collapsed fascicles of basidia. Maturing
specimens exhibit a carrion-like odor, in deserts and sand dunes.
Monotypic: P. inquinans Berk.

52

Queletia Fr.

Fig. 26

Fruiting body epigeous, 9-12 (-15) cm tall, with an oval head 3-4 (-7) cm
broad, dry, white, nearly smooth at first, becoming reddish-brown and
somewhat roughened at maturity, irregularly splitting over the apex to
reveal the gleba. Head nearly sessile at ground level at first. Stalk soon
elongating to 6.5-8 (-15) cm long, 1.5-2 cm thick, tough, somewhat
larger at base and inserted into the head (Fig. 26) to support the gleba,
smooth, white, soon becoming reddish-brown with irregular loose,
somewhat recurved fibrils. Volva absent. Peridium thick, two-layered,
persistent until maturity. Gleba white when young to a rich red-brown
and powdery at maturity.
m abundant, of interwoven, thickwalled, brown, cyanophilic hyphae, 3.4-10.5 J-Lm diam. Spores (4-) 5-7 x
4.5-6 J-Lm, globose to subglobose, thick-walled with prominent warts and
a short pedicel (Julich, 1983), yellow-brown. Decomposer of spent tan
bark, in North America and Europe, rare. Monotypic: Q. mirabilis Fr.

Schizostoma Ehrenb. ex Lev.

Fig. 25

Fruiting body epigeous at maturity, globose, 2-3 cm broad, with a

sand-case exoperidium which sloughs off at the top at maturity,
revealing a dark brown, smooth to pitted endoperidium which splits
from the apex downward, creating lacerate pieces which open up to
expose the gleba (Fig. 25). Stalk 2-3 (-15) cm long, 2.5-4 mm thick, equal,
brown, striate and covered with scales, inserted in a socket at the base
of the peridium, arising from a pad of mycelium (volval pieces sometimes present, according to Dring, 1973). Capillitium of both hyaline
and pigmented, septate hyphae which fragment. Spores 4.5-6 J-Lm,
globose to slightly pip-shaped, smooth to faintly warted. Monotypic: S.
laceratum Ehrenb. ex Lev.

Tulostoma Pers. ex Pers.

PI. VIII A; Figs. 22 & 23

Fruiting body epigeous at maturity, oval to oval-depressed, dull white

to gray-brown or brown, exoperidium often a partial .sand case or a
soft, floccose layer, sloughing from apex first to expose the endoperidium, which has a well-formed pore and either a well-defined or a
undifferentiated peristome. A distinct stalk, composed of longitudinally
arranged, thick-walled hyphae, is inserted in the base of and supports
the spore sac, which develops from a scanty to copious basal mycelium;
no volva is present. Capillitium of thick- or thin-walled hyphae with
swellings at the septa in some species (PI. VIII A). Spores 2~8 (-15) J-Lm,
globose, elliptical to pyriform, hyaline to dark brown, smooth, with fine
to prominent warts, or reticulate and with a short pedicel in some
species. Cosmopolitan decomposers in arid areas or in xeric habitats
(Wright, 1987}.

53

2. Calostomataceae Pilat
fruiting body epigeous, 3-5 cm high, with a globose head 8-15 mm
thick; exoperidium evanescent, transparent to sticky or cottony,
sloughing off soon (fig. 31) to reveal a tough, papery endo- or
mesoperidium, which is red, green, yellow-brown, or dark brown, with
a pore and peristome, forming a suture or radial slits which open at
maturity. Stalk (5-) 15-45 mm tall, 5-10 mm thick, equal, dry or covered
with a transparent, gelatinous layer. Volva absent. Gleba a white sac,
attached only at the peristome, suspended, powdery when mature.
Paracapillitium hyaline, present at first, soon collapsing. Spores 6-20 x
6-8 J,Lm, globose to elliptical, pitted or echinulate, thick-walled, buff at
maturity. Cosmopolitan, decomposers of plant organic debris.
Ca/ostoma Desv.

PI. II f & III A; fig. 31-34

(Characters of the family)

..

~--en

..
~--p

+--- st

I,

, \1

'.\. ,

" i

. .,/

I ~
'.
\

Fig. 22. Tu/ostoma verrucosum.

The globose head is supported by
an inserted woody stalk (st) covered with brown lacerate scales. A
fine fibrillose exoperidium (p)
sloughs off, leaving the papery endoperidium (en). Note the small
ostiole (0) and growth above
ground .

\\\ \

--v:J-- ground line

54

,
.---. .. '------..
~
: '., ..
"

---::~----

en
ground level

\+-----st
\

~,

1------

rh

---0
.,..,I .. . ,.'.' .
. . .. ..: .
.. .' ' . '.'. :. .\ .
. .
, . '. . , '. ." .
,. '.. . , ' .. ' ,.. . , ..
..." .,,",' . " " ".. ' ..
. . '.. . ' . . ' . ..' . .\ .
.... ..
. ... , ... . '... . .....

.'. . .

~ ." ...

'

.. '
'

Fig. 23. Tu/ostoma simulans. The stalk grows from a dense ball of

mycelium and rhizomorphs (rh) below ground. The fruiting body

develops at the ground level, and the covering of sand and soil is soon
shed to reveal the inner whitish endoperidium (en) and small ostiole
(0) .

55

~.:::.~~-- 0

4--en

Fig. 24. Chlamydopus meyenianus.

Mature specimens show the smooth
endoperidium (en), irregular ostiole
(0) , longitudinally grooved stalk (st),
and the small, inconspicuous volva

----st

(v) .
J

en

----v

...

.' '

(\~ ;., .'

Ii

- - _. p (= sc)

\ .

st

Fig. 25. Schizostoma laceratum. The immature specimen, right, has a

sand case (sc) which sloughs off; the endoperidium (en) splits from the
apex downward, creating the lacerate pieces seen in center and left
specimens. The stalk (st) is striate and arises from a pad of basal
mycelium and is inserted into the peridium (p).

56

..
-"

. ', :":.'.

. '.

.~

. .. IV.

. ,... . '
'. '. :.~:.;
.
'. '!.,...
,,'

.
.
., ' " ' ....
. ,: .

...., .
'. '." ,,... , ; . .
J'. ,_
. "

. .. '.

~~

',

'~:
'" ,
~

'~ ~.. \.' ....

h ','

.... ': .::-~?I:.: . .~.. ' ..... :....

_.
. i'...

'.~!, .,

,.-

..

"""*"-p
.
.

:~

..

'.

:# - - - - st

,..---- g
--p
lacerate
fibrils

,
I,'

---h- - - -

st

Fig. 26. Que/etia mirabilis. The coarse, white stalk (st) has loose,
lacerate, often recurved fibrils and is inserted into the exoperidium (p).
The gleba (g) occupies the oval head, which becomes lacerate exposing
it at maturity.

p--+

g -----''T

Fig. 27. Phellorinia inquinans. The

oval gleba (g) is supported by a
woody stalk (st) and covered by an
exoperidium (p) consisting of
thick, raised scales. The specimen
on right is mature and split at the
apex to reveal the gleba (g). The
woody stalk (st) develops from a
large ball of mycelium below the
soil surface.

---st

\
\

st-+

\
\ I

57

Fig. 29

Fig. 28

en

:::.......:,---- en

-----<

--g
>---

Clr

~-en

Clr--'

I
\----- st

st - -

v
v --.;.
....

. ..'
."

. .

~.,

"

,.

. ..
"

', '

Fig. 28. Battarreoidesdigueti. Numerous ostioles (0) develop in the

endoperidium (en) at maturity along with a circumscissile dehiscence
line (cir); however, actual dehiscence along this line does not take
place. A long woody, lacerate stalk (st) originates below ground from a
woody volva (v).
Fig. 29. Battarrea laciniata. The gleba (g) on right specimen is revealed
by the circumscissile dehiscence (cir) of the endoperidium (en). The
naked gleba (g) is shown in center specimen exposed on the surface of
the endoperidium (en) and supported by the lacerate woody stalk (st)
which arises below ground from a woody volva (v).

58

en ---1,:,

me.-:::::::.:=i

--st
---st
- - solid
center

', n
\

--v
v.7~--v

:---rh

) - - - rh

Fig. 30. Dictyocepha/os attenuatus. The specimen on left has an intact

mesoperidium (me) with discrete pyramidal warts (Fig. 4 D), a woody
rimose stalk (st) which is solid, and an exoperidium which remains
below ground as a volva (v), and has proliferating rhizomorphs (rh). The
center and right specimens show the chambered gleba (g) exposed by
the weathering away of the endoperidium (en).

59
Fig. 31. Ca/ostoma cinnabarina. A young
button on right has a thick, gelatinous.
exoperidium (p) which sloughs off (see left
specimen) to reveal the endoperidium
(en), suture-like peristome (ps), and ostiole (0). The young specimen
ps
in center shows the
elongating, viscid
o
stalk (st).

......'"

.~

..

r----en

---st

en ----'
~-p

"

p
ps

o
__- - ps
. . .,
. . ....

en ---+-'-'.<.

st ------\,

.' . . .....

+--- st

Fig. 32

Fig. 33

Fig. 32. Ca/ostoma lutescens. The specimen shows the typical remains of
the recurved pieces of the membranous exoperidium (p) at the base of
the endoperidium (en) and above the viscid stalk (st). A peristome (ps)
surrounds the ostiole (0) . .
Fig. 33. Ca/ostoma cinnabarina at maturity with ostiole (0), peristome
(ps), elongated stalk (st), and without any exoperidial remains.

60

~.~\
.
.
"

'.

; , . '

, . ..
, .

"

_....l'......
\ - - - - en

ps
~/j~~~~~'~~~'. .
"
' . .,
.

'''',. ,.,'. '.

~

1'"

, . , ,

A;i~:: .

.
. . .
.. ... . ''.
. ....::.,..

h!~:"

" '

"

. ', "
..

,
,

'\.. ..

. .'.

"
I

.'

~ ~,
' .... . ~
"

~' : " ".' ", '

. '",

"
;f

. ,
I

".~.

'.

\.

.,
,

, ,

st

'lI i,
:

.
,
I

.:/'f ,

,
..
,.
f

,
"

I I~

.... ,

'

. . ..'

Fig. 34. Ca/ostoma insignis, A persistent, thick, membranous exoperidium (p) adheres to the stalk (st) and surrounds the smooth endoperidium (en ). Close-up on right shows the circular, radially slit peristome
(ps) surrounding the ostiole. The tough stalk (st) arises from a ball of
mycelium and dirt.

61

SCLERODERMATALES Cunningham
General: The earth balls and false earth stars are predominantly

epigeous and partially hypogeous in deep duff or in a well-developed

litter layer. The fruiting body is round to oval-depressed in Scleroderma, irregularly club-shaped to oval in Pisolithus, to stellate or earthstarlike in Astraeus. A one-layered peridium is present in most taxa except
Astraeus, which has a two-layered peridium (Fig. 36). The immature
gleba is pigmented very early and most often is dark purple, purplebrown, to brownish purple. The gleba is organized into locules or
peridioles which are most easily seen when immature, except in
Pisolithus where they are visible through maturity. Plectobasidia often
extend into the locules but are not organized into a hymenium. They
collapse at maturity and give way to paracapillitium which is usually
hyaline, sometimes has clamp connections (PI. VII B), and may have
thickened walls. At maturity the peridium splits, tears, or weathers
away, revealing a naked gleba; no regular stoma is formed. A subgleba
or columella is either very reduced or absent. The spores are usually
ornamented at maturity, ranging from reticulate, reticulate-warted, to
discretely warted (PI. IV C & D), and are thick-walled and usually
globose to ovoid. The spores are either wind-disseminated or washed
into the soil by rainwater. The main glebal pigmentation is derived from
the exosporium as it matures. The species are distributed among three
families which are distinguished from each other by the differences in
morphology and development of the fruiting body.
Ecology: The North American and European species in Astraeus,

Scleroderma, and Pisolithus form ectomycorrhizae with members of

the Pinaceae and Fagaceae. In the Southern Hemisphere, they are quite
probably ,associated with the Myrtaceae as well. The ecological status of
other genera placed in this order is not known, but, as presented by
Dring (1973), it is likely that the species in other genera may be
decomposers or even parasites, and their inclusion within the Sclerodermatales must be reevaluated.

62

Key to the Families of the Sderodermatales

1. Exoperidium splitting at maturity to form an earth star; internal
surfaces of the hygroscopic rays rimose and checkered in appearance; endoperidium fibrillose with an apical tear or slit or with
multiple pores; gleba without a columella, but it may be supported
by columns (Fig. 38) or sessile (Fig. 36) ............ 2. Astraeaceae
1. Exoperidium not as above; if the peridium splits and opens, the
naked gleba is exposed ..................................... 2
2. Fruiting bodies compound (Fig. 39) with numerous spore sacs
seated on a common stroma ............... 3. Broomeiaceae
2. Fruiting bodies single or several in a tight cluster but not on a
common stroma; gleba formed at first in peridioles with common walls (Figs. 35, 37) ............... 1. Sclerodermataceae

1. Sderodermataceae Sebek
Fruiting body epigeous at maturity, oval, sessile or with a fleshy
subgleba on a short stalk, 0.8-15 cm broad, 1.5-10 (-25) cm tall, dry,
brown, nearly smooth to warted, tough, with thick or thin peridium
which splits at the apex in age to reveal the naked gleba. Gleba brown
or white when very young, soon becoming purple with minute to
persistent and well-developed peridioles divided by thin or thick
fragile white walls, powdery in age, brown to purple-brown. Capillilium absent. Spores globose to subglobose with fine to coarse warts or
reticulate ornamentation, thick-walled, brown. Mycorrhizal with
shrubs and trees, often in severely disturbed, dry sites, throughout the
world.

Pisolithus Alb. & Schw.

Fig. 37

Fruiting body epigeous, globose, ovoid, to clavate or pear-shaped, 4-25

cm tall, 3-15 cm broad, olivaceous-yellow to brown, dry, brittle at the
apex and breaking up to reveal the dark brown gleba with persistent,
thick-walled peridioles (Fig. 37) which develop and enlarge as the
fruiting body matures from the top down. Eventually peridioles may
develop in the lower stalk almost to the ground; rhizomorphs proliferate from the base. Capillilium absent. Spores 7-12 J.Lm, globose,
cinnamon-brown, spiny, and covered with a thin gelatinous layer.
Monotypic: P. tinctorius (Mich. ex Pers.) Coker & Couch.

63

Scleroderma Pers.

PI. IV C & D; Fig. 35

Fruiting body 0.8-15 cm broad, 1.5-10 cm tall, with a tough, brown,

smooth to warted peridium and a reduced to well-developed, fleshy
subgleba. Gleba white when very young, soon purple to purple-brown,
with minute peridioles separated by fragile, white walls which break
down early as the spores mature, becoming purple-brown and powdery in age. Base has rhizomorphs which are sometimes stout and
prolific.
m absent. Spores as in the family.

2. Astraeaceae Stanek
Fruiting body epigeous at maturity, sessile, subglobose when young,
1-6 cm broad, exoperidium splitting from the apex and recurving
stellately to form an earth star and exposing the endoperidium which is
dehiscent by an apical slit or by multiple pores. Paracapillitium abundant, hyaline, with clamp connections (PI. VII B). Spores 4-11 JLm, globose, thick-walled, warted, reddish-brown. Mycorrhizal with shrubs
and trees, throughout the world.

Astraeus Morgan

PI. VII B; Fig. 36

Fruiting body epigeous, sessile, subglobose, 1-6 cm broad, exoperidium

splitting stellately (earth star-like) to reveal checkered or rimose,
hygroscopic rays (Fig. 36) and a fibrillose endoperidium with a single,
apical, lacerate slit. Gleba white at first, reddish-brown in age and
powdery at maturity, without a columella (compare Fig. 36 with Geastrum, Fig. 16 B). Paracapillitium as in the family. Spores 7-11 JLm, globose, thick-walled, with warts, brown. Mycorrhizal with trees and
shrubs.

Myriostoma Desv.

Fig. 38

Fruiting body epigeous at maturity, oval-depressed, 2-9 cm broad,

exoperidium of sand and hyphae, splitting stellately like an earth star,
recurved rays smooth or rimose but not checkered; a smooth endoperidium dehisces to form many lacerate ostioles and is supported on the
recurved exoperidium by stout columns (Fig. 38). Gleba white at first,
brown and powdery at maturity. Paracapillitium of spinose, unbranched, thick-walled, hyaline hyphae. Spores 5.8-7.6 JLm, globose,
thick-walled, subreticulate with prominent warts, brown. Rare, occurring in dry habitats, role unknown. Monotypic: M. coliforme (Dicks. ex
Pers.) Corda.

64

3. Broomeiaceae Zeller
Compound fructification, 5-15 cm broad, consisting of numerous spore
sacs seated in the base of the exoperidium which forms an individual
exoperidium surrounding each, or in a cup-like exoperidium surrounding all of the spore sacs. Spore sacs are seated on a thin or thick
stroma (Fig. 39), each spore sac (endoperidium) having a pore or ostiole
with a delimited or indefinite peristome. Gleba dark brown without a
sterile base. Paracapillitium thick-walled, brown to hyaline, branched,
straight or sinuous, hyaline in cotton blue in lactic acid. Spores (4-) 5-7.5
/-Lm, globose to broadly elliptical, thick-walled, minutely punctate to
warted, with a short pedicel. Decomposers or parasites of plant
material, in Africa and Central America.
Broomeia Berk.

Fig. 39

Compound fructification, up to 15 cm broad, 5-900 spore sacs seated in

alveoli on upper surface of a thick, fleshy stroma surrounded by the
lower half of the exoperidium at maturity. Peridium of spore sac
tomentose, covered with sclerotium-like, minute balls of hyphae;
ostiole with clearly delimited, conical, fimbriate peristome. Paracapillitium hyaline, thick-walled, branched and sinuous. Sp~res brown,
elliptical, with numerous, small echinulations, 6.5-8.5 (-11) x 3.5-4.2 (-5)
/-Lm in diameter. Broomeia congregata is possibly parasitic on Acacia.
Monotypic: B. congregata Berk.

Diplocystis Berk. & Curtis

Compound fructifications to 10 cm in diameter, 3-60 spore sacs
crowded together, each one seated in a cup or exoperidium on top of a
flat, thin stroma. Peridium of spore sac smooth, with a circular ostiole
which becomes irregular, peristome indefinite. Paracapillitium thickwalled, refractive , hyaline. Spores brown, globose with low, numerous
warts, 5-6.5 (-7) /-Lm in diameter. Monotypic: D. wrightii Berk.

65

g--H

..

'

4--

.
.'

..

t-r-

~---rh

Fig. 35. Scleroderma po/yrhizon. The delicately chambered immature

gleba (g) is illustrated in upper right. The peridium (p) in lower right
specimen is splitting to reveal the naked gleba (g), shown in the mature
left specimen. Pieces of the peridium have subsequently recurved,
exposing more of the gleba (g). Robust sand-covered rhizomorphs (rh)
proli ferate from the base.

66

---g

---en
r

en

10
't---

10

.'

..

-----en
---me

+--r

---p
,

Fig. 36. Astraeus hygrometricus. The immature gleba (g) in upper right is
delicately chambered, surrounded by an endoperidium (en), and has an
ostiole (0), but has no columella. Mature specimen at bottom under
moist conditions has patches of mesoperidium (me) left on the rays (r)
giving it a checkered appearance. The delicately fibrillose endoperidium (en) is elevated by the recurved rays and has a lacerate ostiole (10),
at maturity. In the middle is a dried specimen with the involute rays (r)
covering the endoperidium (en); the lacerate ostiole (10) is visible.

67

-,..(.if----

pr

..

p--

..

~-g

... ' . , .

," ..
'.:.,,'
,' ..

j~.::;:;:.;,. " .,
'~"

.'

.,

"

"

..:.~.~"
. ."' . .
,

--pr

t,

.,. . .

'.";

.,

f.

..
.
. .
..

',.

,':.'
~"
~

"

st ---t+-

., .

."

;t. .

~ oJ

'"
. ':

I .

'\

1" ~ .
'.

~---sub

.(W.;:..jr': r)':::;""''-..
... ;

. . .. . ,. .,.
.'

,\'

I'

'

.\

,,' s

'.'

pr

.
.'

"

Fig. 37. Pisolithus tinctorius, On left is an immature club-shaped fruiting

body with the exoperidium (p) in place and an irregular short stalk (st),
The sectioned specimen on right shows the peridioles (pr) in the gleba
(g), which are immature toward the base and have disintegrated at the
top, giving rise to a powdery gleba (g) at maturity. At the base is the
subgleba (sub).
'

68

10
10
--en

.:.~~ S::::::~- columns

en -------F.

p
r

Fig. 38. Myriostoma coliforme. The exoperidium (p) splits to form

recurved rays (r), elevating the endoperidium (en) , which is supported

by a cluster of short columns. A number of small lacerate ostioles (10)
forms over the surface of the endoperidium (en).

_ _- f
ps _. ::::::::::::::-

.-<__

t:::;en

-p

Fig. 39. Broomeia congregata. A

large fleshy stroma supports the

tightly packed spore sacs, or endoperidium (en) , surrounded by the
remains of the peridium (p). Each
spore sac has a delimited, fimbriate (f) peristome (ps) surrounding the ostiole (0).

,""<1".,+ -_ _ stroma
.

..,. . . .

.. ,",. ........

.~

.....

;: ~."{.,

.. .

69

NIDULARIALES
General: The "bird's-nest" and "cannon-ball" fungi are aptly named.

The small epigeous fruiting bodies are oval to urn- or trumpet-shaped

(Figs. 40-42) and most often fruit in gregarious clusters. The one- to
three-layered peridium is either open at the top, ruptures and loses its
cover, or splits and folds back as the fruiting body matures. One to
several oval to disk-shaped peridioles (Fig. 41) are revealed within the
!ruiting body. The peridioles are either attached to the inner wall of the
peridium by a gelatinous pad embedded in a gelatinous material, or are
held fast by a middle piece (Fig. 41) connected to a funicular cord. The
peridioles are discharged by raindrops which dislodge them and splash
them out. The funicular cord unfolds and trails behind the peridiole as
it is discharged. In the case of Sphaerobolus, the "cannon-ball", a
single peridiole fits snugly into the outer cavity formed by the peridium
(Fig. 42). The peridioles are thick-walled with one to three walls and
contain the gleba composed of short, clavate basidia and oval to
globose basidiospores. The spores are smooth and thick-walled and
reach their mature size following the collapse of the basidium. The
"cannon ball" forcibly discharges the peridioles in the direction of the
light source. There is no capillitium present in the order.
Ecology: The species are placed in two families (Nidulariaceae and

Sphaerobolaceae) by Dring (1973) and are distributed throughout the

world. The species are few in number but cosmopolitan as decomposers of wood or plant material containing cellulose. Such substrates
include sticks, wood chips, mulch, grass, moss, and leaf litter, to
mention a few. One species, Cyathus stercoreus, is commonly found on
dung, where it invades and colonizes the undigested plant material.

Key to the Families of the Nidulariales

1. Fruiting bodies single to gregarious, with several to many peridioles
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1. Nidulariaceae
1. Fruiting bodies gregarious, with one large dark brown peridiole
exposed by the stellate splitting of the apex (Fig. 42) ............ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2. Sphaerobolaceae

70

1. Nidulariaceae Fr.
Fruiting bodies usually gregarious,small, 5-15 mm broad, 4-8 mm high,
urn- or vase-shaped (similar to a bird ' s nest), with or without an
epiphragm or lid at first, containing one to many disc-shaped peridioles
(Fig. 41), which are attached by a middle piece to the endoperidium or
embedded in a gelatinous material. Capillitium absent. Spores oval to
broadly elliptical, smooth, thick-walled, borne within the peridiole (Fig.
41). Cosmopolitan, decomposers of wood and plant organic matter.
New fruiting bodies often develop within the old ones.

Key to the Genera of the Nidulariaceae

1. Fruiting bodies very small (0.5-2 mm), barrel-shaped; peridium of

loosely interwoven, clamped hyphae; solitary or in small groups

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Mycocalia
1. Fruiting bodies small or more often larger (3-10 mm), urn- or cupshaped; peridium composed of non-clamped hyphae; gregarious
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2
2. Fruiting body 1-1.5 mm broad, white, spherical; peridium splitting stellately to reveal a cadmium-yellow endoperidium .....

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Nidulariopsis
2. Fruiting body larger, gray, brown, tan to cinnamon; peridium
not splitting as above and endoperidium not yellow ........ 3
3. Funicular cord attached to each peridiole ..................... 4
3. Funicular cord absent; peridiole embedded in a gelatinous matrix

.................................. '. . . . . . . . . . . . . . . . . . . . . . . .. 5
4. Peridioles black to dark brown; epiphragm white to lightly pIgmented .......................................... Cyathus
4. Peridioles dull white; epiphragm light tan ....... Crucibulum
5. Fruiting body a cup without an epiphragm; peridium of irregular
spiny cells, not splitting into stellate lobes (PI. VI B) ..... Nidularia
5. Fruiting body urn or cup-shaped with an epiphragm at first; peridium not as indicated above but splitting at apex into four to seven
stellate lobes .......................................... Nidula

71
Crucibulum luI.

Fruiting bodies gregarious, 1.5-10 mm broad, 5-12 mm high, urn- to

cup-shaped, light tan to cinnamon, with an epiphragm at first. Peridium
thick and a single layer, dull white inside. Peridioles disc-shaped, white,
4-6 in number, with a funicular cord individually attached by a middle
piece to the endoperidium. Spores 7-11 x 4-6 J,tm, elliptical, smooth,
thick-walled. Found on decayed wood and wood debris.
Cyathus Haller ex Pers.

Figs. 40 & 41
Fr uiting bodies gregarious, 4-8 mm broad, 7-18 mm high, vase-like,
tru mpet-shaped, to urn-shaped, brown to gray-brown with a white
epiphragm at. first, peridium of three layers with an endoperidium
which is shiny to dull brown and smooth to longitudinally striate (Fig.
40). Peridioles numerous, black or dark brown, with a funicular cord
attached by a middle piece to the endoperidium. Capillitium absent.
Spores 5-15 (-20) x 5-8 (-25) J,tm, elliptical to subglobose, thick-walled,
hyaline or light yellow-brown. Found on decayed wood or wood
debris, one species on woody fragments in cow or horse dung.
Mycocalia Palmer .

Fruiting bodies very small, 0.5-2 mm broad, solitary or in small groups,

white, often brownish at maturity, barrel- to lens-shaped. Peridium of
loosely interwoven, clamped hyphae. Peridioles one to many, discshaped, yellow-brown to red-brown, embedded in a gelatinous matrix
when fresh. Spores 5-9.5 x 3.5-7 J,tm, hyaline to yellow-brown, elliptical,
smooth. Found on herbaceous stems and other plant debris.
Nidu/a White

Fruiting bodies 3-8 mm broad, 5-15 mm high, urn- or cup-shaped, gray,

with an epiphragm at first, splitting into four to seven lobes at maturity.
Peridioles light brown, embedded in a gelatinous matrix when fresh, no
funicular cord. Spores 8-10 x 4-6 J,tm, ovoid to elliptical, thick-walled,
light brown.
Nidu/aria Fr. & Nordholm

Fruiting bodies gregarious, 0.5-6 mm broad, 0.5-3 mm high, irregular in

shape or with a well-formed cup, thin and fragile, without an epiphragm. Peridium with a surface tomentum composed of irregular,
spi ny cells (PI. VI B) over a more compact wall, soon breaking down and
exposing a pile of brown peridioles at maturity. Peridioles are embedded in a gelatinous matrix when fresh (no funicular cord). Spores
5-10.5 x 4-5.5 J,tm, elliptical to broadly elliptical, slightly thick-walled,
hyaline.

72

N;dular;ops;s Greis
Fruiting bodies 1-1.5 mm broad, spherical or subspherical, white or dull
buff, with a stout basal rhizomorph. Peridium of three layers, without a
gelatinous layer, splitting at the apex into 3-8 stellate, slightly recurved
lobes, interior of endoperidium cadmium-yellow. Glebal mass soft and
sticky, brown, becoming nearly black upon exposure to air, divided
internally into many definite chambers lined with the hymenium.
Spores 6-10 x 5-6 J,Lm, oval to broadly elliptical, smooth, thick-walled.
Gregarious on partially decomposed wood and wood debris.

2. Sphaerobolaceae Schroeter
Fruiting bodies gregarious, 1-3 mm broad, subspherical to ovoid, white
to buff, peridium of four to five layers including a gelatinous layer;
exoperidium splitting at the top into 4-7 stellate lobes when mature
(Fig. 42) and exposing the dark brown, single peridiole. Forcible
ejection of the peridiole (up to 14 feet) leaves the evaginated, white
endoperidium as a sphere projecting from the exoperidium. Capillilium absent. Spores 6-10 x 4-6 J,Lm, elliptical to oblong, thick-walled,
hyaline. Cosmopolitan, on dung, wood, and wood debris, usually on
the ground.

Sphaerobolus Tode & Pers.

(Characters of the family)

Fig. 42

73

setae

str
pr

.
. ..
..

pr -----'"

. .
. , .'
~-g

--fun

sh --....:::

/<r<~,
~--mp

~-epl

me

,. .

Fig. 40. Cyathus striatus. The young fruiting body in center has an
epiphragm (epi) in place. Older specimens on left have lost the
epiphragm, and the peridioles (pr) and striations (str) are visible.
Cut-away above shows attachment of the middle piece (mp) to the
mesoperidium (me), sheath (sh), funicular cord (fun), and setae. One
peridiole (pr) is sectioned, revealing the gleba (g).

tu

- - sp

b
Fig. 41. Cyathus striatus peridiole in
longitudinal section showing the tunica (tu), basidia (b), and spores (sp).
The purse (pu) surrounds the funicular cord (fu n) and hapteron (ha),
which is attached to the middle piece
(mp) at the base.

----pu
0 4 - - - - fun

---ha

----mp

74

pr

,.

, .'

"

.'

: :.. . .
..
)U'i"
.
'.
.""...,"
',; :. . .
f , ..
.
'. .. .
.'., ".:. ", .. .
.' '.. ." .

'.

.
.'
~

I.

""'-

.
. .

" . . ~--- en
. . <
...
'.,.

.
.

'

.'

...

~.

.:.

. , ".

tI

......:..~:. ",",.)'.. : .
~

. ... .

...,.., - - - p

'

. '

'.

P ---/f-'...,.-.., .

..' ",.....

.
.
... . . .....,',... ,,'.. ..
.;. .. ...'..

..

..

..

..

..

Fig. 42. Sphaerobo/us stellatus. Lower left shows splitting of the exoperidium (p) into a series of coronate lobes, shown at top with the
peridiole (pr) in place. Forcible ejection of the peridiole leaves the
evaginated endoperidium (en), as shown on the right.

75

PHALLALES Cunningham
General: The "stinkhorns" are usually hypogeous or are partially

hypogeous at first, becoming epigeous as the stalk elongates during the

fruiting process. One group of hypogeous species has no stalks, and
those mature as an oval, oval-depressed, to pyriform fruiting bodies
enclosing the gleba, even after the fruiting body and hymenium are
mature (Fig. 49). The flaccid to almost rubbery, two- to three-layered
peridium protects the developing hymenium until it is mature. The
mesoperidium is gelatinous in many species and protects the immature
hymenium. The receptacle varies from a simple hollow stem (Fig. 43)
with an apically placed hymenium to a complex, clathrate structure
bearing the hymenium on the interior arms (Fig. 62). A fetid odor
ca used by several biochemicals, including methyl mercaptan and hyd rogen sulfide (List & Freund, 1968), is typically produced by the slimy,
olivaceous-brown hymenium at maturity. The slimy hymenium is
present in all families except the Protophallaceae, and the odor attracts
flies, yellow jackets, and other insects. The small rod-shaped basidiospores have a mucilaginous covering and are borne on short, clavate
basidia. The spores are sessile on the basidium, 4 to 8 in number, and
whorled about the apex, tending to be horizontal rather than erect.
Sto ut rhizomorphs proliferate into the substrate and often become
pink to purplish in color upon exposure to air or when handled. There
are six families commonly included in the order. Two families have but
a single genus, Claustulaceae with C/austula and Gelopellidaceae with
Ge/opellis. Two additional families, the Hysterangiaceae and the Protophallaceae, have simple, often hypogeous fruiting bodies, and the
gleba remains unexpanded within the peridium. The showy stinkhorns,
which expand, becoming epigeous, and rely most heavily upon insect
dissemination of the spores, are found in the Phallaceae and Clathraceae. These two families contain the majority of the genera and
species in the Phallales.
Ecology: The Phallales are cosmopolitan, with the greatest diversity and

number of species found in the tropical and humid temperate biomes.

There are no tundra species and very few in the taiga or boreal biomes.
The majority of the species are decomposers of wood and plant organic
matter and are commonly found fruiting in mulch beds. Their rhizomorphs can be traced to buried roots, stumps, and other woody
material. The only totally hypogeous taxa occur in the Hysterangiaceae
and form ectomycorrhizae with trees and shrubs (Miller, 1982). The
Species grow readily in culture and are easily transported in organic

76

matter. Asian species, such as Lysurus mokusin and Aseroe rubra, are
now found in both the eastern and western United States near large
metropolitan areas where exotic plants have been established from
Asia. This trend will probably continue throughout the world in
artificial plantings, parks, and arboretums. It is not known if all of the
species will be able to invade wild plant communities. However,
Pseudoco/us is now found in woodlands in North Carolina, according
to Burk (pers. comm.), and elsewhere in the South in natural habitats.
In any event, new records of exotic phalloids will, no doubt, continue
to be reported on various continents.

Key to the Families of the Phallales

1. Fruiting body hypogeous or epigeous at maturity, indehiscent or
splitting at the apex at maturity (Figs. 48-50); gleba firm to slimy,
olive-green, greenish-gray, to olive-brown, fetid to odorless; columella rudimentary to well developed, fleshy or a transparent gel
(without a stalk or receptaculum) ............................ 2
1. Fruiting body epigeous at maturity with a stalk or receptaculum
(Figs. 44, 61, for example); gleba slimy, olive-gray, green, to
greenish-black, fetid at maturity ............................. 4
2. Fruiting body oval, firm, and epigeous with a club-shaped base
or hypogeous, small, and firm without a transparent gelatinous
columella; white rhizomorphs becoming pink to lavender upon
exposure to air; fetid at maturity ......... 4. Hysterangiaceae
2. Fruiting body hypogeous to partially epigeous, ovoid to globose, flaccid; white rhizomorphs becoming pink or unchanging
when exposed to air ..........,.......................... 3
3. Longitudinal section shows a branched, gelatinous columella which
proliferates among the lobes of greenish-gray glebal tissue; with
white rhizomorphs which remain unchanged upon exposure to air;
odorless or pleasant-smelling when mature ... 3. Protophallaceae
3. Longitudinal section shows a central, white, fleshy columella
unbranched and surrounded by a solid, olive-green gleba; endoperidium a thin, white layer surrounded by a thick, translucent
mesoperidium beneath the tough exoperidium (appearance of an
unopened Mutinus without an unexpanded receptaculum); white
rhizomorphs becoming pink upon exposure to air; fetid odor when
matu re .................................... 6. Gelopellidaceae

77

4. Fruiting body unipileate with external or internal gleba and a

single stalk (Fig. 44) ..................................... 5
4. Fruiting body multipileate with a finger-like to clathrate receptaculum (Figs. 52,62) . . ...................... 2. Clathraceae
5. Fruiting body with gleba external at maturity. . . . . .. 1. Phallaceae
5. Fruiting body an obovate head seated on a fleshy base; gleba internal, with a green layer attached to the inner peridial wall of the
hollow head (rare) ............................ 5. Claustulaceae

1. Phallaceae
Fruiting body a single, chambered, hollow stalk with the olivaceous,
fetid gleba covering the outside of its apex or a well-developed single
head, with pileate development, expanding from a flaccid button, the
re mnants of which act as a volva. Peridium three-layered. Spores
narrowly elliptical, covered with mucilage, olivaceous. Basidia most
commonly 6- to 8-spored. Cosmopolitan, decomposers of wood and
plant organic matter.
Key to the Genera of the Phallaceae

1. Gleba over the outside of a nearly free head (Fig. 45) located at the
apex of the receptacu Iu m ................................... 2
1. Gleba over the apex or upper surface of the tapering receptaculum
(Fig. 47)

................................................... 4

2. Head capped by a white calyptra over an olive-gray, lamellate

gleba (Fig. 45) .................................... Itajahya
2. Calyptra not present .................................... 3
3. Indusium extending from beneath the head (Fig. 44) ............ .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dictyophora
3. Indusium absent (see note under Dictyophora) ........... Phallus
4. Gleba olive to olive-brown, covering the apex of the white to
red receptaculum (Fig. 47) . . ...................... Mutinus
4. Gleba olive-green, looking like a constricted collar one-third
distance from the white, chambered apex (Fig. 46) .......... .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Stahe/iomyces

78

Dictyophora Desv.

Figs. 43 & 44

Fruiting body as in Phallus but in addition has an indusium which

extends from beneath the head to form a long, reticulate, white to
yellow net. Spores 3.5-4 x 1.5-2 Mm. (A questionable genus since the
indusium is not always present or visible; see Dring, 1964.)

Itajahya Moller

Fig. 45

Fruiting body with a single, white to pink, chambered, hollow stalk, a

cylindric head, and an apical gleba capped by a white calyptra to which
the gleba is initially attached (appears similar to species of Phallus).
Spores 3-3.5 x 1.5-2 Mm, narrowly elliptical, smooth, hyaline. North and
South America (long & Stouffer, 1943).

Mutinus Fr.

Fig. 47

Fruiting body a single, white to red, chambered, hollow stalk tapering

toward the apex with an olivaceous, slimy gleba covering the area just
below the apex. Spores narrowly elliptical, 3.5-7 x 1.6-2.5 Mm.

Phallus Hadr.
Fruiting body with a single, white, thick, chambered stalk with a flaring,
free head attached at the apex and covered with a gray-green to dark
olive-green gleba, apex hollow. Spores nearly cylindric, 3-4.5 x 1.2-2
Mm.

Staheliomyces Fischer

Fig. 46

Fruiting body a single, chambered, hollow stalk with a band of olivecolored gleba shaped liked a collar or constriction around the receptaculum just above the center and well below the apex. Monotypic: S.
cinctus Fischer in Central and South America.

2. Clathraceae
Fruiting body sessile or arising from a short, chambered stalk, composed of a receptaculum of upraised, spreading, sometimes arched and
united arms or a latticed (clathrate) receptaculum composed of interconnected tubular or gelatinous arms expanding from a flaccid button
which acts as a volva. Gleba located on the inside and sides of the
clathrate receptaculum, olivaceous-brown to green (multipileate development). Peridium three-layered. Basidia 6- to 8-spored. Spores
narrowly elliptical, covered with mucilage. Cosmopolitan, decomposers of wood and plant organic matter.

79
Key to the Genera of the Clathraceae

1. Receptaculum with a distinct stalk ........................... 2

1. Receptaculum sessile (or with a short, inconspicuous stalk) within
the valva .................................................. 6

2. Receptaculum with upraised arms or finger-like projections

wh ich tend to separate in age ............................ 3
2. Receptaculum with arms or apical chambers which do not
separate at maturity (see also Pseudoco/us) ................ 4
3. With recurved arms which extend out to a nearly horizontal position (Fig. 53) .......................................... Aseroe
3. With short, thick arms which are upright, separating only slightly in
age (Figs. 52 & 54) ............. Lysurus (see also Kalchbrennera)
4. Gleba in an ovoid, apical chamber (Fig. 51) ......... Simblum
4. Gleba on the arched arms which are fused at the apex ..... 5
5. Arms joined at the apex to a fertile, apical network (Fig. 56) ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Co/us
5. Fertile arms arched (3 to 4, rarely 5 arches) and tapered to a common point, usually joined, sometimes not in age (Fig. 55) ........ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudocolus
6. Receptaculum clathrate (of latticed or interconnected tubes)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 7
6. Receptaculum of non-clathrate, arched arms, which are persistently joined at their tips or separate in age (if separating in age,
a short stalk is present) .................................. 8
7. Receptaculum of a tubular latticework (Fig. 62) ......... Clathrus
7. Receptaculum of a gelatinous latticework (Fig. 61) .... IIeodictyon
8. Arched receptaculum with a lantern-like gleba and glebifer suspended below the arch (Figs. 57 & 58) " . . . . . . . . . . .. Laternea
8. Lantern-like gleba not present ........................... 9
9. Arches not separati ng at matu rity ........................... 10
9. Arches separating and recurving almost to the ground; dark
greenish-brown gleba over the inner surface of the arms, with a
short inconspicuous stalk (Fig. 60) .................... Anthurus

80

10. With three to five persistent arches, having somewhat roughened or smooth outer faces ............................. 11
10. With three to six white, orange to yellow, stout arms with corrugated outer faces and a row of teeth on either side .......... .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Blumena"ia
11. With pink to red arches and a roughened outer face of the receptaculum; fetid smell .................................. Linderia
11. With white arches and a smooth outer face of the receptaculum;
aromatic or licorice-like smell .......................... Ligiella

Anthurus Kalchbr. & McOwan

Fig. 60

Fruiting body with a short, inconspicuous stalk from which a receptaculum of 5-8 arms arises in an arch; arms initially joined at the apex but
soon become free and recurve irregularly to expose the inner surface
of each arm, which is unevenly covered with the green, fetid gleba (Fig.
60). Spores 3-4 x 1-1.5p,m, narrowly elliptical, green in mass.

Aseroe labill. ex Massee

Fig. 53

Fruiting body a long stalk culminating in numerous arms which taper to

a point and are outstretched almost at right angles, forming a fertile,
central "plate", hymenium extending out along the upper surface of
the rays (Fig. 53). Spores narrowly elliptical, olive-brown in mass, 4-6 x
1.2-2 p,m.

B/umenavia Moller
Fruiting body 5-7 cm long, 2-2.5 cm broad, receptacle with three white
stout arms tapering upwards from the basal volva and united at their
apices; outer face corrugated transversely; inner face of each arm
marked by two longitudinal furrows and by many transverse corrugations; the angle between the inner and outer faces is marked by a
distinct row of pointed teeth. Gleba dark olive-green along inner face
of the arms. Spores 3-4 x 1-1.5 p,m, cylindric, thin-walled. Smell fetid.
Similar to Linderia, known in South America.

Clathrus Micheli. ex l.

Fig. 62

Fruiting body a latticed (clathrate) receptaculum composed of hollow,

tubular arms which rise directly from the basal tissue within the volva.
Spores 4-6 x 1.5-2.5 p,m, elliptical, smooth, hyaline or with tinted
eplspore.

81

Colus Caval. & Sech.

Fig. 56

Fr uiting body a short stalk from which 6 columns arise to form a red,
arching receptaculum, columns joined at the apex forming a fertile,
cl athrate, apical network. Gleba is on the inside of the receptaculum.
Spores narrowly elliptical, olivaceous, 5-6 x 1.5-2 J,Lm.
lIeodictyon Tul. ex Raoul.

Fig. 61

Fr uiting body a latticed (clathrate) receptaculum composed of gelatino us arms, sessile within the volva. Spores 4-6 x 1.5-2.5 J,Lm, elliptical,
green In mass.

Kalchbrennera Berk.

Fig. 54

Fruiting body a single, red, chambered, hollow stalk 8-12 cm tall, 2.5-3
cm broad, culminating in short, irregular, dichotomously branched
arms. Gleba develops among the apical arms, slimy, olive-brown. Volva
persistent, flaccid, and white. Odor fetid. Spores 3.5-4.5 x 1-2 J,Lm,
narrowly elliptical, smooth, hyaline. Africa (Lysurus of Dring, 1980).
Laternea Turpin

Figs. 57 & 58

Fruiting body unstalked, composed of 2 to 3 tubular arms which arch

from the volva and are joined at the apex; with a glebifer (lantern-like
gleba) suspended below the arch. Spores 4-5 (-5.5) x 1.5-2.5 J,Lm,
cy lindrical, smooth, base obscurely truncate.
Ligiella Saenz

Fruiting body 4.5-6.5 cm long, 2.2-5 cm broad, receptacle with four to

five white stout chambered arms, united at their apices and free at the
base as in Linderia (Fig. 59). Gleba dark olive, divided into separate
compartments located in the middle of each arm. Volva up to 25 mm
high , dull white, persistent, tough, with a gelatinous mesoperidium and
cylindrical, white, basal rhizomorphs (to 1.5 mm thick). Spores 4-6 x
1.5-2 J,Lm cylindric, thin-walled, hyaline. Smell aromatic or licorice-like.
Known only from Costa Rica, (Saenz, 1980). Monotypic: L. rodrigueziana Saenz.
Linderia Cunn.

Fig. 59

FrUit ing body 5-8 cm long, 3-3.5 (-4) cm broad, receptacle with three to
fi ve, stout, pink to reddish, equal arms, united at their apices; outer
fa ce roughened; inner face of each arm irregularly, transversely corrugated ; volva well-formed, white, with abundant basal rhizomorphs.
Gleba dark olive-brown to blackish olive, extending along the inner

82
face of the arms. Spores 4-6 x 1.5-2 J-Lm, cylindric to narrowly elliptical,
thin-walled. Smell fetid. Common in southeastern and southern North
America.

Lysurus Fr.

Fig. 52

Fruiting body a long stalk culminating in 3 or 4 short, thick arms (rarely

2 arms) which are erect and held together at first, opening somewhat at
maturity. Gleba covers the inner surfaces of the arms. Spores narrowly
ellipsoidal, brownish to olivaceous, 4-5 x 1.5-2 J-Lm.

Pseudoco/us Lloyd

Fig. 55

Fruiting body with a stalk shorter or equal in length to the triquetrous

or quadrate arms which arch and taper to a common point, usually 3-4
(rarely 5) fertile arms with olive-green gleba covering the inner faces.
Spores 4.5- 5.5 x 2-2.5 J-Lm, elliptical, thin-walled, hyaline, green in mass.

5imblum Klotzsch

Fig. 51

Fruiting body with a long stalk culminating in an oval, chambered head

which is larger in diameter than the stalk. Gleba develops within the
depressions of the chambered head (Fig. 51). Spores 3.5-4.4 (-7) x 1.4-2
J-Lm, narrowly elliptical, green in mass.

3. Protophallaceae Zeller
Fruiting body epigeous to partially hypogeous, flaccid, ovoid to subglobose, like a stinkhorn button. A translucent firm or jelly-like trama
surrounds the olive-green to olive-brown glebal plates, which are
radially arranged around the point of attachment. A more or less firm
gelatinous basal tissue supports numerous, proliferating white rhizomorphs, which remain white when exposed to the air. Odorless at
maturity or with a smell of passion fruit or rotting oranges. Known from
Asia and North and South America. Biological role not clearly known.
Key to the Genera of the Protophallaceae

1. Fruiting body small (10-15 mm broad), subglobose, chalky white,

staining brown; gleba a brown, powdery mass at maturity (not well
known) ............................................ Caillaruia
1. Fruiting body large, club-shaped to ovoid 3-7 cm high, 2.2-12 cm
broad; gleba tough and not powdery at maturity .............. 2

83

2. Fruiting body club-shaped (3-7 cm high) with a fleshy gelatinous

stipe-columella; chambered gleba brown to yellow-brown ...

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Rhopalogaster
2. Fruiting body ovoid (2.5-12 cm broad) with a gelatinous to
fleshy-gelatinous peridium; gleba olive-green to greenish gray

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3
3. Fruiting body ovoid (3-12 cm broad); peridium rather thin (+ 1 mm);
olive glebal chambers embedded in a translucent, gelatinous,
branched columella (Fig. 50) ......................... Kobayasia
3. Fruiting body ovoid, smaller (2.2-3.5 (-7) cm broad); peridium thick
(3-4 mm); olive glebal chambers embedded in a fleshy to gelatinous
.columella-like central core .................................. 4
4. Fruiting body with a fleshy, non-gelatinous, branching
columella-like central core; Asia ............... Protuberella
4. Fruiting body with a translucent, gelatinous, branching, columella-like core; North and South America (Fig. 49) .. Protubera

Calvarula Zeller
Fruiting body subglobose, 10-15 mm in diameter, chalky white, staining
brown; peridium 2-layered, inner layer embedded in a gelatinous
matrix. Gleba a brown, powdery mass at maturity. Spores elliptical,
3-3.5 x 1.7-2.2 JLm (incompletely known), Zeller (1939).

Kobayasia Imai & Kawam.

Fig. 50

Fruiting body 3-6.5 (-12) cm broad, 2.5-4 (-8) cm high, with a creamcolored, flaccid exoperidium and a very closely adhering endoperidiurn. Gleba of greenish-gray, ovoid tissue embedded in chambers
within a clear, gelatinous matrix which forms a central branched
columella. Spores 4-5.5 x 1.8-2 JLm, cylindric, thin-walled, hyaline. Many
fruitings sampled but no odor detected. Known from Japan and Korea.
Monotypic: K. nipponica (Kobay.) Imai & Kawam.

Protubera Moller

Fig. 49

FrUiting body ovoid to lumpy, 2.2-3.5 (-7) cm broad, 3-4 cm high, with a
thin, dull white, flaccid exoperidium ( 1 mm) and a grayish endoperid
lurn, 3-4 mm thick. Gleba of olive-green chambers separated by a
Whitish, gelatinous tissue radiating from a columella-like, gelatinous,
central core. Spores 3.5-4.5 x 1.5-2 JLm, elliptical, smooth, nearly hyaline.
Smell indistinct or of rotting oranges or like passion fruit. Found in
North and South America.

84

Protuberella Imai & Kawam.

Fruiting body as in family and above genera except that the tissue is
"gelatinous fleshy in the peridium and not so distinctly gelatinous as in
Protubera maracuja, they are less gelatinous in the nature of the sterile
strands than those of P. maracuja" (Imai & Kawamura, 1958). Spores
3.7-5 x 2-2.5 Ilm, cylindric, thin-walled. Monotypic: P. borealis (Imai)
Imai & Kawam. Known only from Asia.

Rhopalogaster (Bosc) Johnston

Fruiting body epigeous, club-shaped, 3-7 cm tall, 1.5-2.5 cm broad,
brown to yellow-brown over the apex, peridium smooth, moist, and
rupturing irregularly at maturity exposing the gleba. Gleba chambered
and chambers lined with the hymenium, yellow-brown at first to dingy,
olive-brown and tough at maturity. Stalk a dry, white columella
extending to the peridium. Flesh firm, subgelatinous. Spores 3.5-4.3 x
5.8-7.2 Ilm, short-elliptical, smooth, yellow-brown. Basidia narrowly
clavate, thin-walled with 4 sterigmata apically oriented. No odor
recorded. Found in organic matter and mulch in the southern United
States, rare. Monotypic: R. transversarium (Bosc) Johnston.

4. Hysterangiaceae
Fruiting body hypogeous to epigeous, ovoid to pear-shaped or shortstalked, with a gelatinized, white columella or sterile stalk which
supports separate, olivaceous-gray to green, glebal plates. These plates
are oriented and attached to the inner peridium around the upper
portion of the fruiting body, which has proliferating, white rhizomorphs which become pink to lavender upon exposure to the air.
Odor at maturity fetid and phalloid-like. Spores small, cylindric to
ellipsoidal, olive to gray-green in mass. Cosmopolitan, with both
mycorrhizal and saprophytic species.

Hysterangium Vitt.
Fruiting body hypogeous, ovoid, buff to pink; peridium membranous
(not gelatinous); columella well developed, branching. Gleba green,
elastic, tough, gelatinized, firm. Proliferating rhizomorphs or a single
rhizomorph present. Odor fetid, stinkhorn-like in older specimens.
Spores elastic, tough, elliptical, with collar and gelatinous surface, 12-17
x 5-7.5 Ilm. Mycorrhizal.

85

phallogaster Morgan

Fig. 48

Fru iting body pear- to club-shaped with a short to well-developed

stalk, 20-55 mm high, 10-30 mm broad, flesh-colored to vivid pink with
ir regular, thin, peridial zones which soon collapse revealing the green
lobes of the gleba separated by gelatinized, some translucent, sterile,
hyali ne to white plates; with copious, basal rhizomorphs which turn
pink to lavender upon exposure to air. Fetid, phalloid odor at maturity.
Spores subcylindric, greenish, 4-5.5 x 1.5-1.8 jLm. Saprophytic. Monotypic: P. saccatus Marg.

5. Claustulaceae

Fruiting body up to 10 cm tall, with an ovate to obovate, soft, finely

chambered, white receptaculum which when expanded leaves a ruptured exoperidium that forms a typical volva. The volva splits to form
five pointed segments which are appressed to the receptaculum
forming a flaccid, fleshy base, 3-4.5 cm diam. The endoperidium is a
gelatinous layer which surrounds the immature receptaculum and
remains in the volva to form a gelatinous base. The gleba is a green
layer, dissected by a membranous, white trama, creating a layer
composed of small chambers attached to the inner surface of the
peridium and leaving a hollow interior. At maturity, the gleba breaks
up and becomes irregular and jagged. The immature, ovate buttons are
white in the soil but change to reddish-brown when exposed to the air.
Spores elliptical, with a short pedicel, smooth, 8-13 x 5-6 jLm. Known
only from South Island, New Zealand under Leptospermum scoparium
Forst. (excellent photographs by Curtis, 1926, PI. VI).

Claustu/a Curtis
(Characters of the family) Monotypic: C. fischeri Curtis.

6. Gelopellidaceae
fruit ing body up to 5 cm tall , ovate to obovate, flaccid with white, basal
rhizomorphs. Exoperidium a thin, dull white, tough layer over a thick,
translucent, gelatinous mesoperidium. Endoperidium a thin, white
layer separating the thick, olive-colored gleba from the gelatinous
rnesoperidium. The fruiting body appears similar to an unopened
MUtinus (see Fig. 47). Columella is central, fleshy, unbranched. Spores
4-~ (-6) x 1.5-2 jLm, cylindric and hyaline to elliptical and brown,
thIn-wailed. Five species found in South America, Africa, and Japan
(Zeller, 1939; Dring & Rose, 1977).

86

Gelopellis Zeller
(Characters of the family)

Fig. 43. Dictyophora duplicata. This

genus differs from Phallus only in the
development of an indusium (i). The
gleba (g) forms in shallow pits separated
by ridges (ri). A hollow (h) stalk (st),
open at the top, expands at maturity
from a flaccid exoperidium (p) and
gelatinized endoperidium (en), usually
referred to as a volva (v).

head

: - - -n

- -' g

-I

--h
--st

.-- p

r------ n

---g

'---- v

't--I

':5~-'~-;

.h+-- v

~"
.
. ' -.
;' .

.. . ..
,

"

'

..

Fig. 44. Dictyophora duplicata.

An almost fully expanded indusium (i) projects from beneath
the gleba (g) which is separated
by ridges (ri). The chambered,
hollow stalk (st), which is open
at the top (h), expands from the
volva (v) as the fruiting body
matures .


87

Fig. 45

ca

'-- g
e;'

;'., "i, '.

' ~,
.' \C.:' ;.~ . ~) ;' .
, i1!1,: .',:' " '"

~ .,

.:,

~.' :.~

Fig. 46

.... "..
I.:' '.

~ ,

~ . ,' :

. "<t
~.'.' ":' t_
"
.',:... ..'
.
. ', ,," "

.. . ;....".",
:

--re

','

"'......+-- st -~,
;C ,
o

"

;' '

.".

--g

"
,.-"

~
, ,:

, .

".~

,.
' ".

+--p

...
".,,'
~
.
.

:\

.'

..

+--h

,+--en
/--- v

' - - - - rh
rh-~
,

:--- rh

Fig. 45. /tajahya ga/ericu/ata. The fruiting body has a white calyptra (ca)
to w hich the gleba (g) is attached. A chambered hollow (h) stalk (st)
expa nds from the peridium (p) and endoperidium (en), both of which
together form a volva (v) with rhizomorphs (rh) at base.
Fig. 46. Staheliomyces cinctus. The gleba (g) forms a band, wh ich
appears constricted around the upper part of the receptaculum (re).
The white stalk (st) is hollow (h) and has rather large perforations. A
volva (v) , from which rhizomorphs (rh) proliferate, surrounds the
base .

88

--g

,......---p
fr--

~--h

me

~-g

r--en
g

!r----

st

--st

-----rh

f---V

Fig. 47. Mutinus ravenelii. The young specimen on left has an exoperidium (p), a gelatinous mesoperidium (me), and a white, membranous
endoperidium (en). The gleba (g) is preformed around the unexpanded
stalk (st) with rhizomorphs (rh) at the base. Older stages show the
hollow (h), expanded stalk (st), gleba (g), and volva (v).

g
.

.',

.!: ;

."

p-~-,

.. .
~

..,.','

.., ...:, . .
'

~-

'.

'

.". "

'.

. ..

g-+

.. .. " '
.'

. ",."

. '.
..

Fig. 48. Phalloga ster saccatus. The

gleba (g) occurs as a series of lobes
separated by sterile, gelatinized tissue and supported by an irregular
short stalk (st) with rhizomorphs
(rh) at the base. At maturity the
apex splits and the exoperidium (p)
is perforate revealing the gleba (g).
~---p

..

:',

~.

:".: ..

.'.
..

st--

..
.'

1-----

rh

89
.

..

.
... ....... . ....
. ... , '
...... . .:"'
.. .'., .
,' - .. ,': .
' . .

I~-

p
rh

IT-

en

I-g
~-

col

' - - - - rh

Fig. 49. Protubera jamaicensis.

The fruiting body has a columella (col) of white, gelatinous tissue which branches
upward between the olivegreen, glebal (g) chambers. A
flaccid, tough exoperidium (p)
and gelatinous endoperidium
(en) surround the gleba (g),
and stout rhizomorphs (rh)
(only one illustrated) proliferate into the soil.

...

.'... .
. . .'
..

t , .:..

.
.

.
"

"

.;; ;.t'
0'.

. . ..
.
.
,.
.
. . .

"r"--

.!'
.'

'.

'.,

'...

rh

en

~-g

Fig. 50. Kobayasia nippoilica.

A watery, gelatinous columella (col) branches repeatedly,
surrounding . the greenishgray glebal (g) chambers encl osed by a flaccid exoperidium (p) and closely adhering
endoperidium (en) of the
fr uiting body. Delicate rhizomorphs (rh) extend from the
base of the fruiting body.

col

..

90
g
Fig. 51. Simbfum sphaerocephafum. The fruiting body has an
oval, chambered head with depressions containing the gleba
(g). The head is supported by a
long, chambered stalk (st) or
receptaculum (re), which is
seated within the flaccid volva
(v) which has fine rhizomorphs (rh) extending from it.

.
, ..

.. 1

i: ' .
I

,.
!

' . .' .'". ~

' . -. " .
' . '..
. .. ", :,

'.

\"" .
\.
I

" .. . ..'... ...

- '" ..
.' r
.. :
' . '~."

'

' 0'

"'; '

:i .'... ...',......
I
\

,I

'~-I'

. ',

"

I
.......
:.
' :.,

. .. .
': ', ; ..: "

: : ':',..

--g

,.

.= ---- st

.- 4 -. v

,
."--=':~
.....
~
,
.".....
.",
'.

..

. z .. ~ . "..... .

. ".

----:':r-- v

'-----rh

Fig. 52. Lysurus borealis. A stout

stalk (st) or receptaculum (re) supports 3 or 4 short, thick receptacular arms (re) of the fruiting
body. The inner surface of each
arm is lined with gleba (g). The
stalk arises from a persistent
volva (v) .

re

"

'.

... . .. .. .
,

"

--re

~".'::

.' .
..
. . . .,:-

. .
'. ; ~""
. .

~--st(=

. '
'.

'

' . ''1
.'.!.

':

.: : :!l

. ....

'"

"

...' . '.

91

re

g
Fig. 53

-i------st

g--+

st - - - :fj'~

v-+.-'

: .

'j.

.. .-~
.. ';':.i':':
.. I ..
. ,'"
",

...

'(' .

,'

~_---v

'.:,
,

} : - - - rh

Fig. 53. Aseroe rubra. The fruiting body

has a stout stalk (st) which supports 6-8
ste llate arms (re) that are erect and position ed at the apex of the stalk (st). At
ma turity the receptacular arms (re)
ope n up to reveal the gleba (g) in the
center and on the surface of the arms.
The stalk arises from a persistent volva
(v) which has stout rhizomorphs (rh) at
the base.
Fig. 54. Kalchbrennera corallocepha/a.
The fruiting body has a single, chambered stalk (st) which produces a distincti ve upraised re'ceptaculum (re)
wit h scattered areas of gleba (g) over
the arms. The flaccid exoperidium (p)
remai ns as a volva (v) and from it rhizom orphs (rh) proliferate into the soil.

---st
Fig. 54

- - v (= p)

, - - - - - rh

92

Fig. 55. Pseudoco/us fusiformis.

The fruiting body has a short
stalk (st) which gives rise to 3-4
receptacular arms (re) lined
with gleba (g) and which arch
and come together at the apex.
The fruiting body is seated in a
flaccid valva (v) which surrounds the base and has numerous rhizomorphs (rh).

~-st

-+-v
~t---v

~---rh

,..---"g

re

it-:----

st

-f---- v

,...----- rh

Fig. 56. Co/us hirudinosus. This species has a short, thick stalk (st) from
which six columns form a red, arching receptaculum (re). The arms are
joined at the fertile, clathrate apex. The gleba (g) is formed on the
inside of the arms and the clathrate apex. A large flaccid valva (v)
surrounds the base and rhizomorphs (rh) proliferate into the soil. An
immature specimen is shown on the left.

93

;:;-~A- tr
~~~...--gl

,....:;...-+- la

t!'::"'!c:r
'--.
\
,I'

re

. .

"

<.

t '.

,'. ..

;~

.' ,. .........' .

-.... ..... .
'

~.

.. . .

. .'
,'

'

F-l-,--

Fig. 57. Laternea pusil/a. The glebifer (gl) is suspended below the
receptaculum (re) by trabeculae
(tr), forming the lantern (Ia). The
receptaculum is sessile within the
exoperidium (p) or volva (v) , and
rhizomorphs (rh) proliferate into
the soil.

(= p)

:.
:.:'('
' ,'

"

. .

----rh

~""<:---- gl
:........l

,-1,,.----

tr

,....--- la

re

. ..

Fig. 58. Laternea triscapa. The glebifer (gl) is held in place and clasped
by the trabeculae (tr), which form
th e lantern (Ia). The receptaculum
(re) is sessile withil1 the volva (v),
fro m which rhizomorphs (rh) proliferate into the soil.

. ..

.........;.......+----v
.

. .', .... .

"~i'. ~;,..

r-----

rh

94
,',. ,"

~:::::::,..---, , ~--

p
en

. . -1 . .

":'" ~..", /;"",,~

;
..:.,' .,,' ,
.~

+--re
""'- -1-- g

'r

"

' ) - - - - - rh

~,---

"

v (= p)

Fig. 59. Linder;a co/umnata. The young

.
- ...
stage on left shows the flaccid exoperidium (p), closely adhering endoperidium
(en), and the receptaculum (re) which is
~:::.... ""'- - - rh
~~~'
.--.......
embedded in a gelatinous matrix. The
gleba (g) develops in the center of the
fruiting body, and abundant rhizomorphs (rh) proliferate from the base.
Mature specimens usually have 3 to 4 stout arms forming the sessile
receptaculum (re) to which the gleba (g) adheres. The exoperidium (p)
'
forms the volva (v).
'"

"" "

,,

re
re

g
g

v ----,

P ----Ii

...;;.;..",t---

Fig. 60. Anthurus archer;. This species has a short stalk from which a
receptaculum (re) of 5-8 arms arches upward with the arms fused at the
apex. The older stage on left shows the characteristically recurved
receptaculum (re) with exposed gleba (g) and shallow volva (v). A
young, oval button with a flaccid exoperidium (p) is in foreground.

95

re

re

."L---

g
d----v

",

~:'

..
..
. ..:
.

'. ' ',~

'.
'~:'.'

...

".

. '.

rh
Fig. 61. Ifeodictyon gracilis. The fruiting body is sessile, with a clathrate
or latticed receptaculum (re), and gleba (g) on inner faces of the
gelatinous arms. Volva (v) and proliferating rhizomorphs (rh) are at the
base. Young stage on left shows the expanding, gelatinous receptaculum (re). A flaccid button is in the foreground.

...
..'.

.'" .
:.
,"

~-

~--rh

Fig. 62. Clathrus. ruber. The fruiting body is sessile with a clathrate
or latticed receptaculum (re) and
has the gleba (g) on the inner
faces of the hollow, chambered
arms. Rhizomorphs (rh) proliferate from the base of the volva (v).

96

l,

GAUTIERIALES

1. Gautieriaceae
Fruiting body hypogeous, ovoid, often with knobs and lumps, 0.5-5 (-6)
cm broad; exoperidium white, buff, to light brown, sometimes with
pigmented areas of red, blue, yellow, green, or brown in dried
specimens, thick to very thin or absent at maturity revealing the glebal
chambers which are minute, up to 6 mm in diameter, irregularly
lacunose, yellow-brown, greenish brown, to brown. Columella when
present white, soon finely branched (Fig. 63) or reduced to a basal pad
of sterile tissue or a thin, inconspicuous, dendroid columella. Spores
10-27 x 6-18 J.Lm, subglobose, ovoid, acuminate, to elliptical, longitudinally ridged, thick-walled with a prominent, orthotropic apiculus,
grayish-olive, yellow-green, dark green, orange-yellow, to brown or
blackish-brown. Widely distributed in Northern and Southern Hemispheres; mycorrhizal.
Key to the Genera of the Gautieriaceae

1. Fruiting body with a prominent columella; exoperidium thin or

evanescent; spores with longitudinal ridges which terminate below the

apex ............................................... Gautieria

1. Fruiting body either without a columella or with an inconspicuous,
dendroid columella; exoperidium thick; spores with longitudinal
ridges which converge at the apex .............. Austrogaut;er;a

Gaut;er;a Vitt.

Fig. 63

Fruiting body 1-5 (-6) cm broad, exoperidium buff to light brown, very
thin, often absent at maturity, revealing the characteristic, bright
yellow-brown to brown glebal chambers. Columella present, white,
soon finely branched. Rhizomorphs present. Spores 10-22 x 6-13 J.Lm,
longitudinally ridged, ridges terminating below the spore apex or tip,
thick-walled with a prominent, orthotropic apiculus, rusty brown.
Distributed in the Northern Hemisphere (Zeller & Dodge, 1918).

Austrogaut;er;a Stewart & Trappe

Fruiting body 0.5-2 (-3) cm broad, exoperidium thick, white to light
brown, often with pigmented areas of red, blue, yellow, green, or
brown in dried specimens. Glebal chambers minute, lacunose, yellow-

...
97

brown, greenish brown, to brown. Columella either absent and fruiting

body with a basal pad of sterile tissue, or present and thin, inconspicuo us, and dendroid. Rhizomorphs present. Spores 10-27 x (8-) 9-18 jJ.m ,
ovoid to acuminate, longitudinally ridged, ridges converging at the
spore apex or tip (best seen by means of scanning electron microscopy),
thick-walled, grayish-olive, yellow-green , dark green, orange-yellow, to
brown. Known only from Australia (Stewart & Trappe, 1985) .

, ~ ""

.,,}

,-

.: .:.',,'!t M~
41' ". "

! ... ~;

-~

l 'r-

I~:c 'Jtj;,.
'.... if,;,,;t<'
",:.:. -t:l,VfI ~.:
. .'t..

!,.J.. ~

~"-J' 'oJ
:
' ;'1..::..

.~

Y..i

>.,~-~-.........

.., ...,;.

..

..

-:

'..

"

............~. ,..

(;.

.... ....

.,.'J

,.'
"

'::'1"':":"-

.'

. ,."

""

'"i\" .'

: . "'

.....

..

'.: . 0
_ .'

. ' _

'/tIft
' .;
,a " 1,.., .
)0

"

".

...

\
,' ~
~

.J'

"., ... -.

' ~.,. ", -,.-:,:

,,

..

",'

',.
._,

'.
.

-:.

. ..

,-,

"to

~,"

'.

_. . ....... ~" .
", ' ..
. :"j~'J

...'

. "!'".c/_
..' ... .'. . ...
. ..

.".-. ,~ ...)~
\

'

..

.:.. :r.-. ".

. . '..,' ",:

."

' l '!'
'
~ ~

'r : ~

"

'

"lKv -#
-~

(l"

,',- -

- - col

Fig. 63. Gautieria graveo/ens. The fruiting body is irregularly ovoid,

firm, and with a very thin exoperidium (p) and irregularly lacunose
gleba (g) (above right). A short columella (col) branches and proliferates into the gleba (g).
--g

,"

. ...

f---P

.'

.'

--en

..

"

.,: .

....... . .
. .. ' .. .

.
." .
"

~--rh

"--- p

I----rh
Fig. 64. Me/anogaster variegatus. This species has an oval to lumpy
e xoperidium (p) and a closely adhering endoperidium (en). The gleba
(g) is black, gel-filled, and tough. Rhizomorphs (rh) proliferate from the
e xoperidium (p).

98

MELANOGASTRALES Svrcek
General: Fruiting body hypogeous, sometimes epigeous at maturity,

oval to irregularly shaped, often lumpy, tough and rubbery, dull white
to black. Gleba loculate and locules filled with a translucent gel, orange
or black, unchanging or changing to deep maroon-red when a fresh
surface is exposed to the air. Sterile base and capillitium absent.
Pseudostem present in Torrendia. Spores 4.5-22 x (1.8-) 5-18 (-18) ILm,
globose to elliptical, smooth, thick-walled, brown to dark brown, often
with a collar.

Key to the Families of the Melanogastrales

1. Either marine and fruiting body embedded within decaying wood
or terrestrial and epigeous at maturity, minute, and expanding to
form a powdery gleba, pseudostem and minute volva present at
matu rity ..................................... 2. Torrendiaceae
1. Terrestrial; hypogeous or epigeous at maturity with a tough, gelfilled, orange to black, loculate gleba; without a volva .......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1. Melanogastraceae

1. Melanogastraceae luI.

Fruiting body oval to lumpy, light brown, brownish-olive, to rusty

brown. Gleba lacunose, gel-filled, either black and unchanging or
yellowish-pink to yellowish-brown, darkening when cut to orange-red
and finally to deep maroon. Spores elliptical, thick-walled, base truncate with a distinct collar, no germ pore. Cosmopolitan, ectomycorrhizal with trees and shrubs.
Key to the Genera of the Melanogastraceae

1. Fruiting body oval, brown, with black gel-filled locules (Fig. 64) ...

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Melanogaster

1. Fruiting body oval, light brown, with yellow-brown to orange

gel-filled locules ...................................... Alpova

99

Alpova Dodge
Fruiting body oval, 0.5-6 cm broad, light brown to brownish-olive.
Gleba yellowish-pink to yellowish-brown, darkening when cut to
orange-red or deep maroon. Spores 5-8 (-15) x 3-4 (-5) p,m, elliptical to
cylindric, smooth, yellow to yellow-brown, hyaline, thick-walled. Hyphae with or without clamp connections.

Me/anogaster Corda

PI. III C; Fig. 64

Fruiting body oval to lumpy, 1-8 cm broad, brown to rusty brown.

Gleba black. Spores 4-18 (-22) x 3-10 p,m, elliptical to broadly elliptical,
thick-walled, dark yellow-brown to purple-brown. Hyphae with clamp
connections.
I

2. Torrendiaceae Malen~on
Fruiting bodies small, 5-10 mm broad, embedded within decaying
wood, subglobose to ovoid, remaining ovoid or expanding at maturity
to form a pseudostem and minute volva, gray-brown to orange. Gleba
within minute chambers which are not filled with gel, spores released
into a gelatinous matrix and dispersed in water. Capillitium absent.
Spores tetraradiate, sessile, and hyaline in Nia. Spores oblong or
elliptical with a basal collar and hyaline in Torrendia. Known from
North America, Europe, and North Africa. Decomposers. The two
genera are probably not related.
Key to the Genera of the T orrendiaceae

1. Marine; fruiting body embedded in ocean wood ............ Nia

1. Terrestrial; fruiting body hypogeous in soil when young, becoming
epigeous at maturity, expanding at maturity to form a pseudostem
and volva .......................................... Torrendia

Nia Moore & Meyers

Fruiting body within wood in the ocean, 5 mm broad, orange. Peridium
a single layer, splitting to release the gleba, which is a spore mass
embedded in a gelatinous matrix. Capillitium not present. Spores 11-12
x 7-8 p,m, ovoid to elliptical with 4 to 5 radiating appendages, hyaline.
Known from North America and Europe. Monotypic: N. vibrissa Moore
& Meyers.

100

Torrendia Bres.
Fruiting body hypogeous, oval, 5-10 mm broad, gray-brown; stalk 8-10
mm and expands and elevates the chambered head; some exoperidium
initially persistent, but splitting at the apex at maturity leaving a minute
volva surrounding the stalk base. Gleba a series of locules with a hymenium developed in each. Capillitium absent. Spores 13-17 x 5-7 J.Lm, elliptical , orthotropic, thin-walled with a basal collar. Clamp connections
present. Bas (1975) considers this genus to be a secotioid relative of
Amanita, and therefore mycorrhizal. Known from Europe and North
Africa. Monotypic: T. pulchella Bres.

101

LEUCOGASTRALES

1. Leucogastraceae Moreau ex Fogel

f ruiting body hypogeous, oval to uneven with lumps, white to buff, dry.
Gleba lacunose with round pores or labyrinthiform, gel-filled or
with a milk-like latex, unchanging when cut and exposed to the air.
Sterile base and capillitium absent. Spores globose, thick-walled, with
spinose to reticulate ornamentation, collar not formed. Ectomycorrhizal with trees and shrubs. Known from Europe and North America.
Key to the Genera of the Leucogastraceae

1. Spores globose, finely warted or with reticulate ornamentation;

gleba glistening white and gel-filled ................ Leucogaster
1. Spores globose, with long spines; gleba glistening white, gel-filled
or with a milk-like latex .......................... Leucophleps
Leucogaster Hesse

f ruiting body oval to irregular, 1-5 cm broad, white to buff in age, dry.
Gleba lacunose, with round pores, gel-filled, translucent white. Spores
10-21 JLm, globose, finely warted or spiny with a reticulate pattern of
ornamentation, hyaline, collar absent.

Leucophleps Harkness
fruiting body oval to irregular, 1-3 (-4) cm broad, white to buff, dry.
Gleba lacunose (Iabyrinthiform), 3-5 mm wide, gel-filled or with a
milk-like latex. Spores 10-20 JLm, globose with long spines, thick-walled,
hyaline, collar absent (fogel, 1979).

102

LOCULATE, SECOTIOID, AND AGARICOID

GASTEROMYCETES
(Note: see also section on Excluded Taxa, p. 122)
Key to the Genera of the Loculate, Secotioid, and Agaricoid
Gasteromycetes

1. Fruiting body with the stature of an unopened Coprinus comatus

(Fig. 65), with a stipe-columella and a loose, brown to blackishbrown powdery gleba at maturity; spores obovate (10-17 x 9-12 j.Lm),
thick-walled with a large apical pore (PI. III D) ................. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. see Podaxis (PODAXALES)
1. Fruiting body agaricoid to secotioid or bolete-like, with a lacunose,
poroid, or more frequently a lamellate to sublamellate hymenium
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2
2. Peridium and/or gleba with heteromerous trama; spores with
amyloid warts or reticulations; flesh often with a latex; fruiting
body terrestrial, epigeous to hypogeous, lacunose toagaricoid
in form (see Singer & Smith, 1960b; Miller, S., 1985; Smith, 1973)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. RUSSULALES (not included)
2. Peridium and/or gleba without heteromerous trama in combination with spores which have amyloid warts and ornamentation

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 3
3. Spores angular (PI. IV E), nearly cubical to nodulose ........... 4
3. Spores not angular, cubical, or nodulose ..................... 5
4. Fruiting body sublamellate with a stipe-columella; spores angular to nod u lose .............................. Rhodogaster
4. Fruiting body loculate with a small sterile point of attachment
but no stipe-columella; spores nearly cubical to angular ..... .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Richoniella
5. Fruiting body agaricoid, secotioid, or bolete-like, with or without a
columella; spores ornamented to smooth with or without an apical
pore; if loculate, spores not as in choice 5 below .............. 7
5. Fruiting body loculate with or without a reduced, branched, fleshy
columella; spores narrowly elliptical, smooth, without an apical
pore but often with a discrete collar (PI. IV F) .................. .
........................................ [Rhizopogonaceae 16

103

6. Fruiting body loculate, with a fleshy, rudimentary stipecolumella (Fig. 67) ....................... Truncocolumella
6. Fruiting body loculate, without a stipe-columella ........... .
. . . . . . . . .. . . . .. Rhizopogon (not included) (Smith et aI., 1981)
7. Fruiting body lacunose (loculate), gel-filled or not, columella lacking; spores orthotropic, verrucose (PI. IV B), rugulose, to echinulate, with or without an apical beak and apical pore; usually
hypogeous ................................................ 8
7. Fruiting body lacunose, agaricoid to secotioid, or bolete-like, with a
columella or stipe-columella; spores orthotropic or heterotropic,
ornamented or not, with or without an apical germ pore ...... 10
8. Gleba solid with gel-filled locules; spores verrucose, grayishyellow to light olive, cyanophilic ... Destuntzia (not included)
8. Gleba solid but not with gel-filled locules; spores verrucose to
rugulose or echinulate, white to brown in age, acyanophilic

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 9
9. Gleba white at first, becoming light yellow-brown in age, with a
watery latex; spores echinulate, non-amyloid .................. .
. . . . . . . . . . . . . . . . . . . . . ; . . . . . . . . . . . .. Octavianina (not included)
9. Gleba white at first, becoming brown in age, latex absent; spores
verrucose to rugulose with or without an apical beak (PI. IV B),
brown, non-amyloid ......................................... .
. . . . . . . . . . .. (see also Sclerogaster) Hymenogaster (not included)
10. Fruiting body bolete-like, tubular to loculate, with a columella
or a prominent stipe-columella; spores subfusiform to elliptical,
smooth with a prominent apiculus and no apical germ pore, or
with longitudinal ridges and with or without an apical germ
pore .................................................. 11
10. Fruiting body lacunose, agaricoid, to secotioid, with a columella
or prominent stipe-columella; spores variously shaped (but not
subfusiform to elliptical), smooth, rugose, or warted, with or
without an apical germ pore ............................ 12
11. Fruiting body hypogeous to partially epigeous, subglobose with a
small columella; gleba irregularly tubular and often with minute
chambers; spores with longitudinal ridges, occasionally with isolated warts, apical germ pore usually present ........ Chamonixia
11. Fruiting body epigeous, bolete-like with a prominent stipecolumella; spores smooth, no germ pore .................... 14

104

12. Fruiting body epigeous or partially hypogeous, agaricoid (Fig.

75) (related to the Gomphidiaceae); spores subfusiform to
broadly elliptical, germ pore absent; cystidia prominent; associated with members of the Pinaceae in the Northern Hemisphere ................................................ 13

12. Fruiting body epigeous, agaricoid, to secotioid (not related to

the Gomphidiaceae); spores variously shaped, smooth, rugose,
to warted, some with an apical germ pore; widely distributed in
many habitats ......................................... 15
13. Context pale orange with amyloid hyphae, fresh flesh purple in
ETOH; clamp connections only on amyloid vegetative hyphae at
stipe base (Fig. 75) ................................ Brauniellula
13. Context white with non-amyloid hyphae ........ Gomphogaster
14. Fruiting body epigeous, bolete-like; gleba of contorted tubes,
red to yellow, bruising blue; stipe-columella projecting below
gleba or not, surface smooth; spores elliptical to subfusifrom,
non-amyloid, no germ pore; cosmopolitan. . .. Gastroboletus
14. Fruiting body epigeous, bolete-like; gleba lacunose or with
contorted tubes or chambers, pale brown, not bruising blue;
spores elliptical to amygdaliform, non-amyloid; South America
and New Zealand, in Nothofagus forests ....... Austrogaster
15. Fruiting body epigeous, agaricoid; gleba contorted-Iamellate;
spores elliptical, rugose, brown (Cortinarius-like), apical germ pore
absent; ectomycorrhizal with North American conifers (Fig. 68) ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Thaxterogaster
15. Fruiting body epigeous, agaricoid to secotioid; gleba loculate to
lamellate; spores variously shaped, smooth to warted but not
rugose, apical germ pore present or absent .................. 16
16. Fruiting body epigeous, agaricoid, oval-depressed (pholiotalike); spores elliptical, thick-walled, germ pore absent; on
conifer limbs, twigs, and logs near or in melting snowbanks;
western North America (Fig. 71) ............. Nillatogastrium
16. Fruiting body epigeous, agaricoid to secotioid, spores variously
shaped; not on conifer wood near or in melting snow. . . .. 17

I'

,
1

105

17. Fruiting body epigeous, conical, apex conic to pointed and mucronate (Figs. 69-70); gleba contorted-Iamellate to intervenose, yellow, reddish, to rusty-brown, with soft tissue or collapsing and
deliquescent at maturity; spores elliptical or broadly elliptical,
smooth, thick-walled with an apical germ pore; decomposers of
grass or plant debris; found in either deserts or high prairies .. 18
17. Fruiting body epigeous, differently shaped or with a contortedlamellate gleba which is differently colored, often firm or becoming
tough and persistent in age; spores variously shaped, smooth to
warted, with or without an apical pore ...................... 20
18. On plant debris in hot, dry, high p ;'airies and deserts; pileus
narrowly conical, pointed, wrinkled and lacerate in age; gleba
yellow-brown, persistent (Fig. 69) . . . . . . . . . . . . . . .. Ga/eropsis
18. In grass, lawns (alpine meadows), and debris in temperate areas
during warm, wet periods .............................. 19
19. Gleba bright rusty brown; fruiting body collapsing and deliquescent at maturity; widely distributed ................. Gastrocybe
19. Gleba reddish-brown; fruiting body wrinkled and the margin
crimped when mature, but not deliquescent; in alpine meadows in
western North America (Fig. 70) ...................... Weraroa
20. Spores with an apical germ pore (PI. III D), smooth to warted,
thick-walled ........................................... 21

20. Spores without an apical germ pore, smooth, thin to thickwalled ................................................ 25

21. Spores globose to subglobose, thick-walled with prominent warts;

grassy areas in prairie habitats in North America .... Neosecotium
21. Spores . elliptical, smooth, thick-walled; in temperate, xeric, and
desert habitats ............................

0 -'

22

22. Fruiting body epigeous, coprinaceous; gleba black at maturity;

spores thick-walled with a broad apical germ pore (PI. III E) ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 23

22. Fruiting body epigeous, secotioid, not coprinoid, without

gussets in age; spores orthotropic or heterotropic, with or without germ pore, brown, yellow-brown, but not black ...... 24
23. Gleba forming gussets in age (Fig. 66); in xeric and desert habitats
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Montagnea

23. Gleba remains covered by peridium, gussets not formed; in disturbed areas in temperate habitats ................ Panaeolopsis

106

24. Spores 7.5-9.5 ILm long, small apical germ pore present but not
always visible (PI. III F); gleba chocolate-brown at maturity; terrestrial in xeric microhabitats in temperate areas or in arid,
desert habitats (Fig. 73) ................. . .... Endoptychum
24. Spores 11-141Lm long, apical germ pore visible on some spores;
gleba pale brown in age; terrestrial in sand steppes of South
Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Secotium
25. Volva present, fragile, shallow; gleba gill-like (Fig. 72), light gray to
black-brown in age; spores 5-7 ILm, globose to irregular, thickwalled; in arid regions or in xeric habitats ...... Gyrophragmium
25. Volva absent, not as above ................................. 26
26. Fruiting body epigeous, Agaricus-like (Fig. 74); gleba lamellate,
chocolate-brown to blackish-brown; flesh white, bruising yellow, giving a positive Schaeffer reaction; spores 4.5-7.5 ILm,
globose to subglobose; common in North America, in xeric

habitats .......................................... Longula

26. Fruiting body epigeous, agaricoid or Agaricus-like; gleba pale
brown; spores larger, elliptical; known only from the Southern
Hemisphere ........................................ , .. 27

27. Fruiting body Agrocybe-like with cystidia similar to those of an

Agrocybe; spores 9-11 x 5-6 ILm, elliptical, pale brown; New Zealand under Podocarpus ............................ Agrogaster
27. Fruiting body Agaricus-like; spores 13-16 x 9-10 ILm, broadly elliptical, pale brown; known only from Norfolk Island ............... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Gasteroagaricoides

107

PODAXALES

1. Podaxaceae
Fruiting body epigeous at maturity, 6-15 cm tall, head ovate-oblong to
obovate, 3-7 cm tall, 1-3 cm thick, dry, with appressed to slightly raised
scales, white to bay-brown (similar to Coprinus comatus in appearance). Stalk 4-8 cm long, 2-10 mm thick, equal, dry, with white,
imbricately arranged scales, fleshy when young but soon woody,
percurrent, with a bulbous base composed of hyphae and sand. Gleba
cottony, white at first, slowly turning yellow then yellowish-olive,
becoming reddish-brown to black in age, maturing from the base
upwards to the apex. A slight tendency to deliquesce in wet weather
like Coprinus. Paracapillitium deeply colored reddish-brown, infrequently septate. Spores 10-17 x 9-12 JLm, obovate, thick-walled, truncate
with an apical pore, with a short pedicel or none at all. Cosmopolitan,
decomposer of organic matter in xeric habitats and desert soils.

Podaxis Desv.

PI. III D, IV A; Fig. 65

(Characters of the family.) Monotypic(?): P. pistil/aris (L. ex Pers.) Fr.

HYMENOGASTRALES
(Note: see also section on Excluded Taxa, p. 122)

1. Rhizopogonaceae
Truncocolumella Zeller

Fig. 67

Fruiting body epigeous to partially hypogeous, obovate to irregular,

15-40 mm tall, 20-30 mm thick, surface smooth, dry, greenish-yellow,
yellow to dingy yellow in age. Gleba loculate with small, irregular
chambers, gelatinous to tough, olive-gray to gray-brown. Stipe-columella thick and fleshy over the narrow base, white, with proliferating
radial, fine branches extending throughout the gleba (Fig. 67). Spores
6.5-10 x 3.5-4.5 JLm, elliptical, smooth, thin-walled, yellowish, no apical
pore. Mycorrhizal with conifers in western North America. Monotypic:
T. citrina Zeller.

108

AGARICOID GASTEROMYCETES
Agaricales
Agrogaster Reid
Fruiting body epigeous, secotioid, up to 4 cm tall, pileus 1-1.5 cm
broad, conic to convex, smooth, waxy, sienna brown, drying lighter
brown. Gleba lamellate to contorted and chambered, pale brown,
attached to the pileus. Stipe-columella 1.5-4 cm long, 2-2.5 mm thick,
nearly equal, attached to the peridium, cylindric or tapering below.
Context white. Exoperidium cellular, composed of sphaeropedunculate cells, 8-30 tLm wide. Spores 9-11 x 5-6 tLm, elliptical, thin-walled,
non-amyloid, pale brown. Cystidia on tramal plates similar to those in
Agrocybe. New Zealand under Podocarpus dacrydiodes and P. spicatus.
Monotypic: A. coneae Reid (Reid, 1986).

fndoptychum Czern.

PI. III F; Fig. 73

Fruiting body 1-8 cm tall, 1-6 cm thick, oval, obovate or irregular in

shape with a very short stalk (Fig. 73). Pileus dry, white when young to
light brown, pinkish-brown, or chocolate-brown in age, with appressed
scales which are scattered or in regular rows, becoming shredded, torn
and splitting open. Gleba of irregular lamellate to anastomosing,
intervenose plates, becoming tough and brittle in age, white when
young to yellow-brown in age. Stipe-columella 1-3 cm long, 0.8-2 cm
thick, protruding a short distance or barely protruding below the
peridium, dull white, dry; bruises yellow in some cases. Spores 7.5-9.5
(-12) x 5-7 (-8.5) tLm, ovoid to broadly elliptical, smooth, thick-walled,
with a small apical pore, chocolate-brown. Decomposers in disturbed,
usually xeric areas. Endoptychum agaricoides Czern. is common and
widely distributed in the Northern Hemisphere (Singer & Smith,
1958b).

Galeropsis Velen.

Fig. 69

Fruiting body epigeous, 3-8 cm tall, pileus 1-3 cm thick, 2-4 mm broad,
narrowly conical to mucronate or pointed at apex, glabrous, base of
cap wrinkled and lacerate in age, ochre-gray to light gray-brown. Gleba
with numerous, contorted, anastomosed tramal plates, ochre to yellowbrown, attached only to the apex and appressed to the stipe-columella.
Stipe-columella 2-5 cm tall, 1-2.5 mm thick, enlarging somewhat toward
the base, dry, flexuous, fragile, dull white; volva absent. Spores 12-17 x
5-9 tLm, elliptical, smooth, thick-walled with an apical germ-pore,
.

109

yellow-brown. Decomposers of plant debris in warm, dry, high prairie

and xeric habitats. Ca/eropsis desertorum Velen. & Dvorak is in North
America, and several species occur in Africa.

Gasteroagaricoides Reid
Fruiting body epigeous, agaricoid, up to 5 cm tall, pileus 2-3 cm broad,
with convex, strongly incurved margin, dry, minutely asperate,
cinnamon-brown. Gleba lamellate but strongly intervenose, very thin,
pale brown, attached to the peridium. Stipe-columella 3 cm long, 1.5
mm wide, nearly equal, slightly larger at base, pale white, with a few
surface scales. Exoperidium with warts composed of globose, ovate, to
vesiculose, thin-walled cells. Spores 13-16 x 9-10 /J-m, elliptical, smooth,
hyaline, no germ pore. Taxonomic position unknown. Norfolk Island,
growing between rotting logs. Monotypic: C. ralstoniae Reid (Reid,
1986).

Gastrocybe Watling
Fruiting body 6-15 cm tall, pileus 5-30 x 2-10 mm, conic to narrowly
campanulate, striate, slightly viscid, light brown, collapsing and deliquescent at maturity (in several hours) to form a gelatinous mass. Gleba
irregularly lamellate with contorted tramal plates, bright rusty brown,
attached at apex and appressed to the columella. Stipe-columella 5-13
cm long, 1-2 mm thick, white, dry, shining, hollow, fragile and somewhat larger at base. Spores 10.5-12 (-14) x (6-) 7-8 /J-m, elliptical to broadly elliptical, thick-walled, with a prominent apical germ pore, yellowbrown. Pileipellis hymeniform. Found in temperate areas as decomposers in lawns and grassy areas under wet conditions during the
summer. Monotypic: C. lateritia Watling.

Gyrophragmium Mont.

Fig. 72

Fruiting body epigeous at maturity, round to egg-shaped when young,

10-22 cm tall when fully expanded. Pileus convex with a broad umbo
(Cetto, 1979, PI. 1198), 3-3.5 cm broad, 1.5-2 cm thick, smooth or
roughened in the center, light to dark brown, margin smooth to torn or
shredded from the separation of the partial veil which remains near the
center of the stalk as a ragged annulus. Gleba light gray at first, soon
black-brown, gill-like, contorted and anastomosing, free from the stalk,
attached to the lower side of the pileus, tough and brittle in age.
Stipe-columella 10-20 cm long, 7-22 mm thick, dry, dingy white to
yellowish, with a ragged annular zone and scattered loose fibrils or
scales below annular zone, base surrounded by a dish-like, fragile
membranous volva (Agaricus-like in appearance). Spores 5-7 /J-m, glo-

110

bose to ovoid or irregular in shape, thick-walled, no germ pore, dark

brown. Decomposers in arid regions and xeric habitats. Cyrophragmium delilei Mont. is cosmopolitan.
Longula Zeller

Fig. 74

Fruiting body epigeous, 5-18 cm tall with an oval to broadly convex

pileus, (4-) 5-7.5 cm broad, dry, white, with yellow-brown or browntipped, slightly raised squamules, often coarsely scaly flesh at first,
drying, splitting, and shredding in age over the lower part of the glel;>a,
leaving a ragged annular zone on the stalk and exposing the lamellate,
chocolate-brown to blackish-brown gleba. Gleba irregularly lamellate,
anastomosing, and forming cavities, white and fleshy but soon darkening to drab brown or brown, then to blackish-brown, becoming brittle
and tough, free from the stem. Stipe-columella 4-15 cm long, 1-2 cm
thick at the apex, broadening up to 4.5 cm at the base, smooth, white,
fleshy at first, becoming striate and woody in age and staining yellow
when bruised, with a mycelial covering around the base but no volva.
Strong, positive Schaeffer reaction on surface and flesh of the stalk.
Spores 4.5-7.5 x (4.5-) 5.5-6.5 J,Lm, globose to subglobose, thick-walled,
no germ pore, brown to black-brown. Common in North American
xeric habitats. Monotypic: L. texensis (Berk. & Curt.) Zeller.
Montagnea Fr.

PI. III E; Fig. 66

Fruiting body polymorphic, epigeous at maturity, 4-30 cm tall, head

conical to campanulate, dry, grayish brown; cap 1-6 cm broad with a
very thin endoperidium which breaks up and falls away at the margin
(Fig. 66) revealing the black, indurate tramal plates of the gleba. These
plates or gussets, on which the spores are borne, recurve, harden, and
become tough; desert wind-blown sand erodes away pieces of the
gussets, distributing the spores. Stalk 2.5-28 cm long, 3-9 (-15) mm thick,
white and fleshy at first, soon woody and dingy white to light brown
with scattered loose fibrils, sometimes with a ring of fibrils on the upper
stalk where the peridium was attached, base contained in a thick,
double volva consisting of an exoperidium, with embedded sand, and a
tough, fibrous mesoperidium. Gleba a black, tough, lamellate-like
tissue with a hymenium of collapsed basidia at maturity, and abundant
spores. Spores 15-28 x (7-) 9.5-14 I-tm, elliptical, thick-walled, orthotropic, with a broad, apical germ pore and prominent apiculus, black in
mass. Decomposer of plant organic matter under xeric conditions or in
desert regions. (Smith et aI., 1981; Cleland, 1934; H6110s, 1904; Pilat,
1958). Montagnea arenaria (DC ex Fr.) Zeller is known from North
America.

111

Neosecotium Singer & Smith

Fruiting body epigeous, 1-3 cm tall. Pileus 1-3 x 1-2 cm, nearly globose,
margin remaining firmly attached to the stipe-columella, smooth, dry,
avellanous to whitish. Gleba chambered with obscure lamellar arrangement at first, becoming somewhat powdery when mature, cinnamon to
light brown. Stipe-columella 2 cm long, 2-3.5 mm thick, percurrent,
dry, dull white, projecting slightly beneath gleba, context firm and dull
white. Spores 12-18 J,Lm, globose to subglobose, thick-walled (3-5 J,Lm)
with a hyaline perisporium covering a warted to deeply warted exosporium, with a distinctive, broad, hilar appendage (Singer and Smith,
1960; Figs. 2-4), red-brown in Melzer's solution. Clamp connections
rare. N. macrosporium (Lloyd) Singer & Smith is found in prairie and
xeric habitats among grass in sandy soil of central and southwestern
North America (Singer & Smith) 1960a).

Nivatogastrium Singer & Smith

Fig. 71

Fruiting body epigeous, oval-depressed, 1.5-4 cm tall, 10-24 mm thick,

broadly convex, smooth, slightly viscid, light yellow-brown to light
brown fading to dull white in age, margin appressed against the stalk,
sometimes pulling away to reveal the gleba which is composed of
cinnamon-brown to dark brown, contorted, anastomosing tramal
plates (Fig. 71). Stipe-columella 5-25 mm long, 5-15 mm thick, very short
to barely protruding, dry, fibrillose, dull white. Spores 7.5-10 x 5-7 J,Lm,
elliptical, thick-walled, smooth, no germ pore, yellow-brown. Decomposer of conifer woods, fruiting in snowbanks in western North
America. Monotypic: N. nubigenum (Hark.) Smith & Singer (Singer &
Smith,1959b).

Panaeolopsis Singer

Fruiting body 4-6.5 cm tall, pileus 10-15 x 15-20 mm, fusiform to

ellipsoid, surface glabrous, dry, gray to brown, with margin firmly
appressed to the stipe forming a sterile flap around it, at maturity
expanding slightly or not at all, margin lacking veil remains. Gleba of
crowded, ascending, grayish-black lamellae with a paler margin. Flesh
thin, white to pale brown. Stipe-columella 30-50 long, 2-3 mm thick,
cylindric, dry, solid, pruinose at first, smooth in age. Spores inequilateral, 12-14 x 8.5-9.5 x 7-7.5 J,Lm, broadly elliptical in face view, elliptic in
side view with large germ pore, black, not discoloring in concentrated
sulfuric acid. Cheilocystidia numerous, large. Pileipellis a polycystoderm. Decomposers in disturbed areas in South America (Singer, 1969)
and Australia (Watling & Young, 1983).

112

Rhodogaster Horak
Fruiting body epigeous, 2-3 cm tall, agaric-like with a pileus 1.2-2 cm
broad, subglobose, slightly depressed in the center, with a dry to
slightly viscid peridium, glabrous, sometimes radially rimose, whitish at
first, the center brown in age, becoming pinkish over the margin, no
veil. Gleba sublamellate to lacunate, up to 2 mm broad, attached to the
peridium or upper stipe, pinkish. Stipe-columella 2-3 cm long, 3-4 mm
thick, equal, solid, fragile, white, becoming brown, with dense fibrils,
separated somewhat from the gleba. Spores 9.5-11.5 x 6.5-7.5 /-Lm,
angular to nodulose, thin-walled, with a distinct apiculus, no germ
pore. On ground in the Pacific Rain Forest, Chile, South America
(Horak, 1964). Monotypic: R. chilensis Horak.

Richoniella Cost. et Dufour

PI. IV E

Fruiting body hypogeous to partially epigeous, (1-) 4-6 cm broad,

1.5-4.5 cm high, cream to ochraceous or darkening in age to tawny or
light brown, no columella or stipe. Peridium 0.1-0.4 mm thick. Gleba of
globose to labyrinthiform chambers, 0.5-1.5 mm wide, empty. Spores
10-12 x 7-10 /-Lm, angular to nearly cubical, smooth, tinted rose-colored
(PI. IV E). Odor indistinct or of potatoes. Richoniella asterospora (Coker
& Couch) Zeller & Dodge is known from North America under
hardwoods (Quercus) under the name Nigropogon. Other species are
present and widely distributed in Europe and New Zealand.

,
.

Secotium Kunze
Fruiting body epigeous, 3.5-11 cm tall, agaric-like. Pileus 3-6 x 3-4 cm,
conic to campanulate, surface dry, white to buff, sometimes not
completely covering gleba over the margin, which is deeply sinuate;
veil not described. Gleba with irregular, contorted, non-Iamellate
chambers, white to buff or pale brown, brittle, not powdery, surrounding but not attached to the stipe-columella. Stipe-columella 4-7 cm
long, 5-10 mm thick expanding to 2-2.5 cm at the base, smooth, white,
volva present and remains as a cothurnate basal zone, context fleshy
and white. Spores (8.5-) 11-14 x (6.3-) 8.2-9.7 /-Lm, short-elliptical but
ovate in face view, thick-walled, apical pore sometimes present, weakly
metachromatic in cresyl blue. Clamp connections present (Singer &
Smith, 1960). Terrestrial in sand steppes of the Cape of Good Hope,
South Africa. Monotypic: S. queinzii Kunze.

113
Thaxterogaster Singer

Fig. 68

Fruiting body hypogeous when young, epigeous at maturity, 3-7.5 cm

tall, pileus 1-5 x 1-3 cm, convex, convex-depressed, to convexumbonate, slimy to dry, glabrous, olive-yellow, yellow-brown, to
brown. Gleba lamellate, often contorted with anastomosing veins and
locules, cinnamon to dingy brown when mature, attached near apex of
stipe-columella; peridium separating and pulling away somewhat at the
base of the gleba. Stipe-columella 1-4.5 cm long, 8-30 mm thick, fleshy,
white, slimy when moist, with cortinate partial veit fibrils often seen as
peridium separates from the columella, volva absent. Spores 14-17 x
8-9.5 p.m, elliptical to broadly elliptical, rugose or warty and wrinkled,
no apical germ pore, brown. Ectomycorrhizal with conifers and
angiosperms. Thaxterogaster pingue (Zeller) Singer & Smith is
distributed in western North America; other species are known from
South America, Australia, etc. (Singer & Smith, 1963).

Weraroa Singer

Fig. 70

Fruiting body 6-16 cm tall, pileus 9-44 x 4-15 mm, narrowly conical,
acuminate or mucronate-pipped at apex, with remains of a fibrillose
veil, otherwise glabrous, dry, with a sheen, wrinkled and cracking over
the crimped margin of the pileus, surface orange-yellow, mustardyellow to yellow-brown, context brown. Gleba of contorted, anastomosing ridges, and intervenose, reddish-brown unexpanded lamellae
appressed to the columella. Stipe-columella 5-12 cm long, 1-4 mm
thick, slightly enlarged toward base, flexuous, dry, buff, yellow-brown
to brown; volva absent, with fine rhizomorphs at the base. Spores 11-14
(-15) x 6-7 (-8) p.m, ellipsoid to almond-shaped, thick-walled, smooth
with a prominent apical germ pore, dark reddish-brown in mass.
Decomposers of grass in alpine meadows of western North America. W.
cucullata (Seaver & Shope) Thiers & Watling is the only species in in
North America. Four other species are reported from New Zealand,
Australia, and South America (Singer & Smith, 1958c; Singer, 1958).

Boletales
Austrogaster Singer
Fruiting body epigeous, boletoid, up to 6 cm tall, pileus 3-5 cm broad,
2-3 cm high, convex to conical, pale brown, dry, sometimes areolate to
rimose. Gleba adnate or decurrent, lacunose or with contorted tubes or
chambers 1-2 mm in diameter, closely appressed or attached to the
columella, pale brown, brittle and not powdery at maturity. Stipe-

114

columella 5-6 cm long, 1-1.5 cm wide, nearly equat attached to the

peridium, dry, somewhat lacerate in age, pale brown, gleba exposed

near the stipe or closed at maturity. Spores 9-14 x (5.5-) 6-7.2 p,m,
elliptical to amygdaliform, slightly thickened, orthotropic with a central
or lateral apiculus, non-amyloid, under SEM surface roughened. Related to Paxillus (Singer, 1962; Reid, 1986). South America and New
Zealand in Nothofagus forests. Three known species.

Braun;ellula Smith & Singer

Fig. 75

Fruiting body epigeous or partially hypogeous, 2-7 cm tall. Pileus 1.5-4.5

cm broad, ovoid to convex, moist, glabrous, ochraceous-orange, attached to stipe by a fibrillose veit opening slightly or not at all. Gleba
contorted, sublamellate, anastomosing, light gray, soon darkening to
drab gray or smoke-gray at maturity. Stipe-columella 2-3 cm long, (5-)
7-10 mm thick, equal or slightly narrowed at base, dry, glabrous or with
appressed fibrils, ochraceous to vinaceous-red in age. Context pale
orange with amyloid hyphae in Melzer's solution, fresh flesh purple in
70% ETOH. Spores (15-) 16-20 x (7-) 8-9.5 p,m, broadly ellipticat thickwalled, smooth, no germ pore. Clamp connections only on amyloid,
vegetative hyphae at stipe base. Mycorrhizal with conifers in western
North America. Monotypic: B. albipes (Zeller) Smith & Singer (Smith &
Si nger, 1958).

Chamon;x;a Rolland
Fruiting body hypogeous to somewhat epigeous, in litter or leaf mold,
globose to subglobose, 1-5 cm broad, often in clusters, attenuated or
folded below to a reduced columella or fleshy point of attachment
which is white or light in color. Peridium dry, smooth or with some
rhizomorphs appressed to the surface, convex or flattened, brown,
olive-brown, to olive-ochre, sometimes staining blue when bruised.
Gleba of minute chambers within a thin, branching columella (as illustrated in Fig. 67L sometimes extending to the peridium but variable
among the species, irregularly tubular in construction, empty but lined
with the hymenium, white, yellow, reddish-brown, cinnamonbrown, to chocolate-brown usually darkening in age; sometimes staining blue when cut. Spores 9-18 (-25) x 9-16 p,m, broadly ellipsoid to
ovate, with (or without?) an apical pore, episporium thick with longitudinal ridges or sometimes isolated warts, orthotropic with a prominent hilum, brown, non-amyloid. Associated with the Pinaceae (Picea
and Abies) and the Fagaceae (Quercus agrifo/ia) in North America and
Europe. Smith & Singer (1959) reported and discussed four species from
Europe and North America.

115

Gastrobo/etus Lohwag
Fruiting body epigeous, bolete-like in aspect, 7-12 cm tall. Pileus 2-5 (-8)
cm broad, broadly convex, 2-4 cm thick, surface dry, brown, redbrown, to golden yellow, glabrous or obscurely squamulose. Gleba of
contorted tubes and pores, readily separable from the peridium, free
from the upper stipe, deep red or yellow, some turning blue when
bruised. Stipe-columella 2-7 cm long, 8-25 mm thick, equal or tapered,
dry, yellowish to reddish over the lower part, without a veil, even,
slightly projecting or elongating well beyond the gleba. Context firm,
yellow in peridium, pinkish in stipe-columella, or deep red at base.
Spores (9.5-) 13-20 x 5.5-9.5 JLm, broadly elliptical to subfusiform, thickwalled, with a prominent apiculus, germ pore absent, golden brown.
Clamp connections absent in fruiting body. Mycorrhizal in deep
conifer duff, cosmopolitan. Gastrobo/etus turbinatus (Snell) Smith &
Singer is common in North America (Thiers & Trappe, 1969).

Gomphogaster (Smith & Singer) O. K. Miller

Fruiting body epigeous at maturity, 16 mm tall. Pileus 15 mm broad,
convex-depressed, dry or subviscid, pale vinaceous. Gleba chambered
and with contorted tramal plates, drab gray, attached to pileus and
upper stipe. Stipe-columella 12 x 6 mm, dry, dull white with a lemonyellow base, no veil. Context white, firm, non-amyloid in Melzer's
solution. Spores 17-22 x 8.5-10 JLm, elliptical, thick-walled, smooth, no
germ pore. Clamp connections absent in fruiting body. Mycorrhizal,
probably with Pinus contorta in western North America, rare. Monotypic: G. leucosarx (Smith & Singer) O. K. Miller (Miller, 1973).

116

~--p

stc

-->;

- - stc

Fig. 65. Podaxis pistillaris. The fruiting body has raised scales over the
exoperidium (p), cottony gleba
(g), and a tough, almost woody,
stipe-columella (stc) at maturity.

p
-----:T

Fig. 66. Montagnea arenaria.

The fruiting body on left has
the conic exoperidium (p) in;
tact, a well-formed volva (v),
and fine rhizomorphs (rh).
The exoperidium (p) collapses, except in the center
and i nd urate tramal plates
form distinctive gussets (gu)
at maturity.

gU

gu

rh - - - ; 1
v

117

. . ..'.....
. :'?\
. '.
'

','

..
..

"

~-- . g

.
,
.. ',.
.'.; .
'

: .. .
... ..,
.. .,". .,
'"

..

=---- stc

Fig. 67. Truncoco/umella citrina. The fruiting body is obovate to

irregu lar with a smooth exoperidium (p), loculate gleba (g) , and a fleshy
stipe-columella (stc) from which fine branches radiate throughout the
gleba.

p
,

>

it: .
.j:

'.;",. .

..

-If

.:
; ".

','.

. .

-g

:;

. . -.:........;,.---- stc

..1-1

.",

t! !
;-"

ve

Fig. 68. Thaxterogaster pingue. This species has a convex pileus with a
smoot h exoperidium (p) and a fibrillose (cortinate) veil (ve) which is
best seen in the button stage, as illustrated on left. The gleba (g) is of
brown, anastomosing, contorted lamellae with a stout, firm stipe-colume ll a (stc).

118

It - - -

Fig. 69. Ga/eropsis desertorum. The

fruiting body has a conical pileus
with a wrinkled to smooth exoperidium (p) , contorted tramal plates
which form the gleba (g), and a wellformed stipe-columella (stc) which
supports the fruiting body.

stc

.::

( , ,.
'

"

,.. "."i: .~

1---

Fig. 70. Weraroa cucullata.

The fruiting body has a narrowly conical pileus with a
pointed apex, fragile exoperidium (p) , flexuous stipecolumella (stc), and gleba (g)
of contorted, anastomosing
lamellae.

stc

/
stc-H
:'

119

~--p

.
. : I,:

"

..

. ..

'

.
.
I

. . ........
, "

'-/--- g

, .--';;'-",

.. '

stc

Fig. 71. Nivatogastrium nubigenum. The fruiting body is convex and has
a smooth exoperidium (p), a gleba (g) of anastomosing tramal plates,
and a thick, short stipe-columella (stc).

Ar---

Fig. 72. Gyrophragmium delilei. The fruiting

stc

.-r-v

body has a convex-umbonate pileus with a

smooth exoperidium (p), gill-like, anastomosing gleba (g), fibrillose stipe-columella (stc), and
a shallow, fragile, membranous volva (v) .

120

---p

, '.

"'-rJ~"..
",

...--- stc

.~,.

."

. .. . .

. :; . ..
"

r---

stc

Fig. 73. Endoptychum agaricoides. The fruiting body has a conical to

convex pileus with an exoperidium (p) of appressed scales, a short,
thick stipe-columella (stc), and an irregular, lamellate gleba (g).

g
. ,"""'~

'.

:.-. ' .1
r-P
.., ..::Jf;
,: rf'l;~~'".
:.,~.,

..

Fig. 74. Longula texensis. The

fruiting body has an oval
pileus with a wrinkled, squamulose exoperidium (p), and
gleba (g) of blackish brown,
brittle gussets (gu). The stipecolumella (stc) is tough and
longitudinally striate in age.

,,

;,. '
.

'---- stc

...

~~',-_3 .

121

~---p

I !.
,'

,, ,

. ,.

.. .
~

.'

-\-- g
"

!.
I ,'

, .

'

stc

'.

..

; I '

. '
;

', '

stc

;' .

ve

Fig. 75. Brauniellula albipes. The fruiting body has an ovoid to convex
pile us with a smooth exoperidium (p), gleba (g) of anastomosing
lame llae, and a robust stipe-columella (stc). Note the fibrillose veil (ve)
between the stipe and margin of the pileus on the specimen on the
righ t.

122

EXCLUDED TAXA
The emphasis in covering the extensive number of taxa of loculate,
secotioid, and agaricoid genera of Gasteromycetes was on epigeous,
common taxa in the Northern Hemisphere and those which can be
encountered frequently in particular habitats. A few hypogeous taxa
are included, such as Richoniella (Nigropogon) , because of a unique
characteristic, such as the spore morphology, or their position near a
related, epigeous taxon. The large number of taxa in the Russulales or
the Astrogastraceous series of Singer and Smith (1960) is beyond the
scope of this work; these taxa are being studied by Pegler and Young
(1981), S. L. Miller (1985), Thiers (1979), and others. Much of the
Rhizopogonaceae is not included. Rhizopogon, Hymenogaster, and
Sclerogaster are hypogeous, loculate Gasteromycetes and are at least
partially covered by Smith et al. (1981) and others in regional works or
are in need of revision.
The Cribbiaceae as presented by Smith (1973), including Cribbea
(Smith & Reid, 1962), Singeromyces Moser apud Horak and Moser
(Horak & Moser, 1965), Paxillogaster Horak and Moser (Horak &. Moser,
1965), and Setchelliogaster Pouzar (Singer & Smith, 1959) are not
covered. In one way or another information is missing which would be
necessary to illustrate and describe them adequately. Many are found
in the Southern Hemisphere (Singer, 1962). Collections are not readily
available, and some are known only from the holotype or type locality
in exotic situations. They are rare or at least infrequently collected, and
the group needs critical restudy.
In addition, several secotioid, hypogeous taxa of uncertain position
are not included. Protogautieria A. H. Smith (Smith, 1965), Brauniella
Rick. ex Singer (Singer et aI., 1963), Hypogaster Singer (Singer, 1964),
Gymnopaxillus Horak (Horak, 1966), Smithiogaster J. E. Wright (Wright,
1975), and Hypogaea Horak (Horak, 1964) were excluded for the
reasons mentioned above. Most genera of the latter group are probably related to the Agaricales, Boletales, or other Hymenomycetes, but
their exact position is uncertain. Little or no SEM (scanning electron
microscopy) or TEM (transmission electron microscopy) studies of
spores have been carried out, or other critical data are missing.

I
I

123

GLOSSARY
acolumellate (of gleba): refers to a fruiting body which lacks a basal
columella, e. g. in Astraeus, Fig. 36.
acuminate: refers to gradual narrowing to a tip, as in the apex of the
pileus in Weraroa, Fig. 70.
acyanophilic: refers to the negative reaction of the cell wall or spore
wall, which does not stain blue in 1% cotton blue in lactic acid.
agaricoid: . refers to a fruiting body with the aspect of a lamellate
mushroom, but with strong evidence of an affinity to a particular
family of the Agaricales (see secotioid). Figs. 71,72,73.
angiocarpic development: a type of development in which the basidia
mature while enclosed within the fruiting body, typical of the
Gasteromycetes.
apophysis: a swelling, such as the ventral swelling of the endoperidium
of Geastrum, Fig. 19.
asperate: having a surface roughened with small granules, e. g. spore
sac of Lycoperdon pulcherrimum, Fig. 7.
aulaeate development: hymenium formed on the surface of downgrowing primordial tissue which arises from the peridium, Fig. 1 D.
aulaeothecium: the downgrowing primordial tissue formed during
aulaeate development.

ballistosporic: refers to a spore which is forcibly discharged from the

basidium. Gasteromycetes, by definition, do not have ballistospores.
binding hyphae: highly branched, thin- or thick-walled, cylindric
flexuous cells which intertwine, creating a firm but often flexible
tissue.
bovistelloid: refers to a type of forate development which produces a
sterile base and a capillitium of discrete, tapering units, Fig. 3 A.
bovistoid: refers to a fruiting body which is usually ovoid and lacks a
sterile base, but has a capillitium of discrete, branched, tapering
units, Fig. 3 B.
button: the youngest, usually unopened, immature stage in the development of the fruiting body, Fig. 31, right.
byssoid: refers to a delicate pattern of radiating bundles of hyphae (=
fibrils), characteristically found in the gleba of Sclerogaster.

124

caducous: refers to a structure which sloughs off or falls away, as

happens to the exoperidium of Lycoperdon at maturity, Fig. 6.
.
calyptra: tissue which covers the apex of the fruiting body of /tajahya
ga/ericu/ata and to which the gleba is attached, Fig. 45.
capillitium: coarse thick-walled cells found in the gleba, which mature
late and may develop pores or slits in the thick secondary wall,
usually branched, PI. VII.
centrifugal: maturing or developing from the center outward.
centripetal: maturing or developing from the outside inward.
circumscissile dehiscence: opening or separating on a circular or
equatorial line, e. g. in Battarrea, Fig. 29.
clamp connection: a small hyphal branch which grows back from the
hyphal tip to join the hypha I cell. The passage created allows one
of the nuclei resulting from mitosis to move into the dorsal cell, PI.
VII B.
clathrate (clathroid): refers to a latticed or interconnected receptaculum which supports the gleba, e. g. in C/athrus, Fig. 62.
collar: a ridge of tissue which surrounds the pedicel in some species of
Geastrum, Fig. 18, or forms the "little cup" referred to by Coker
(1928) on the basidiospore of Rhizopogon and shown here for
Hysterangium, PI. IV F.
columella: sterile tissue, usually in the base of the gleba, extending up
into or through the gleba (see dendritic or percurrent columella
and stipe-columella, Fig. 73).
connivent: see fibrils, Fig. 4 B & c.
coronate: splitting to form a crown-like structure, like the exoperidium
of Sphaerobo/us, Fig. 42.
cortex: a general term for the outer layer or layers of a sporocarp,
including the peridium; in Nidulariales, the main wall of the
peridiole.
cortinate: having a cobweb-like, fibrillose partial veil; typically found in
the Cortinariaceae and in Thaxterogaster, Fig. 68.
cothurnate: having a tissue which sheathes the base of the stipe and
usually has a thickened roll of tissue (like a rolled sock) at the top,
p. 112.
cyanophilic: refers to the positive reaction of the cell wall or spore wall,
which stains dark blue in 1% cotton blue in lactic acid.
deciduous warts: warts which can become readily detached from the
exoperidium, usually leaving a visible scar, e. g. in Lycoperdon,
Fig. 5 A & B.

125

dehiscence: the splitting of the peridiumin mature fruiting bodies in

order to expose the gleba and facilitate the discharge of the

spores, Fig. 10 (in general dehisce to split open).
delimited peristome: a zone around the peristome which is distinct
from the exoperidium, usually small and sometimes slightly raised,
often different in color, Fig. 21 (see also disc).
deliquescence: the process of dissolving into an "inky" fluid, common
in Coprinus.
dendritic columella: sterile tissue with repeated branches into the
gleba, usually from a solid basal tissue, Fig. 67.
diaphragm: a homogeneous wall of hyphae which separates the gleba
from the sterile base, e. g. in Vascellum, Fig. 11.
dimitic system: used to describe the development of true capillitium
and paracapillitium within the same gleba.
disc: a delimited peristome.
earth ball: common name for the members of the Sclerodermatales,

which include Scleroderma and Sclerogaster.

echinulate: covered with small spines, PI. II F.
elaters: spirally thickened capillitial elements in the gleba of Battarraea,
PI. VI C & D.
emplacement: a cushion or pad of closely packed hyphae beneath the
sporocarp, usually incorporating bits of substrate; occurs in the
Nid u lariales.
endoperidium: inner wall of a multi-walled peridium.
endosporium: the innermost wall layer of the basidiospore, adjacent to
the cytoplasmic membrane.
epigeous (epigaeous): growing or developing on the surface of the
ground.
epiphragm: an apical portion of the peridium, a lid, typical of the
Nidulariales, which detaches at maturity exposing the mature
peridioles, Fig. 40 .

eplsporium: wall layer of the basidiospore just beneath the exospor

lum.
evanescent: thin and fragile, as the exoperidium of Gautieria, Fig. 63.
exoperidium: outer wall of a multi-walled peridium.
exosporium: outer layer of the basidiospore wall after the perispore

disappears, usually forming the spore ornamentation, PI. II E.

farinose: covered with meal-like particles.
fibrillose-truncate wart: a flat-topped wart adorned with fibrils, Fig. 4 F.

126

fibrils: exoperidial ornamentation in the form of thin, hair-like structures, typical of Lycoperdon and Calvatia, Fig. 4 A, B & c.
fimbriate: delicately toothed, often used in reference to a stoma, Fig.
21.
forate development: hymenium formed by the differentiation of primordial tissue from the peridium, Fig. 1 C.
fornicate ray: an arched ray formed when the mesoperidium separates
from the exoperidium, adhering only at the margin; the ray arches
to support the elevated endoperidium, e. g. in Geastrum, Fig. 17.
funicular cord: spirally twisted or braided cord within the funicular
sheath in some species of the Nidulariales, Fig. 41.
funiculus: a structure composed of a sheath, middle piece, and cord,
which connects the peridioles to the endoperidium; typical of the
Nidulariales, Fig. 41.
gastroid: general adjective applied to all Basidiomycetes which have
angiocarpic development.
gelatinous peridium: a peridiLlm composed of a hyaline, pectinous
matrix with embedded hyaline hyphae or hyphal fragments,
infrequent, e. g. in Calostoma, Fig. 31.
gelatinous tissue: a translucent, pectinous matrix with or without
embedded hyphae, common in the Phallales, e. g. Kobayasia, Fig.
50.
generative hyphae: simple, thin-walled, regularly septate cylindric
hyphal cells which differentiate to form the hymenium and
related tissues.
germ pore: a pore which develops in the spore at the end opposite the
apiculus or hilar scar, e. g. in Podaxis, PI. III D and Montagnea, PI.
III E.
gleba: the spore-bearing tissue in an angiocarpic sporocarp, which
includes sterile tissues such as capillitial elements which form as a
sporocarp matures, Fig. 5.
glebal chambers: cavities within the gleba usually lined with hymenial
elements, either empty or gel-filled, e. g. in Me/anogaster, Fig.

64.
glebifer: a special hymenial structure borne in the arch of the receptaculum, called a "lantern". The glebifer is held in place by either a
piece of tissue or by trabeculae which anchor it to the arch, as in
Laternea, Fig. 57.
gussets: regions of the tramal plate which thicken over the point of
attachment or anastomose with the peridium or another tramal
plate. The individual upraised, mature plates of Montagnea, Fig.
66 are gussets.

127

hanging collar: a ridge of tissue surrounding the stalk and attached to

the bottom of the endoperidium, typical of some species of
Geastrum, Fig. 18.
hapteron: the ball of hyphae to which the funicular cord is attached in
Cyathus, Fig. 41.
heteromerous: refers to tramal tissue which is composed of clusters of
globose to ovoid cells intermixed with filamentous hyphae, especially noted in the pi leal trama. Typically found in the Russulales
(p. 30).
heterotropic spore: a spore which is borne somewhat obliquely to the
sterigma. This type of attachment facilitates spore discharge.
hilar appendage: the apiculus, hilum, or small knob-like point of
. attachment of the spore to the sterigma of the basidium, PI. III F.
hilar scar: a slightly recessed area on the spore where it is attached to
the sterigma of the basidium. Common in the Phallales.
homogeneous development: differentiation of the basidia in primordial tissue evenly throughout the gleba; typically found in species
of Bovista, Fig. 3 B.
hygroscopic: applied to an exoperidium which recurves to enclose the
endoperidium when dry but opens when moistened to expose
and elevate the inner spore case.
hymeniform: refers to a layer of narrowly clavate cells which appear
similar to basidioles which make up the hymenium.
hypha (ae): a single fungus cell. Several hyphae are shown in Plate VIII
A&C.
hypogeous: growing or developing below the surface of the ground.

indehiscence: the condition of not rupturing or splitting at maturity

(see dehiscence), often referring to the peridium.
indurate ~rama: trama which grows hard or hardens with age, such as
the tramal plates in mature Montagnea sporocarps, Fig. 66.
ind usium: a veil which flares out in a pendant fashion from the base of
the receptaculum in Dictyophora, Fig. 44 .

Invagination: an inward growth from the peridium, as in forate development, to form multiple cavities in which the clusters of basidia
are found, Fig. I C.
Involute ray: a ray which in profile has the outer peridium concave and
inrolled at the edge, as in Astraeus, Fig. 36.
o

128

labyrinthiform: maze-like with irregular chambers, like the glebal

chambers of Gautieria, Fig. 63.
lacerate peristome: a peristome with an irregularly torn edge around
the stoma (mouth), e. g. in Geastrum, Figs. 17 & 18.
lacunar development: formation of regular chambers lined by the
hymenium (see locular), Fig. 1 B.
lacunose: composed of or covered with pits, as in Calostoma basidiospores, PI. . III A; also refers to the chambered gleba in the
Hymenogastrales, as in Gautieria, Fig. 63.
lantern: an ovoid, deep-orange glebifer suspended from the columnar
receptacle and held in place by trabeculae, often numerous. At
maturity coated with a blackish-green basidial layer, as in Laternea, Fig. 57.
loculate: composed of chambers as in lacunar.
lumen: the central portion of the hypha I cell within the cell wall. PI. VII I D.
Iycoperdoid: refers to a fruiting body having a gleba which is powdery
at maturity and usually supported by a sterile base, Fig. 2 A.
mesoperidium: middle wall of a multi-walled peridium.
metachromatic: refers to e. g. a cell wall or inner spore wall which
stains violet to reddish in cresyl blue.
middle piece: a stout, short tissue which anchors the purse to the
sheath in some species in the Nidulariales, Fig. 41.
mucronate: having a sharp, often abrupt point, as in the pileal apex of
Weraroa and Ga/eropsis, Figs. 69 & 70.
multipileate development: development in which many individual
stalks with small pilei, each with its own hymenial tissue, form
from a central stalk, Fig. 1 F.
mycelium: the mass of hyphae forming the asexual body of a fungus,
the vegetative thallus.
.
myco: prefix meaning "pertaining to fungi".
mycorrhizae: the symbiotic combination of the hyphae of certain fungi
with the roots of a plant.
mycosclerids: irregular, thick-walled skeletal hyphae, occasionally present in the peridiallayers or around the ostiole in Vascellum.
myriostomes: many preformed pores, as in the endoperidium of
Myriostoma, Fig. 38.
naked: applied to a stoma which does not have a differentiated
peristome, e. g. in Tu/ostoma, Fig. 23.
non-persistent attachment: occurs in fruiting bodies which break the
attachment to the ground and are free at maturity to be blown
about by the wind, e. g. in Bovista.

..
129

obovate: oval with a broader apex, like an egg standing on the narrow
end.

one-spored basidium: a basidium which develops a single sterigma on

which a single spore develops.
orthotropic spore: a spore which is borne erect and centered on the
sterigma. Discharge is passive and a spore collar is often visible at
the point of detachment from the sterigma, PI. III D.
ostiole: opening or pore usually at top of the peridium through which
the spores escape (see stoma), Figs. 7 & 8.

papyraceous: paper- or parchment-like, used to describe the endoperidium of Bov;sta, Fig. 9.

paracapillitium: sterile, colorless, thin-walled, septate cells similar in
function to the capillitium but derived from thin-walled or occasionally thick-walled generative hyphae, PI. VIII C; see capillitium.
pedicel (of spores): a narrow, delicate extension of the basidium on
which the sterigma and spores are formed. Some species retain
the pedicel and are described as pedicellate, e. g. in Bov;sta,
Lycoperdon, and Geastrum, PI. II D & E.
pedicel (of sporocarps): a short stalk which elevates the endoperidium
in Geastrum, Figs. 18 & 19.
pedicellate: having a pedicel.
percurrent columella: a sterile column of tissue which extends from the
base of the gleba through it to the peridium at its apex, as in
Podax;s, Fig. 65.
perforate at maturity: refers here to a thin peridium which splits or
develops holes and thereby exposes the mature gleba, e. g. in
Phallogaster, Fig. 48.
peridioles: small, oval to rounded bodies ("eggs") composed of a thick
wall; or tunica, containing the glebal elements, e. g. in Nidulariales and Sclerodermatales, Figs. 37, 40, 41.
peridium: outer wall surrounding the fruiting body; may consist of one
or several layers, Figs. 5, 16 A.
perisporium: outer spore wall of basidial origin, PI. lIB.
peristome: clearly delimited edge or area around the opening of the
stoma in some Gasteromycetes; see Geastrum, Figs. 20, 21.
persistent attachment: occurs in fruiting bodies which remain attached
to the substrate after they are mature, e. g. Bov;stella.
pileate development: development resulting in a fruiting body with a
single stalk and pileus, with hymenial tissue often in the form of
gill-like tramal plates, Fig. 1 E.

130

pileipellis: the outer layer of the pileus, usually composed of anticlinal

or decumbent hyphae.
pits or pitted: capillitial elements with round holes (PI. VII C) or the
surface depressions of the peridium, such as the endoperidium of
Lycoperdon perlatum, Fig. 5 A.
plectobasidium: a basidium which arises irregularly from mycelial
tissue, PI. IV A.
polymorphic: variable in either shape or size or both.
pore: an opening in the top of the peridium through which spores
escape (see ostiole, stoma), Figs. 13, 16 B.
pseudocolumella: a stalk composed of sterile tissue, often chambered,
which is surrounded by gleba, e. g. in Geastrum and Radiigera,
Fig. 16. See also Lycoperdon, Fig. 5.
pseudohymenium: a fertile layer composed of plectobasidia; a hymenium which has not evolved for the forcible discharge of the
basidiospores and has no free air space to facilitate the forcible
type of dissemination.
pseudoparenchyma: tissue of hyphae which have lost their individuality and superficially resemble parenchyma tissue of plants.
pseudostem: a well-developed subgleba which forms a more or less
stem-like base supporting the gleba. The tissue is spongy, not
oriented in a parallel fashion, and is often modified glebal tissue,
as in Lycoperdon, Fig. 5.
puffball: name given to species of the Lycoperdales which discharge
clouds of spores when the outer peridium (usually endoperidium
at maturity) is compressed rapidly.
purse: the upper part of the funiculus which contains the coiled
funicular cord in some species of the Nidulariales, Figs. 40 & 41.
pyramid-shaped wart: a wart with a square base and three triangular
sides, pointed or nearly so, often with zone lines, e. g. in Ca/vatia,
Fig. 4 E.
quadrate arm: a receptaculum or arm in the Phallales which is 4-sided
in cross section, Fig. 59.
radiating plates: veins of sterile tissue which proliferate throughout the
gleba and radiate from a point of attachment or from a columella,
e. g. in Kobayasia, Fig. 50 and Truncoco/umella, Fig. 67.
rays: recurved portions of the mesoperidium and exoperidium, often
in triangular segments, remaining attached to the sporocarp and
supporting the elevated endoperidium, e. g. in Geastrum and
Astraeus, Figs. 17, 18, 36.

131

receptacle (receptaculum): as in the Phallales, the stalk or stipe and

pileus or the clathrate structure which supports the reproductive
unit, e. g. in C/athrus, Fig. 62.
recu rved rays: rays in which the outer peridium is convex in profile,
Figs. 18 & 19.
reticulate: having a net-like pattern, PI. IV D.
revolute rays: rays in which the outer peridium is recurved and also
inrolled at the edge, Figs. 20 & 21.
rhizomorph: a cord of mycelium which penetrates the substrate, often
white or yellow to pink or purple, single (Fig. 30) or branched;
common in Gasteromycetes, Lycoperdon, Fig. 8.
rimose: having coarse, often deep or wide splits and cracks, as e. g. in
. the peridium of Astraeus, Fig. 36, right.
rugose: rather uneven or wrinkled.
saccate rays: rays in which the outer peridial walls recurve and split to
form a shallow bowl surrounding the endoperidium, Figs. 20 &
21.
sand-case: the outer peridium of some puffballs which consists of
loosely interwoven hyphae intermixed with gravel or sand particles, as in Disciseda, Fig. 13, and Gastrosporium, Fig. 12.
Schaeffer reaction: the bright orange-red reaction at the point where
the flesh is streaked with aniline and crossed with a streak of nitric
acid; occurs in Longu/a and Section Arvensis in the Agaricales.
sclerotium: a hard knot of fungal tissue often with a hard dark crust and
a lighter interior.
secotioid: refers to a fruiting body having the appearance of an
unopened agaric or bolete, usually with a stipe-columella and a
gleba often composed of contorted lamellulae; strongly correlated with orthotropic spore development, Figs. 65 & 67, but not
necessarily with obvious affinities to known agarics.
sessile: refers to a fruiting body which is appressed to the surface of the
substrate or emerging from a volva without a stalk (stipe), Figs. 61
& 62.
setae: stout, pointed, usually thick-walled hairs, distinctly seen around
the edge of the cup in Cyathus, Fig. 40.
setose exoperidium: an exoperidium in which some cells are transformed into setae; typical of some species of Bovista and Lycoperdon, PI. V A & B.
setose: having setae.
.
sheath: also called basal piece, the structure to which the middle piece
is attached in some species of Nidulariales, Figs. 40 & 41.

132

sinuous (sinuate): wavy, undulating or with indentations or furrows.

skeletal hyphae: hyphae which are thick-walled, infrequently
branched, and often have pigmented cells, PI. V E, PI. VIII A. C.
sloughing exoperidium: an exoperidium which becomes detached and
falls away, e. g. in Lycoperdon marginatum, Fig. 5,6.
sphaerocysts: variously shaped inflated cells characteristic of the exoperidium of Bovista and Lycoperdon, PI. V C, D.
sphaeropedunculate cells: cystidia with a nearly globose apex and a
narrow stalk at the base.
splash cup: an open cup-like structure, often with flaring side-walls, in
which raindrops land and forcibly discharge the peridioles; typically found in the Nidulariales, Fig. 40.
spore collar: a distinctive cylinder of tissue, derived from the upper
part of the sterigma, which remains on many detached orthotropic spores; typically seen in Hysterangium, PI. IV F.
sporocarp: a fruiting body in or on which spores are formed.
squamules: small shingle-like scales often present on the peridium, e.g.
in Endoptychum, Fig. 73.
stalk: sti pe.
statismosporic: having spores which cannot be forcibly discharged
from the basidium. Gasteromycetes, by definition, are statismosponc.
stipe or stem: a more or less parallel tissue system which supports the
gleba but has no direct structural connection with it, e. g. in
Tu/ostoma, Fig. 22. See also pseudostem.
stipe-columella: a regular stipe or stalk which supports the receptaculum and, in addition, penetr<ltes the gleba, forming a percurrent columella, e. g. in Podaxis, Fig. 65.
stoma: a mouth or ostiole through which spores are discharged, e. g. in
Ca/ostoma, Fig. 34.
striate: with lines or striae, which are often parallel or nearly so, of
different density or color from the surrounding tissue, sometimes
used to mean having minute furrows; e. g. in Nidulariales, Fig. 40
and Lycoperdales, Fig. 19.
strigose: having coarse, long, rather st iff hairs, more or less appressed,
e. g. in Lycoperdon, Fig. 7.
stroma: a fleshy mass of fungal tissue upon which the fruiting bodies
develop, e. g. in Broomeia, Fig. ~9.
subgleba: sterile, filamentous or chanbered tissue which supports the
gleba, e. g. in Lycoperdon and C:a/vatia, Fig. 5 (see pseudostem).
sulcate: grooved or delicately groo'/ed, frequently referring to the
tissue surrounding the stoma, e. g. in Geastrum, Fig. 19.

133

taiga: a northern biome including dwarf birch, willow, and conifers;

forms a transition zone between the arctic biome to the north and
the conifer biome to the south.
tomentum: covering of dense, fine hairs over the surface of a fungus.
trabeculae: stout columns which extend from the peridium to the
central core of the fruiting body, e. g. in Mesophelliaceae, Fig.
15.
tramal autolysis: process in which the tramal plates collapse at maturity,
forming a viscous or slimy mass of tissue and spores; typical of
species in the Phallales.
tramal plates: layers of tissue partially or totally devoted to the
production of basidia, Figs. 66 & 71.
triquetrous arm: a receptaculum or arm in the Phallales which is
3-cornered in cross section, Fig. 55.
truncate spore: an elliptical or oval spore which is flat on one end as
though it were cut off.
truncate wart: a wart of any shape with the top flat as though it were cut
off, e. g. in Calvatia and Lycoperdon, Fig. 4 D & F.
truncate, pyramid-shaped: a pyramid-shaped wart with the top cut off,
e. g. in Calvatia, Fig. 4 D.
tunica: the peridiole wall in the Nidulariales, Fig. 41.

vaulted ray: a ray which arches over the exoperidium forming a

chamber, usually conic in profile, Fig. 17.
volva: the remains of the peridium which form a persistent cup
surrounding the base of the fruiting body, e. g. in Phallales, Figs.
43 & 44 and Chlamydopus, Fig. 24.

wart: a small cone-shaped ornamentation on the spores, PI. " A, or on

the e?<operidium of Lycoperdon, Fig. 4 D, E, F.
wrinkled peristome: a peristome with uneven tissue, which is neither
sulcate nor ridged and grooved, Fig. 20.

134

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144

INDEX
Abies 114
Abstoma 33, 34
Acacia 64
acolumellate 123
acuminate 47, 123
acyanophilic 11, 17, 123
Agaricales 108, 122
agaricoid 123
agaricoid gasteromycetes 30, 102, 108
agaricoides, Endoptychum 108, 120
Agaricus-like 106, 109

agrifolia, Quercus 114

Agrocybe 27, 106, 108
Agrogaster 106, 108
albipes, Brauniellula 114, 121
alcohol 25
Alpova 98, 99

Amanita 100
amyloid 26, 27
angiocarpic development 23
aniline blue 11
Anthurus 23, 79, 80, 94
apical germ pore 7, 8
apophysis 46, 123
Arachniaceae 32, 37

Arachnion 38
archeri, Anthurus 94
arenaria, Montagnea 110, 116
Artemesia 50
Aseroe 23, 79, 80, 91
asperate 123
asterospora, Nigropogon 112
asterospbra, Richoniella 112
Astraceae 62, 63
Astraeus 9, 12, 16, 19, 22, 26, 61, 63, 66
Atriplex 50, 51
attenuatus, Dictyocephalos 51, 58
aulaeate development 21, 13, 123
aulaeothecium 23, 123
Austrogaster 104, 113
Austrogautieria 96

145

ballistosporic 10, 123

basidia 8, 9
basidiospores 8
Battarraeoides 12, 20, 50, 57
Battarrea 12, 15, 19, 50, 57
binding hyphae 11, 123
bird's nest 69

Blumenavia 80
Boletales 113, 122

borealis, Lysurus 90
borealis, Protuberella 84
Bovista 6, 17, 19, 23, 24, 26, 33, 34
Bovistella 23, 33, 34
bovistelloid 23, 24, 123
bovistoid 24, 123

Brauniella 122
Brauniellu/a 104, 114, 121
bri II iant cresyl bl ue 28
Broomeia 64, 68
Broomeiaceae 62, 64
brown rots 28

bryantii, Geastrum 46
button 59, 123
byssoid 123
caducous 31, 124
Calbovista x, 34, 35
Ca/ostoma 6, 7, 20, 48, 53, 59, 60
Calostomataceae 48, 53

Calvarula 82
Calvatia x, 6, 14, 15, 23, 26, 31, 33, 34
calyptra 77, 87, 123

candida, Disciseda 44
cannon-ball 69
capillitium 14, 16, 17, 26, 48, 123

Castoreum 38
centrifugal 23, 124
centripetal 22, 124
chambered subgleba 40
Chamonixia 103, 114

chilensis, Rhodogaster 112

Chlamydopus 12, 17, 19, 50, 51, 55

146

cinctus, Staheliomyces 78, 87

cinnabarina, Calostoma 59

circumscissile dehiscence 20, 48, 50, 57, 124

citrin a, Truncocolumella 107, 117
clamp connection 12, 124
Clathraceae 23, 75, 77, 78
clathrate 75, 124
Clathrus 23, 79, 80, 95
Claustula 75, 85
Claustulaceae 75, 77, 85
coliforme, Myriostoma 63, 68
collar 46, 124,
collar, spore 7, 8, 124
columella 45, 124
column 63, 68, 124
columnata, Linderia 94
Colus 81, 92
comatus, Coprinus 102, 107
coneae, Agrogaster 108

congo red 25
congregata, Broomeia 64, 68
connivent fibrils 14, 18, 39, 124
contorta, Pinus 115
corallocephala, Kalchbrennera 91

coriaceous 18
coronate 74, 124
cortex 124
Cortinarius-like 104

cortinate 117, 124

cothurnate 112, 124
cotton blue 11, 17, 26
cresyl blue 28, 112
cretaceum, Castoreum 44
Cribbea 122

Cribbiaceae 122
Crucibulum 70, 71
cucullata, Weraroa 113, 118
cyanophilic 11, 26, 103, 124
Cyathus 10, 70, 71, 73

147

dacrydiodes, Podocarpus 108

deciduous warts 40, 124
decomposers 1, 31, 75, 77, 78, 99, 107, 108, 109, 111
dehiscence 42, 125
defilei, Gyrophragmium 110, 119
delimited peristome 64, 68, 125
deliquescence 107, 125
dendritic columella 96, 117, 125
desert biome 31, 125
desertorum, Ga/eropsis 109, 118
Destuntzia 103
dextrinoid 26, 27
diaphragm 36, 43, 125

Dictyocepha/os 50, 51, 58

Dictyophora 23, 77, 78, 86
digueti, Battarraeoides 50, 57
dimitic system 11, 125
Diplocystis 64
disc 36, 125
Disciseda 19, 33, 35
double connivent fibrils 39
duplicata, Dictyophora 86
earth ball 61, 125
earthstar 31, 32
echinulate 6, 125
ectomycorrhizae 28, 61, 75, 98, 101, 113
ectosporium 3
elaters 12, 15, 48, 125
emplacement 125
endoperidium 18, 125
Endoptychum 7, 20, 106, 108, 120
endosporium 3, 125
epigeous 2, 122, 125
epiphragm 73, 125
episporium 3, 125
ethanol 25
evenescent 96, 125
Excluded Species 122
exoperidial fibrils 39
exoperidium 18, 125
exosporium 3, 6, 125

148

Fagaceae 61, 114

false earthstars 61
farinose 125
fetid odor 75
fibrillose truncate-wart 39, 125
fibrils 18, 39, 126
fimbriate 46, 68, 126
fischeri, C/austu/a 85

forate development 21, 22, 126

forcible spore discharge 4
fornicate ray 45, 126
fornicatum, Geastrum 45

funicular cord 73, 126

funiculus 73, 126
fuscog/eba, Radiigera 45
fusiformis, Pseudoco/us 92
galericulata, /tajahya 87
Galeropsis 105, 108, 118
Gasteroagaricoides 106 109
Gastrobo/etus 104, 115
Gastrocybe 105, 109

gastroid 1, 126
Gastrosporiaceae 32, 37
Gastrosporium 7, 23, 37
Gautieria 96, 97
Gautieriaceae 96
Gautieriales 22, 30, 96
Geastraceae 32, 36
Geastrum 19, 31, 37
gelatinous peridium 86, 126
gelatinous tissue 86, 126
Gelopellidaceae 75, 76, 85
Ge/opellis 75, 85
generative hyphae 11, 126
germ pore 7, 126
gigantea, Ca/vatia 35
gigantea, Langermannia 35

gleba 86, 126

glebal chambers 107, 126 .
glebifer 81, 126
globaroid 23, 24

149

Gomphidiaceae 104
Gomphogaster 104, 115
gracilis, lIeodictyon 95
graveo/enes, Gautieria 97

gum guaiac 27
gussets 5, 110, 116, 120, 126
Gymnopaxillus 122
Gyrophragmium 106 109, 110, 119

hanging collar 46, 127

hapteron 73, 127
heteromerous 30, 127
heterotropic spore 9, 127
hilar appendage 7, 127
hilar scar 127
hirundinosus, Co/us 92

Holobasidiomycetidae 9
holobasidium 9
homogen 22
homogeneus development 21, 22, 127
hygrometricus, Astraeus 66
hygroscopic 62, 127
hymeniform 109, 127
Hymenogaster 8, 103, 122
Hymenogastrales 23, 30 107
hypha (ae) 17, 127
hyphal systems 11
Hypogaea 122
Hypogaster 122

hypogeous 2, 122, 127

Hysterangiaceae 75, 76, 84
Hysterangium 8, 84
lIeodictyon 79, 81, 95

indehiscence 127
indurate trama 110, 116, 127
indusium 77, 86, 127
inquinans, Phellorinia 51, 56
insect-disseminated 4, 9, 75
insignis, Ca/ostoma 60

invagination 21, 127

inverse koralloid 23

150

invertebrate herbivory 4
involute ray 66, 127
/tajahya 77, 78, 87
jamaciensis, Protuber 89
Kalchbrennera 79, 81, 91
kammerartig 22
KOH 27
Kobayasia 83, 89
koralloid 22

labyrinthiform 96, 128

laccase enzyme system 27, 28
lacerate ostiole 68
lacerate peristome 36, 63, 128
laceratum, Schizostoma 52, 55
laciniata, Battarrea 57
lactic acid/phenol 11, 26
lacunar development 21, 22, 128
lacunose 7, 96, 103, 128
lamellate fungi 99, 128
Langermannia x, 33, 35
lantern 79, 81, 93, 128
lateritia, Gastrocybe 109
Laternea 79, 81, 93
Lentinellus 27
Leptospermum scoparium 85
Leucogaster 101
Leucogastraceae 101
Leucogastrales 29, 101
Leucophleps 101
leucosarx, Gomphogaster 115
Ligiella 80, 81
Linderia 23, 80, 81, 94
lipid body 7
loculate 102, 128
longitudinally grooved 55
Longula 20, 28, 106, 110, 120
lumen 11, 17, 128
lutescens, Ca/ostoma 59
,

151
Lycoperdaceae 32, 33
Lycoperdales 22, 23, 28, 30, 31, 48
Iycoperdoid 23, 24, 128
Lycoperdon 14, 16, 19, 22, 23, 26, 33, 35
Lysurus 79, 81, 82, 90
macrosporium, Neosecotium 111
mammaeforme, Lycoperdon 41
maracuja, Protubera 84
marginatum, Lycoperdon 41
Me/anogaster 7, 97, 98, 99

Melanogastraceae 98
Melanogastrales 29,
Melzer's reagent 26, 114, 115
mesoperidium 18, 47, 128
Mesophellia 38, 44
Mesophelliaceae 32, 38
metachromatic 28, 112, 128
metachromatic spores 28
meyenianus, Chlamydopus 51, 55
middle piece 70, 71, 73, 128
mirabilis, Que/etia 52, 56

9""

mokusin, Lysurus 76

monomitic 11
monopileat 23
Montagnea 7, 20, 105, 110, 116
Morganella 33, 36
mucronate 113, 118, 128
multipileate development 21, 23, 128
Mutinus 9, 23, 76, 77, 78, 88
mycelium 27, 52, 53, 54, 55, 128
Mycenastrum x, 14, 16, 34, 36
myco 128
Mycocalia 13, 70, 71
mycorrhizal 62, 63, 84, 96, 100, 107, 115, 128
mycosderids 36, 128
Myriostoma 63, 68
Myrtaceae 61

152

naked stoma 128

Neosecotium 105, 111
Nia 99
Nidu/a 70, 71
Nidu/aria 15, 70, 71
Nidulariaceae 69, 70
Nidulariales 29, 69
Nidu/ariopsis 70, 72
Nigropogon 8, 122
nipponica, Kobayasia 83, 89
nitric acid 28
Nivatogastrium 20, 23, 104, 111, 119
non-persistent attachment 19, 33, 128
Nothofagus 104, 114
nubigenum, Nivatogastrium 111, 119
nurse hyphae 9
obovate 85, 129
Octavianina 103
odor 4, 75, 129
one-spored basidium 129
open-pored hilum 7
oranges 83
orthotropic spore 6, 7, 8, 9, 129
ostiole 20, 40, 41, 45, 129
Panaeolopsis 105, 111
papyraceous 18, 129
paracapillitium 11, 12, 17, 48, 129
parasites 64
passion fruit 83
Paxillogaster 122
Paxillus 114
pectinatum, Geastrum 46
pedicel (of spores) 3, 6, 7, 129
pedicel (of sporocarps) 36, 45, 46, 129
pedicellate 3, 129
percurrent columella 107, 129
perforate at maturity 88, 129
peridioles 9, 61, 62, 63, 67, 73, 129
peridium 14, 18, 129
perisporium 3, 6, 129

153
peristome 36, 45, 46, 47, 68, 129

per/atum, Lycoperdon 40
persistent attachment 129
Phallaceae 23, 75, 77
Phallales 23, 29, 75
Phallogaster 85, 88
Phallus 9, 78, 86
Phellorinia 49, 51, 56

Picea 114
pi/a, Bovista 42
pileate development 21, 23, 129
pileipellis 109, 130
Pinaceae 6 1 , 1 0 4 .
pingue, Thaxterogaster 113, 117

Pinus 115
Pisolithus 19, 22, 61, 62, 67
pistil/aris, Podaxis 107, 116
pitted spores 7
pits 7, 40, 130
plectobasidium 8, 9, 130
Podaxaceae 107
Podaxales 102, 107
Podaxis 7, 8, 10, 20, 23, 49, 102, 107, 116
Podocarpus 106, 108
polycystoderm 14
polymorphic 130
polypileat 23, 130

po/yrhizon, Scleroderma 65
pore 12, 16, 130
pored fungi 1, 115, 130
potassium hydroxide 25, 27

pretense, Vascellum 43
proteinaceous core 44

Protogautieria 122
Protophallaceae 75, 76, 82
Protubera 83, 84, 89
Protuberella 83, 84
pseudocolumella 35, 36, 37, 40, 45, 130
Pseudoco/us 76, 79, 82, 92
pseudohymenium 130
pseudoparenchyma 12, 130
pseudostem 99, 130
puffball 31, 48, 130

154

pulchella, Torrendia 100

pulcherrimum, Lycoperdon 41
purse 73, 130
Purshia 50
pusilla, Laternea 93
pyramid-shaped wart 18, 39, 130
quadrate arm 82, 130
queinzii,Secotium 112
Que/etia 49, 52, 56
Quercus 112
radiating plates 130
radicata, Bovistella 42
Radiigera 19, 31, 37
ralstoniae, Gasteroagaricoides 109
ravenelii, Mutinus 88
rays 36, 45, 46, 47, 62, 66, 68, 130
receptacle 75, 131
receptaculum 79, 93, 131
recurved rays 46, 131
reticulate 8, 131
revolute ray 47, 131
rhizomorph 19, 41, 43, 58, 131
Rhizopogon 103, 122
Rhizopogonaceae 102, 107, 122
Rhodogaster 102, 112
Rhopalogaster 83, 84
Richoniella 102, 112, 122
rimose 62, 63, 130
rimose stalk 58
rodent herbivory 4
rodrigueziana, Ligiella 81
ruber, Clathrus 95
rubra, Aseroe 76, 91
rugose 131
Russulales 26, 30, 102, 120

155

saccate ray 47, 131

saccatum, Geastrum 47
saccatus, Phallogaster 85, 88
sand-case 35, 131
saprophytes 84
savanna 31, 131
scanning electron microscopy (SEM) 1
Schaeffer reaction 28, 110, 131
Schizostoma 19, 49, 52, 55
Scleroderma 8, 19, 61, 63, 65
Sclerodermataceae 62
Sclerodermatales 30, 61, 62
Sclerogaster 103, 122 . .
sclerotia 64, 131
Secotiaceae 23
secotioid 10, 102, 131
Secotium 28, 106, 112
sessile 81, 95, 131
seta (-ae) 14, 73, 131
Setchelliogaster 122
setose 14, 131
setose exoperidium 131
sheath 73, 131
Simblum 79, 82, 90
simplex, Gastrosporium 43
simu/ans, Tu/ostoma 54
Singeromyces 122
sinuous 132
skeletal hyphae 11, 132
slits 12, 15
sloughing exoperidium 40, 41, 132
Smithiogaster 122
Sphaerobolaceae 69, 72
Sphaerobo/us 69, 72, 74
sphaerocepha/um, Simblum 90
sphaerocysts 14, 132
sphaeropedunculate cells 108, 132
spicatus, Podocarpus 108
spiny capillitium 14
splash cup 4, 40, 132
spore collar 3, 8, 132
spores 3

156
sporocarp 132
squamules 132
Stahe/iomyces 77, 78, 87
stalk 132
stalked puffball 48, 132
statismosporic 9, 132
stellatus, Sphaerobo/us 74
stem 48, 132
stercoreus, Cyathus 69
stinkhorn 75, 132
stipe 48, 132,
stipe-columella 108, 116, 117, 118, 132
stoma 19, 132
striate (striae) 46, 55, 73, 132
striatus, Cyathus 73
strigose 41, 132
stroma 64, 68, 132
subapical germ pore 7
subgleba 34, 35, 36, 132
sulcate 46, 132
swollen septa 17
syringaldazine 27
taiga biome 75, 133
tan bark 52
tapering fibrils 39
texensis, Longu/a 110, 120
textura angularis 12
textura epidermoidea 12
textura globosa 12
textura intricata 12
textura obi ita 12
textura prismatica 13, 14
Thaxterogaster 104, 113, 117
tinctorius, Pisolithus 62, 67
tomentum 71, 133
Torrendia 98, 99, 100
Torrendiaceae 98, 99
trabeculae 44, 93, 133
tramal autolysis 133
tramal plates 119, 133
transmission electron microscopy (TEM) 1

157

transversarium, Rhopa/ogaster 84
triplex, Geastrum 47
triquetrous arm 82, 133
true capillitium 11
truncate pyramidal warts 39
truncate spore 133
truncate warts 39, 133
Truncoco/umella 103, 107, 117
Tu/ostoma 17, 19, 22, 49, 52, 54
Tulostomaceae 48, 49
Tulostomatales 30, 48
tunica 73, 133
turbonatus, Gastrobo/etus 115
variegatus, Me/anogster 97
vascelloid 23, 24, 133
Vascellum 17, 23, 33, 36
vaulted ray 45, 133
verrucosum, Tu/ostoma 53
vibrissa, Nia 99
volva 55, 58, 86, 87, 119, 133
warts 6, 7, 8, 15, 133
Weraroa 105, 113, 118
white rots 28
wind-disseminated 4
wrightii, Diplocystis 64
wrinkled peristome 47, 133
xeric habitats 5, 109, 110, 111, 133

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091

Gasteromycetes: MorphologIcal &

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