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Livro Miller Amp Miller 1988 Gasteromycetes Morphological and Development Features With Keys To The Orders Families and Genera PDF Free
Livro Miller Amp Miller 1988 Gasteromycetes Morphological and Development Features With Keys To The Orders Families and Genera PDF Free
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l
I
GASTEROMVCETES
Morphological and Development
Features
with
Keys to the Orders, Families, and Genera
by
Orson K. Miller, Ir.
and
Hope H. Miller
ISBN 0-916422-74-7 I
1
TABLE OF CONTENTS
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
General Morphology
. ..
Spores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
. .
Basidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 .
Hyphal Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Peridium ...... · . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Development · . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Chemical Reagents ·.. .......... . .... . .. . ..... . ............. . .. 25
Key to the Orders of Gasteromycetes . . . . . . . . . . . . . . . . . . . . . . . . . .
~ 29
lycoperdales 31
• • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • •
Tulostomatales ............................................. . 48
Sclerodermatales • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 61
N idu lariales • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 69
Phallales • • . . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
~ ~ ~
Gautieriales • •• • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 96
Melanogastrales • •••• • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 98
.. .
leucogastrales . . . • • . . . . . , • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 101
~
•
•
"'---------- -------------
•
>
LIST OF PLATES
LIST OF FIGURES
•
$
..
VII
ap apophysis pr peridiole
b basidia ps peristome
ca calyptra pc pseudocolumella
•
Clr circumscissile dehiscence pu purse
co collar r ray
col columella re receptacle
dia diaphragm rh rhizomorph
•
en endoperidium n ridges
•
epJ epiphragm s stoma
f fimbriate sc sand case
fun funicular cord se secotioid
g gleba sh sheath
gl glebifer sp spores
gu gussets • sr saccate ray
h hollow center st stalk (stipe)
ha hapteron stc sti pe-col umella
•
J indusium str striate (striae)
la lantern sub subgleba
10 lacerate ostiole sui sulcate
me mesoperidium tr trabeculae
mp middle piece tu tunica
0 ostiole v volva
p peridium (exoperidium) ve veil
pe pedicel
• ••
VIII
PREFACE
There are many macrofungi which are classified under the ambi-
guous 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 informa-
tion 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 investiga-
tion.
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 excel-
lent 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, BARC-
West, 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 transla-
tion of many passages in the "Gasteromycetes" by Dr. Pilat from
Czechoslovakian to German by the late Dr. Ruzina Hilber. We acknowl-
edge the cooperation and support of the Biology Department at Virgin-
ia Polytechnic Institute and State University and the University of Mon-
tana 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.
•
x
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 puff-
balls), Sclerodermatales (earth balls), Phallales (stinkhorns), and Nidu-
lariales (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 Gastero-
mycetes".
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 dis-
charge 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 transmis-
sion 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 dis-
charge 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
2
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 rec-
ognizable 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.
•
p
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 charac-
terized 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 hetero-
trophic 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 basi-
dium 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 mucilagi-
nous 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 de-
scribed above. The Nidulariales have short elliptical, ovoid, to globose
4
•
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 basidio-
spore of Calostoma with echinulations.
7
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.
8
c.
D.
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 stink-
horn. 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 sterigma-
tic 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 Gastero-
mycetes. Moreover, this cbaracteristic is usually accompanied by dra-
matic alterations in the morphology of the fruiting body to accom-
modate 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
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 differen-
tiation peculiar to or repeatedly encountered within the Gastero-
mycetes.
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 Gastero-
mycetes 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 gen-
erative 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 deve-
lopment, 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
•
14
~
• ~ ,
• I .~
•
\
D.
, '", '
,
D.
17
t.
c.
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 develop-
ing 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 dis-
ch.uged. Spore discharge usually occurs when droplets of rain or dew
strike the surface of the endoperidium, causing small clouds of spores
,
19
A. B. .- h
h
. . ' . .. . . •. . . ..
"
. '.
.' ".
• •• • I . ' ••
c. D.
h
E. F. ~h
+-h
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 dissemina-
tion. 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 de-
velopment. Pilch (1958) was the only one who recognized the homo-
geneous type. Each of these developmental types is described below
and illustrated in Figure 1.
4. Aulaeate (Invers koralloid) (Fig. 1 0): This type occurs when plecto-
basidia differentiate on the surface of down-growing primordial tissue
I\ which usually grows from the upper surface of the peridium. As matura-
I tion develops, small areas of tissue may break away to form locules.
Hymenial development could be characterized as centrifugal, since the
!II hymenium forms on the inner surfaces of the primordial tissue. Gleba
developing in this manner is the aulaeothecium of Kreisel (1969), and it
q
I. 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,
I as in an embryonic agaric button, with gill-like tramal plates (Podaxales,
r Podaxis, Fig. 65; Secotiaceae, Nivatogastrium, Fig. 71), or with the gleba
A. B.
c.
A•
•
,
\
)
25
CHEMICAL REAGENTS
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 en-
hanced 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.
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.
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 Homobasidio-
mycetes 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
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
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
LYCOPERDALES Clements
1. Fruiting body small (0.5-2 cm broad), ovoid; gleba with minute perid-
ioles which resemble sand grains .............. 4. Arachniaceae
1. Fruiting body larger; gleba without peridioles, powdery at maturity
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2
1. Lycoperdaceae Smarda
Fruiting body epigeous or becoming so at maturity, ovoid, elliptical,
pyriform, 0.5-140 cm broad, exoperidium sloughing to reveal a well-
formed 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
Abstoma Cunn.
Fruiting body up to 4 cm broad, depressed-ovoid to ovoid, exoperid-
ium 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.
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.
2. Geastraceae Corda
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.
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)
A.
B.
'.. '
c.
D.
E.
F.
~-B
......-:
o
en
....--- g
pc
sub
. .•.. . ... . .
• • ... ...
. . . . .,
. •
.- • •
.' " .
,....-- 0
. -. ....
•
•
•
.....~~.",
.' .::"':~,.,
' .... ,.. .. '::
..... : . .
.. . . .
'.
. '...
..
.
.' ' ...
• •
--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
p
Fig. 8. Lycoperdon mam-
maeforme with the
unique cottony patches it:? P
formed as the exoperi-
dlum (p) breaks up, re-
vealing the ostiole (0)
and endoperidium (en).
Rhizomorphs (rh) prolif-
erate from the steri Ie
base. ')---rh
42
en
g
.. ,
. .'
. .. . ..'. . .
. . ." .
. " ..
..
p
. .. ..
!. :
en - ---'-0-
>-----rh
.: . , .. . . en 10
. . .
'.
. '. .:, . .
. .. .. .'. '
"
. .'
.'
~.~.,
.
•
' .....
.• ~-:':.,;.to.,\ ..:. / .... . . ~-,,'
. .
• • " ••• ')' •.•• ". • . • • ',
.. . . . . . . ,~..j,:....'
:::.;.~~"
',-'~'
:',-,- ..
., I '.,
.. ' ";' ,
.... .' "•.•,. •
.\.~
I .
;
'" "~.::"'.
"~",
. ........ .
.• :.~: "
. ~. .~'
'J t
. .(".
,
".
. / ."
....:,..
'" "
........ •: .. 1. "
.. ,.'.:........ :;,' . ." ,
• or, " '.", ',_' ..\.. ••
"., : ..... ':; :: -'." . ~'.:
...' .' . ." ,
·,., · .
. ;.A
". ~..,
., .• ,' ......
. . .
.
·
,-
... . ,'·
' ..
' .
.-....
'
·
....
. .• .., .
.. :.!:
.'
~
. . p
"\.'
...... ,.. . ' ,'",.", " , ' ' ''' . '
.
...
..
. .,. .....'
. ••
:'.',
~.. . '.
. ..', ;. . , . ...........
,. ' ' . ,
.,r ..
...•• . . .
. .
•• '
.'
• A',
~.,
..........
.." ':'"
'..
..... ,';!'..... ."
,.:...~
. ...'·r'·,
:.:.... ,::
... . ." .
. '.,;, ... "'" " .'
..
' " ' ".:"
• '. '. ... . . ,.' . '. '. ',;.,.)/1
. '
',; '.'- .'.:." :.:.....": .. :.'
:;.~/,~"'.;"
. .•i{~ ,; . .;,.:.:..' ,:'. ,.:!-.: " ::. :.~:.' ,
, .
"", ,..... ,.;,
'
• :~.
"r
'J
..... , . : .. .'..,....
. ..•...
.;'
.. ,:,.,
"
.... .. ..":
' .
... : . .. i."
,;"
.i·
• ' . , \........~:.; .. , . .
.'
.
::.....
••
'.:>...... ......., .... ......... .....
.
''',~
'
•• ~ ....' ••• "lo: ....... ,> . . . . . . .
~:.~·.~·.>!.,:-:·:~;.\:· t~::i:.:::;"':::::·' " .
' .. '..
'" "'.' . ~
'
\ .
•
. .. ...
, : '
..' .. '..
I · · ·..... · · ' .... " , · ·
. ~~.:~~.:}( ~ . ... .
.• .
'
.., ..
'. .'
. ". "
"
A, '.'
,. '. "
.
' .
,. .
' , ; r ... ,; ..
..•..'. -. .'..... . .. .
, ,
. .
':'",
. .' . : :.
'
. ~
.
"',
"
.
.
" . ..,
... ". ", ,.. -, , ...... : ....,,:;.
".~.. ,'~'
,-".,,,,,,~;,.
.' . . : . '... . .
• . •• ". ..Iot"'' ' \"'". • ~
.
.
.. . . (ilP:' ... " •
. " ~ -:~,... ~:~~,~,":' .'.:' . .
....... ,.. -..... :,
. "
.
,
.~.,:.~
":'1...
,~
.'
. . ..
'.i!.
.
'"#<'
; .. ,;:
..',' .- ' ......
'. '".;,
~. "
.'~'.
.'.
.::
..
."
..+
" f.l'
.;2'
~:',
. '.
A.",;;.
~;.
. .
' .
..'
. . . .'",",
," ...
.'
.' .." ft-g
. ·,:·,i-;iif.t'
, ,.',
. ...,.;:'
g--l 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.
rh
44
..;;;......",....--- 0
en Fig. 13. Disciseda can-
dida. At maturity the
exoperidium, which is
a sand case (sc), re- .
mains persistent over
the base, and endo-
peridium (en) and osti-
ole (0) at apex are ex-
posed. "
sc
p g
.
.. .
the exoperidium (p) at
maturity to reveal the
gleba (g).
p
tr
g
core
• en
.'. . . '
,
~
.. . ~----p
.' :
:. .
"
:;i~ -"'J. •
./
,
.
.,''/i;~~:;~:~
,".
... ;:~i... :
"
..
~~
".~.,.
.'
;
.,
'.'
",. '.
:• .o'
•
.
•
," .,",'"
"
"}o,'
' ,'
.
f·\
, ',
... ....,
,', '• .'.:1-'"
.
. •
. ,
•
.:.~
~'f..' ,.
•
,
•
' , g
. '. :. , '
.
,
. ' - - 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
0----
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
~--~ me
sr ----,.<..,...".~
..
•
...•
",
: . ': :
" ;,,'
",
•
r
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
-~......"..-- en ps
--me
sr
.,-o-f--- me
Fig. 21. Geastrum triplex. The saccate rays (sr) become revolute, with
the mesoperidium (me) breaking completely apart, leaving the endo-
peridium (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
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 connec-
tions present. Cosmopolitan, decomposers of plant litter.
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.
.. . ~--en
. . ..
o ~--p
,
.---. .. '------..
·~
: '., ..
"
---::~---- en
ground level
o
\+-----st
\
~,
•
• 1------ rh
f
---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
en
¥----v
...
.' '
o·
(\~ ;., .' - - - _. p (= sc)
Ii
I
1
•
\ .
st
..
. ', :":.'. . ,
. '. """*"--
. .~
. • .. IV.
. ,... . ' -" ...., . p
'. '. :.~:.;
• .
•
.
'. '!.,...
., ' " ' • ....
·,,'
. ~~. .. '.
. ,: • .
• • '~ ~.. \.' ....
',
'~:
'" ,
'. '." ,,... , ; . .
~
J'. ,_
.
. "
h ',' •
. . ..
:# - - - - 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--+
p
g -----''T
Fig. 29
•
Clr--'
\----- st
st - -
v
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
g
en ---1,:,
me.-:::::::.:=i
--st
---st
- - solid
', n center
\
--v
v.7~--v
:---rh
) - - - rh
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) . .
~.~\
. . "
o
/ •. ; , • . '•
'.
.
.
•.
•
, . ..
, .
"
, . _....l'......
\ - - - - en
1'"
~
~/j~~~~~'~~~-
'''',. ,.,'. '.
_• • ps
h!~:"
, . , ,
. . " '
,
,
,
I
A;i~:: .
.'
'. .
' . .,
~' : " ".' ", '
"
~ ~, .
"
•
~
"
:
. ,
I
• , ' .... . ~ ;f
, , , . '", ".~.
,
. . .
\ \.
'.
• •• .
.. ... . ''.
~
, ,
. ....::.,..
;
'\.. ..
"
. ', "
..
. .'.
•••• •
"
I
., ,
•
•
,• ,
\
f st
'lI i,
: .
, •
, ,
.:/'f , I
\
• ,
\
,. .. f
,
"
- •
I I~
•
.... , •' •
. · . ••·..'
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 Scleroder-
ma, irregularly club-shaped to oval in Pisolithus, to stellate or earthstar-
like 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, purple-
brown, 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.
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. Capil-
lilium 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.
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 abun-
dant, hyaline, with clamp connections (PI. VII B). Spores 4-11 JLm, glo-
bose, thick-walled, warted, reddish-brown. Mycorrhizal with shrubs
and trees, throughout the world.
3. Broomeiaceae Zeller
g--H
g .. '
4-- P
. .
.' ..
t-r- P
~---rh
---g
---en
en
10
't--- r
.' 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 endoperid-
ium (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
... ' . , .
- pr
j~.::;:;:.;,. " ., ," .. "
'~"
..:.~.~"
.,
,' ..
'.:.,,' .
" I
.' t, --pr
. ."' . .
, .,. . .
'."; .,
• •
~
,
•~ • f.
.... •
.I . '\
.
',.
,':.' ~
. .
~"
~•
" .
1" ~• .
'.
.
~---sub
st ---t+-
.
.(W·.;:..jr': ·r)':::;·""''-..
... ;
., . . . .. . ,. .,.
.'
,\'
.
• I'
'
.\
,,' s·
'.'." .
;t. .
~ oJ
.
,
. ':
'"
.'
" :
.
•
en -------F.
_ _- f
ps _. ::::::::::::::-
.-<__ 0
t:::;en
-p
..••,. . . .
spore sac has a delimited, fim- • ;: ~."{., • ••
.. .
.. ,",. ........
briate (f) peristome (ps) surround- •
NIDULARIALES
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.
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.
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, disc-
shaped, 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.
-
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
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
----mp
74
pr
...,.., - - - p
. . ... .
, .' ,. . •
· : :.. . . ·.
• .'
•)U'i" .
.""...," ..
•
'. '.
',; :. . .
"
. .•
.
f ·, ..
~
'.· .. - .
~
· " . . ~--- en
I. •
.
·.'., ".:. ", .. .
.' .· . <
·· .' '.. ." .
~
~ . ...
'.,.
. , ". .
""'- • ••
. tI . ' ,
......:..~:. ",",.)'.. : .
~
.'
'.
'
~ ... ~. .:. ' .
..' ",..... . ..
P ---/f-'...,.-- •• ..
.., .
•
. .
... . . .....,',... ,,'.. .. ,
, • ..
.;. .. ...'..
•
~
••
..
• ,
..
•
• •
Fig. 42. Sphaerobo/us stellatus. Lower left shows splitting of the exope-
ridium (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.
2
75
PHALLALES Cunningham
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 rhizo-
morphs 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.
77
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.
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
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.
2. Clathraceae
Fruiting body sessile or arising from a short, chambered stalk, com-
posed of a receptaculum of upraised, spreading, sometimes arched and
united arms or a latticed (clathrate) receptaculum composed of inter-
connected 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 de-
velopment). Peridium three-layered. Basidia 6- to 8-spored. Spores
narrowly elliptical, covered with mucilage. Cosmopolitan, decom-
posers of wood and plant organic matter.
-
79
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 corruga-
tions; 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.
Ligiella Saenz
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.
3. Protophallaceae Zeller
83
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).
4. Hysterangiaceae
Fruiting body hypogeous to epigeous, ovoid to pear-shaped or short-
stalked, 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 rhizo-
morphs 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
5. Claustulaceae
•
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).
D
86
Gelopellis Zeller
(Characters of the family)
.-- p
•
r------ n
---g
'---- v
•
't--I
Fig. 45 ca
'-- g
e;'
' ~, ;'., "i, '. g
.' \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
~--h
fr-- me
~-g
r--en
g
!r---- st
--st
-----rh f---V
Fig. 47. Mutinus ravenelii. The young specimen on left has an exoperid-
ium (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).
.
apex splits and the exoperidium (p)
'.
.. .. " ' · . ",."
.". " ·.' . '.
..
. '. is perforate revealing the gleba (g).
. .. .
g-+ ~---p
..•
•:',
~.
•
:".: .. g
· .'. ..
..
st-- .'
1----- rh
89
· .
•
..
..·. ....... . ....
. •
•
. ... , '
· ...... . .:"'
~
~
.. .'., .
,' - .. ,': .
' . .
p
I~- rh Fig. 49. Protubera jamaicensis.
IT- ·en
The fruiting body has a colu-
I-g mella (col) of white, gelati-
~- col nous tissue which branches
upward between the olive-
' - - - - rh green, glebal (g) chambers. A
flaccid, tough exoperidium (p)
and gelatinous endoperidium
(en) surround the gleba (g),
and stout rhizomorphs (rh)
(only one illustrated) prolifer-
ate into the soil.
•
•
p
. ...
· .'... .
. . .'
..
t , .:..
·• •. •
.;; ;.t'
. . 0'. •
·.
" "
..
.!'
,.
.'. . . ..
'., "r"-- en
. . .
'. .~
'...
rh ~-g
. ....
'. '"
receptaculum (re), which is \"" . "
• - '" ..
.. :
.' r
'
•
~--st(=
\.
I • ' 0'
. '•
'. •
~
'~-I'
·I .
.
. ', • •
"
.
----:':r-- v
. .. .
... . .. .. .
•,
': ': ', ; ..: "
: : ':',.. '.
•• •
...' .
. . . .,:- "
'-----rh
.= ---- st
•
.• .
'. ; ~""
.• . .- 4 -. v
" ,
'. ."--=':~
.....
.".....~
,
.", ..
. • ••• ·z .. ~ . "..... .
.
. ". ...' . '.
• • • v
re 91
g
Fig. 53
-i------st
g--+
: . 'j. ••
st - - - :fj'~ - •
..• .-~
.. ';':.i':':
~
'(' . ,' ~¥_---v .
-
•
. ,'" ...
.. I ..
",
,
'•.:,
• , ••
v-+.-'
} : - - - rh
~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 Fig. 57. Laternea pusil/a. The gleb-
ifer (gl) is suspended below the
t!'::"'!c:r
.
'--- re receptaculum (re) by trabeculae
. •. •
,I'
\ "
•
. (tr), forming the lantern (Ia). The
<.
t '.
receptaculum is sessile within the
,'. ..
••
;~
. .' F-l-,-- V (= p)
.
.. . .
.
~. "
:.
:.:'('
. .
' ,'
,'
----rh
~""<:---- gl
:........l ,-1,,.---- tr
,....--- la
re
. ..
. ..
Fig. 58. Laternea triscapa. The gleb-
.........;.......+----v
ifer (gl) is held in place and clasped . •y
. .', .... .
"·~·i'.·· ~;,..
by the trabeculae (tr), which form
th e lantern (Ia). The receptaculum
(re) is sessile withil1 the volva (v),
fro m which rhizomorphs (rh) prolif-
r----- rh
erate into the soil.
94
,',. ,"
. . -1 . • .
~:::::::,..---- p ,
' ) - - - - - rh
" ~,--- v (= p)
Fig. 59. Linder;a co/umnata. The young .
'"
"" "
- ... ,
,
,
,,
v ----,
P ----Ii ...;;.;..",t--- v
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
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 recepta-
culum (re). A flaccid button is in the foreground.
...
..'.
. .'" .
:.
,"
GAUTIERIALES l
j
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, longitudin-
ally 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 Hemis-
pheres; mycorrhizal.
. "
, ~ "" .., ...,;. >.,~-~-.........
.: .:.',,'!t M·~
41' ". " -~ -~
"lKv ·-# : . "'
! ... ··· ·~; · · l 'r- ..
•
. '.... if,;,,;t<¥'
I~:c 'Jtj;,.
",:.:. -t:l,VfI ~•.:
.:.. :r.-. ".
.,,} !,.J.. ~ . .'t.. . . '..,' ",:
• ,- ~"-J' 'oJ
:
.. ' ;'1.-
.::..
..
'
." .'
..
.....• •
,' ~
\ .•
-:
' ..•..........~. ,..
' l '!'
~ ~ ... ...'
~
-
. ..
',.
._, (l"
,',- -
- - col
--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
1. Melanogastraceae luI.
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
,
•
1
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. Hy-
phae with or without clamp connections.
2. Torrendiaceae Malen~on
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 i
J
initially persistent, but splitting at the apex at maturity leaving a minute
volva surrounding the stalk base. Gleba a series of locules with a hymen-
ium developed in each. Capillitium absent. Spores 13-17 x 5-7 J.Lm, ellip-
tical , 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
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. Ectomycor-
rhizal with trees and shrubs. Known from Europe and North America.
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
1
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 (pholiota-
like); 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
105
17. Fruiting body epigeous, conical, apex conic to pointed and mu-
cronate (Figs. 69-70); gleba contorted-Iamellate to intervenose, yel-
low, 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 contorted-
lamellate 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 deliques-
cent 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 thick-
walled ................................................ 25
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; ter-
restrial 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, thick-
walled; 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 yel-
low, 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 Zea-
land under Podocarpus ............................ Agrogaster
27. Fruiting body Agaricus-like; spores 13-16 x 9-10 ILm, broadly ellipti-
cal, 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 appear-
ance). 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, infre-
quently 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.
HYMENOGASTRALES
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-col-
umella 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 sphaeropeduncu-
late 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).
.
j
109
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 deli-
quescent 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 some-
what larger at base. Spores 10.5-12 (-14) x (6-) 7-8 /J-m, elliptical to broad-
ly elliptical, thick-walled, with a prominent apical germ pore, yellow-
brown. Pileipellis hymeniform. Found in temperate areas as decom-
posers in lawns and grassy areas under wet conditions during the
summer. Monotypic: C. lateritia Watling.
Panaeolopsis Singer
•
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.
Secotium Kunze
Fruiting body epigeous, 3.5-11 cm tall, agaric-like. Pileus 3-6 x 3-4 cm, 1
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
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 illus-
trated 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, cinnamon-
brown, to chocolate-brown usually darkening in age; sometimes stain-
ing 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 longi-
tudinal ridges or sometimes isolated warts, orthotropic with a promi-
nent 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, red-
brown, 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, thick-
walled, 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).
~--p
p
stc -->;
- - stc
p
-----:T gU
p
Fig. 66. Montagnea arenaria.
The fruiting body on left has
the conic exoperidium (p) in;
gu
tact, a well-formed volva (v),
and fine rhizomorphs (rh).
The exoperidium (p) col-
lapses, except in the center
and i nd urate tramal plates
form distinctive gussets (gu) 7
at maturity.
rh - - - ; 1
v
117
. ~-- . g
..•
. . ..'..... .. .•
... ..,
: .. .
..
"
'
',' ~ , .
. :'?\ .. ',.
'
'"
.. .,". .,
. '. .'.; .
•
/
=---- stc
p
· ,
>
it: . '.;",. .
-g
.•j:
..•
-If
•
.: •
; ".
.
. .
','. :;
••
•
. . -.:........;,.---- 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-col-
ume ll a (stc).
118
It - - - stc
.::
•
( , ,.
'
"
• •
,.. "."i: -
. .~
p
,
1--- stc
/
Fig. 70. Weraroa cucullata.
The fruiting body has a nar-
rowly conical pileus with a stc-H
pointed apex, fragile exope- :'
ridium (p) , flexuous stipe-
columella (stc), and gleba (g)
of contorted, anastomosing
lamellae.
119
~--p .
. .
. : I,: "
.. . ..' .
-
. '-/--- g
. I
. . ........
, ", .--';;'-",
.. '
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--- P
.-r-v
120
---p
, '. .
"'-
-rJ~···".··.
", ." .~,. ...--- stc
. .. . .
. :; . ..
"
r--- g
stc
. ,"""'~ '. .
:.-. ' • .1 r-P
,: r·f'l;~~'".
.., ..::Jf;
:.,~., ;,. '
.. ,, .
.
Fig. 74. Longula texensis. The
fruiting body has an oval
'---- stc
pileus with a wrinkled, squam-
ulose exoperidium (p), and
gleba (g) of blackish brown,
brittle gussets (gu). The stipe-
columella (stc) is tough and
longitudinally striate in age.
...
. /
·~~',-_·3 .
121
~---p
I !.
,'
,, ,
p
. ,.
.. .
~ .'
-\-- g
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
spores have been carried out, or other critical data are missing. I
1
123
GLOSSARY
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
1
»
127
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.
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 par-
ticles, 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 corre-
lated 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 trans-
formed into setae; typical of some species of Bovista and Lycoper-
don, 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
-
134
BIBLIOGRAPHY
Fischer, Ed. 1921. Staheliomyces cinctus nov. gen. Mitteilungen der '
Naturforschenden Gesellschaft in Bern aus dem Jahre 1920, p. 35.
Fischer, Ed. 1924. 27. Zur vergleichenden Morphologie der Frucht-
korper von Staheliomyces, Xylophallus und Mutinus. Mykologische
Beit. Nr. 18, Mitteilungen der Naturforschenden Gesellschaft in Bern
aus dem Jahre 1923, Heft V.
Fischer, Ed. 1933. Gastromyceteae Stahelianae. Ann. Mycol. 31 :113- 1
125. 1
Fitzpatrick, H. M. 1913. A comparative study of the development of the
fruit body in Phallogaster, Hysterangium, and Cautieria. Ann. MycoL !
11:119-149. l,
Fogel, R. 1979. The genus Leucophleps (Basidiomycotina, Leuco- l
gastrales). Can. J. Bot. 57:1718-1728.
Fogel, R., and J. M. Trappe. 1985. Destuntzia, a new genus in the .
Hymenogastraceae (Basidiomycotina). Mycologia 77:732-742.
Furtado, J. 5., and D. M. Dring. 1967. The rediscovery of Protubera ~
maracuja, with additional descriptive notes. Trans. Brit. MycoL Soc.
50: 500-502.
Garner, J. H. B. 1956. Gasteromycetes from Panama and Costa Rica.
Mycologia 48:757-764.
Giumann, E. 1926. "Vergleichende Morphologie der Pilze". Verlag von
Gustav Fischer, Jena, 626 p.
Guzman, G. 1970. Monografia del genero Scleroderma. Darwiniana
16: 233-407.
Harding, P. R., Jr. 1957. Notes on Longu/a texensis var. major. Mycologia
49: 273-276.
Harkin, J. M., M. J. Larsen, and J. R. Obst. 1974. Use of syringaldazine for
detection of laccase in sporophores of wood-rotting fungi. Mycolo-
gia 66:469-476.
Herrera, T. 1953. Un hongo nuevo procedente del estado de San Luis
POtOSI Battarreoides potosinus gen. nov. sp. nov. An. Inst. BioL,
Mexico 24:41-46.
Hollos, L. 1904. "Die Gasteromyceten Ungarns". Leipzig. 278 p.
,
Horak, E. 1964a. Fungi austroamericani III. Rhodogaster gen. nov.-a .
new link from Chile towards the Rhodophyllaceae. Sydowia 17:190-
192.
Horak, E. 1964b. Fungi austroamericani V. Beitrag zur Kenntnis der
Cattungen Hysterangium Vitt., Hymenogaster Vitt., Hydnangium
Wallr. und Me/anogaster Cda. in Sudamerika. Sydowia 17:197-205.
137
Mil ler, O. K., Jr. 1983. Ectomycorrhizae in the Agaricales and Gastero-
mycetes. Can J. Bot. 61 :909-916.
Mi ller, o. K., Jr., and H. H. Miller. 1981. "Mushrooms in Color". E. P.
Dutton, Inc., New York. 286 p.
Miller, o. K., Jr., H. H. Burdsall, Jr., G. A. Laursen, and I. B. Sachs. 1980.
The status of Calvatia cretacea in arctic and alpine tundra. Can. J. Bot.
58:2533-2542.
Mi ller, O. K., Jr., and B. Askew. 1982. The genus Gastrosporium in North
America. Can. J. Bot. 60:364-368.
Mi ller, O. K., Jr., and R. Watling. 1987. Whence Cometh the Agarics? A
reappraisal. pp. 435-448 in: "Evolutionary Biology of the Fungi",
Rayner, Brasier and Moore, eds. Univ. of Cambridge Press, Cam-
. bridge, England. 465 p.
M iller, S. L. 1985. Basidiosporogenesis and Developmental Anatomy of
Spore Release in the Russulales-a Systematic Interpretation. Doc-
toral dissertation, VPI &, Blacksburg, VA, 193 p.
M itchell, D. H., S. W. Chapman, and G. Grimes. 1975. Studies of
Disciseda (Gasteromycetes in Colorado). Mycologia 67:586-596.
Moller, A. 1895. Brasilische Pilzblumen, pp. 146-148, illus. In: "Bota-
nische Mittheilungen aus den Tropen, Heft 7." A. F. W. Schimper, ed.
Verlag von G. Fischer.
Morse, E. E. 1933. A study of the genus Podaxis. Mycologia 25: 1-33.
Morse, E. E. 1935. A new puffball. Mycologia 27:96-101.
Palmer, J. T. 1968. A chronological catalogue of the literature to the
British Gasteromycetes. Nova Hedw. 15:65-178.
Pa rady, B. J., and W. J. Sundberg. 1977. A survey of the Nidulariaceae
(bird's nest fungi). Trans. III. State Acad. Sci. 70:314-322.
Pegler, D. N., and T. W. K. Young. 1981. "Basidiospore Morphology in
the Agaricales". Beih. Nova Hedwigia 35. J. Cramer, Lehre. 210 p, 53
pI.
Pi lat, A. 1934. Sur Ie genre Gastrosporium Mattirolo (Gasteromycetes).
Bull. Soc. Mycol. Fr. 50:37-49.
Pi lat, A. 1958. "Gasteromycetes." Prace Ceskoslovenske Akademie ved
Praha. 862 p, illus.
Po nce de Leon, P. 1970. Revision of the genus Vascellum (Lycoperda-
ceae). Fieldiana: Botany 32:109-125.
Po nce de Leon, P. 1971. Revision of the genus Morganella (Lycoperda-
ceae). Fieldiana: Botany 34:27-44.
Rea, P. M. 1942. Fungi of Southern California. I. Mycologia 34:563-574.
140
•
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
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
invertebrate herbivory 4
involute ray 66, 127
/tajahya 77, 78, 87
jamaciensis, Protuber 89
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
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
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
•
.-
--';""
,.
,..,
.. ,
..." "
,,.'.
,.
•
•
111
091 ,
Gasteromycetes: MorphologIcal &
Developmental Features
" - •
•
....
•