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MORPHOLOGY

AND

TAXONOMY OF FUNGI

MORPHOLOGY

AND

TAXONOMY OF FUNGI

Ernst Atnearn Bessey, Pk.D.

Distinguisnea Professor or Botany, Retired, ana

Dean Emeritus or tne Scnool or Graduate

Studies, Micnigan State College

Pkiladelpliia • THE BLAKISTON COMPANY • Toronto

1950

This hook is fully protected by copyright, and no

part of it, with the exception of short quotations

for review, may be reproduced without the written

consent of the publisher

printed in the united states of AMERICA
BY the maple press COMPANY, YORK, PA.

Dedicated to the memory of two
great teachers:

Charles Edwin Bessey

and

Georg Klebs

PREFACE

In the over fifteen years that have elapsed since the pubhcation of the
author's "A Text-book of Mycology,"^ the subject has experienced many
changes. The subject of Medical Mycology has gained greatly in interest
and there has arisen a great realization of the importance of those fungi
whose antibiotic products have become indispensable in the treatment
of many maladies of Man and other animals. It has therefore become
apparent that an entirely new work is necessary, not merely as a textbook
for students but also to assist the many persons now undertaking research
in the various fields of study in which the knowledge of fungi is funda-
mental to their work. Although in some features the present work follows
the framework of the older Text-book it has been entirely rewritten and
greatly enlarged.

Aside from the above-mentioned special fields of study, investigations
of the microscopic soil and water fungi, both parasitic and saprophytic,
which comprise the simplest forms, in structure at least, have revealed
hundreds of species and many new genera, families, and even orders. Thus
great numbers of fungi are now known where but few were recognized less
than two decades ago. This has compelled radical revisions in the classi-
fication of the lower fungi.

In the higher fungi, especially the Basidiomyceteae, many new species
and genera have been described. Above all, studies in the anatomy and
chemical reactions of these new as well as of the old, long known species
have shown the artificiality of the old Friesian system of classification.
This has led to radical revisions which beyond doubt indicate far more
correctly the true relationships and phylogeny of the members of this
class. Yet here, there still remains so much to be done that mycologists
are not at all in agreement as to the ultimate system of classification to
be used.

With so many points still in disagreement, it is impossible for a book
of this type to settle these matters beyond dispute. The author, therefore,
while taking a middle ground in many disputed areas, has attempted to
show those using this book the various points of conflict and opposing

1 Bessey, Ernst Athearn: A Text-book of Mycology, xv -f- 495 pp., 139 tigs.,
Philadelphia, Blakiston, 1935.

vii

viii PREFACE

ideas so that they may judge for themselves what seems to be the more
logical. Accordingly numerous references are given at the close of each
chapter from which these conflicting ideas may be sought. It is the hope
of the author that the recognition of these conflicts may stimulate further
study until the points at issue may be settled by a fuller knowledge of the
facts.

In this work an attempt has been made to give the user the oppor-
tunity to learn the more important characters of almost all the generally
recognized orders of fungi, and to have a foundation upon which the more
intensive study of any of these groups may be based. So many of these
organisms are still known but scantily, so that their true relationships are
very doubtful. As examples, the Protomycetales and Eccrinales may be
mentioned. Many species of each order have been recognized, but the
gaps in knowledge still make their relationships to other fungi uncertain.

Little more than mere mention is made of those fungi that make up
the subject of Medical Mycology, as the author does not feel competent
to enter far into that field. Furthermore, he feels that a rather broad
foundation in the knowledge of fungi in general, such as this book should
give the student, is absolutely necessary before reasonable progress can
be made in the study of such a specialized subject. Similarly, but little
discussion is made of the physiology of fungi, as that subject, too, requires
a good knowledge of the fungi themselves as well as of the physiology of
green plants and an adequate training in chemistry. The genetics of fungi
is scarcely touched upon except where it is necessary to understand some-
what the relationships of certain fungus groups. In other words, this book
must not be expected to be a complete encyclopedia of fungi, but rather
the foundation upon which to lay the various superstructures of the edi-
fice of Mycology.

So much has been published in the last fifteen years or more in the way
of manuals and monographs that the final chapter, "Guide to the Litera-
ture for the Identification of Fungi," is already large in size, in spite of
the fact that many older references have been omitted where newer ones
are readily available. Here, especially, the effects of World War II have
been deplorable. Contacts with mycologists in many portions of the world
have been broken; continuity of exchange of literature has been inter-
rupted. The mycologic publications in many of the countries where fight-
ing was intense have been destroyed, so that much that was of value can-
not even be located. As a consequence, in this chapter some glaring omis-
sions doubtless Mdll become apparent. It is hoped that these will not lessen
the value of the book.

As before, the author owes a great deal to the generous cooperation of
many botanists in the New World and in portions of Europe. To begin to
name them would result in a list too long to be printed here. Furthermore,

PREFACE IX

he expresses his gratitude to the many mycologic friends who have assisted
him by granting permission to use illustrations from their publications.
To the management of Mycologia especial thanks are due for their blanket
permission to use illustrations from that indispensable periodical. To
M. B. Walton, of Cleveland, Ohio, the author's thanks are due for the
generous gift of prints of some of his wonderfully fine photographs of fungi.
To his wife, especially, the author owes thanks for the continued
stimulus to persist in the arduous work of preparing this new book, for
writing the manuscript, for assistance in reading proof, and for attending
to the many other tiring details that go into the making of a book.

Ernst A. Bessey

Distinguished Professor of Botany, retired

Michigan State College
East Lansing, Michigan
Jiine 1950.

CONTENTS

Preface vii

1. Introduction 1

2. Mycetozoa and Related Organisms 22

3. Phycomyceteae : Chytridiales and Hyphochytriales 42

4. Phycomyceteae: Blastocladiales and Monoblephar-

IDALES 78

5. Phycomyceteae: Lagenidiales and Saprolegniales 94

6. Phycomyceteae: Peronosporales and Protomyce-

tales 126

7. Phycomyceteae: Mucorales, Entomophthorales, Zoo-

pagales, Eccrinales 150

8. The Higher Fungi: Carpomyceteae 192

9. Class Ascomyceteae: Laboulbeniales and Discomy-

cetes 200

10. Class Ascomyceteae: The ''Pyrenomycetes" 262

11. Class Ascomyceteae: Erysiphales, Aspergillales,

Myriangiales, Saccharomycetales 307

12. Class Basidiomyceteae: Subclass Teliosporeae 366

13. Class Basidiomyceteae: Subclass Heterobasidiae. .. . 436

14. Class Basidiomyceteae: Subclass Eubasidiae, ''Hy-

menomyceteae " 464

15. Class Basidiomyceteae: Subclass Eubasidiae, ^'Gas-

teromyceteae " 530

16. Fungi Imperfecta The Imperfect Fungi 572

17. The Phylogeny of the Fungi 628

XI 1 CONTENTS

18. Guide to the Literature for the Identification of

Fungi 660

List 1. General Works Covering the Whole Field of Systematic

Mycology 662

2. Host Indexes, Local Fungus Lists with Host Indexes,

Lists of Fungi on Special Hosts or Substrata, Bibli-
ographies, Fungi of Man and Other Animals 668

3. Mycetozoa, Including Myxogastrales, Plasmodiophor-

ales, Acrasiales, Labyrinthulales 672

4. General Works on Phycomyceteae 673

5. Chytridiales 673

6. Hyphochytriales 675

7. Blastocladiales and Monoblepharidales 675

8. Lagenidiales 676

9. Saprolegniales (Including Leptomitales) 676

10. Peronosporales, Also Protomycetales 677

11. Mucorales, Entomophthorales 678

12. Zoopagales, Eccrinales 680

13. Ascomyceteae, Miscellaneous 682

14. Laboulbeniales 682

15. Lecanorales and Pyrenulales 683

16. Pezizales Operculati and Inoperculati, Including "Pha-

cidiales" 688

17. Tuberales 693

18.' Taphrinales 694

19. Hysteriales 695

20. Sphaeriales 695

21. Hypocreales 699

22. Dothideales 700

23. Pseudosphaeriales 701

24. Hemisphaeriales 702

25. Erysiphaceae 703

26. Meliolaceae (Perisporiaceae) 704

27. Remainder of Erysiphales: Capnodiaceae, Ehglerula-

ceae, Trichothyriaceae, Atichiaceae 705

28. Myriangiaceae 706

29. Aspergillales (Plectascales) 707

30. Saccharomycetales and Asporogenous Yeasts 707

31. Ustilaginales (Including Graphiolaceae) 709

32. Uredinales 712

33. Heterobasidiae 718

34. " Hymenomyceteae " : General Works 720

35. Thelephoraceae and Exobasidiaceae 721

CONTENTS Xm

36. Clavariaceae 723

37. Hydnaceae 724

38. Meruliaceae 725

39. Polyporaceae 726

40. Boletaceae 729

41. Agaricaceae (in the Broader Sense) 730

42. Gasteromyceteae — General Works 742

43. Hymenogastrales, Sclerodermatales, etc 743

44. Lycoperdales, including Tulostomataceae and Podax-

aceae 744

45. Nidulariales, including Arachniaceae 746

46. Phallales 746

47. Sphaeropsidales 747

48. Melanconiales 750

49. Moniliales: Moniliaceae 750

50. Moniliales: Dematiaceae 753

51. Moniliales: Tuberculariaceae, Stilbellaceae, and My-

celia Sterilia 755

Index 757

INTRODUCTION

THERE are many kinds of living organism.s whose interactions serve to
maintain or to break the balance of Nature. Some of them, green
plants, build up organic substances from the inorganic matter at their dis-
posal. They are basic to all life on the earth, for without them almost all
other organisms would eventually perish. The animals reach probably the
greatest degrees of complexity but they must take their basic organic
foods directly or indirectly from the chlorophyll-containing plants. Be-
sides these two great groups of organisms there are the numerous bacteria
— minute breakers-down of the complex organic substances built by the
green plants and by the animals and also builders-up of some of the
inorganic substances that the green plants need. Aside from these there
is the great horde of organisms called fungi. Perhaps there are 100,000
species of them, according to Bisby and Ainsworth (1943), but of the
100,000 named fungi probably not over 40,000 are, according to these
authors, valid species, leaving 60,000 or more to be recognized and de-
scribed in the future. It is with the fungi that mycology has to do. Ordi-
narily they are considered by most botanists to be plants, but Martin
(1932) and others who followed him, have suggested that fungi may be
neither animals nor plants but a third kingdom, of common origin with
them, which has undergone a parallel development to the animal and
plant kingdoms.

Fungi

There is no general agreement as to the limits of the forms that should
be called fungi. Some of the earlier mycologists included the bacteria with
them, but that is rarely done nowadays. The majority of botanists, or to
be more specific, of mycologists, include the Slime Molds (Mycetozoa or
Myxomycetes) among the fungi. Yet the great German mycologist and
plant pathologist Anton de Bary (1831-1888) said (1887), "I have since
the year 1858 placed the Myxomycetes under the name Mycetozoa out-
side the limits of the vegetable kingdom, and I still consider this to be

2 INTRODUCTION

their true position." With this viewpoint the author is in agreement, but
since most mycologists accept these organisms as fungi they are included
in this textbook.

From the foregoing it is apparent that the fungi do not possess chloro-
phyll and also that the chlorophyll-less bacteria and Mycetozoa are ex-
cluded from their ranks. There are many other organisms which lack
chlorophyll and still are not fungi. Thus in the diatomaceous genus
Nitzschia some species are known that possess no chloroplasts while the
majority of species possess them. Otherwise they are so similar that they
are retained in the same genus. In the Red Seaweeds (Florideae) there
are several species that lack chlorophyll and are parasitic upon other
Florideae — e.g., Harveyella mirahilis (Reinsch) Schmitz & Reinke (see
Sturch, 1899). Yet in their modes of sexual and asexual reproduction
they can be assigned definite positions among these algae. Among the
higher plants (Anthophyta or Angiospermae) many widely separated
chlorophyll-less species are found, e.g., in the Orchidaceae and Bur-
manniaceae among the Monocotyledoneae, and Cuscuta, Cassytha, Mo-
notropa, Rafflesia, Orohanche, and many others in the Dicotyledoneae.
Thus it is necessary to delimit the fungi by further characters than
merely the lack of chlorophyll. Such a definition is in the main negative.

Definition. As a group the fungi may be defined as chlorophyll-less
nonvascular plants whose reproductive or vegetative structures do not
permit them to be assigned to positions among recognized groups of
algae or higher plants, and as excluding the Bacteria (which are typically
one-celled and lack a typical nucleus) and the Mycetozoa (which have
an animal type of structure and reproduction).

Whether the fungi represent a single phylum of organisms with a
common origin or have arisen in independent lines from several ancestral
types of plants is still a matter of debate among students of their phylog-
eny and classification. They range from very simple short-lived, one-
celled structures whose single cell becomes the organ of reproduction
(e.g., Olpidiopsis, which at maturity becomes the zoosporangium from
which escape the zoospores) to massive perennial mycelia giving rise to
great spore fruits as in some of the puffballs, pore fungi, etc. Except for
the lack of chlorophyll and the saprophytic or parasitic mode of life
thereby necessitated, these two extremes have no single character in
common: manner of reproduction, structure of the vegetative body,
chemical composition of the cell wall, etc. The extreme simplicity of the
one type of fungus might be considered to indicate a low position in
evolution, i.e., great primitiveness, but on the other hand it might be the
result of a great degree of simplification from a much more complex
fungus. The lack of any good fossil record of these lower fungi prevents
us from obtaining direct evidence in this matter.

FUNGI 6

Structure. The vast majority of fungi consist vegetatively of more or
less elongated, septate or nonseptate filaments. These are called indi-
vidually hyphae (singular, hypha) and collectively the mycelium. They
may be uniform in thickness or tapering from broad to slender portions
in the same hypha or in different portions of the same mycelium. They
may be branched or unbranched (simple). In thickness they may be less
than 0.5 jj. up to over 100 n (in some Saprolegniales). In size the whole
mycelium may be only a few microns in length or it may produce great
sheets or strands that extend many meters.

The composition of the cell wall is very variable among the different
fungi and sometimes in the same individual at different stages of ma-
turity. Basically the chief components appear to be various types of
carbohydrates or mixtures of these: cellulose, pectose, callose, etc. Mixed
with these and probably often in chemical combination with them there
may be other substances. Cellulose predominates in many of the Phyco-
myceteae so that the characteristic cellulose reaction is shown upon treat-
ment with chloriodide of zinc, but sometimes where it makes up the bulk
of the wall it does not respond to this reagent until certain fatty deposits
in the outer portion of the cell wall are first dissolved away, as in Mono-
hlepharis. In a great many fungi, especially the Ascomyceteae, Basidio-
myceteae and the higher Phycomyceteae, either cellulose is entirely
lacking, being replaced by some other carbohydrate, or the considerable
amount of chitin with which the wall is impregnated prevents the cellulose
from showing its presence. Chitin is never alone in the wall but it may
form a considerable portion of the component substances. Aside from the
foregoing substances calcium carbonate or other salts may be deposited
upon or within the wall. Although von Wettstein (1921) and others have
identified the chief component of the cell wall of many fungi, apart from
the carbohydrates, as chitin, identical with the chitin of the Arthropods,
Dous and Ziegenspeck (1926) after a careful comparative study of the
animal chitin with the fungus chitin conclude that these are parallel
compounds derived from different basic substances, but with much the
same general characters. Throughout this book, wherever the word chitin
is used it should be understood as referring to this fungus chitin, not to
the true animal chitin.

There are two main types of mycelium ; in one the hyphae are cellular
and in the other, coenocytic. A cellular hypha usually contains either one
or two nuclei per cell and the division of the cell is initiated by the division
of the nucleus or by the simultaneous division of both nuclei, respectively.
In a coenocyte there are many nuclei and the formation of septa occurs
without immediate reference to any preceding nuclear division. A coeno-
cytic hypha may be "tubular," i.e., lacking septa, or septate. In the
latter case each segment is multinuclear. A tubular coenocyte, such as is

4 INTRODUCTION

characteristic of the majority of the Phycomyceteae, produces septa to
set apart the reproductive organs (sporangia or gametangia) from the
hypha, or to fence off an injured region, or to separate an empty portion
of the hypha from those portions still containing protoplasm. A cellular
mycelium may have multinucleate cells in the younger portions but by
intervention of septa the older portion is transformed to cells with one
or two nuclei, depending upon the phase of the mycelium. An old my-
celium, no longer able to continue normal growth, may undergo nuclear
division without septum formation so that the old cells may have several
nuclei. In the Higher Fungi that generation of development in which the
cells are uninuclear is sometimes called the monocaryon phase and that
in which the cells are binuclear the dicaryon phase, or primary and
secondary mycelium, respectively. In general the growth in length and
the formation of new cells takes place in the terminal portion of the
hypha or of its branches.

Septum formation occurs, as in most algae, by the production of a
circular shelf which gradually grows inward until (in the Phycomyceteae)
it makes a complete septum or (in the Higher Fungi) leaves a small
central perforation through which there is a continuous protoplasmic
connection from cell to cell. In Allomyces, one of the Phycomyceteae,
septa may be formed but these are imperfect with large openings and are
called pseudosepta.

The mycelia of the majority of fungi are hyaline, especially the hyphae
that are embedded in the substratum and functioning as organs for ob-
taining nutriment. These are sometimes colored but the color is much
more frequent in those hyphae that are external. The pigment causing
this dark color is related to melanin and is largely confined to the cell
walls. The hyphae that bear the conidia or that protect the other types
of reproductive structures are especially apt to be dark-colored.

Aside from their elongated thread-like form the hyphae may be
packed together tightly so that they adhere in elongated strands which
sometimes have a hard black external layer and creep long distances.
These are called rhizomorphs. In ArmiUariella mellea (Vahl) Karst., these
black, shoestring-like strands creep under the bark of roots and trunks
of trees and through the soil. Storage organs called sclerotia are frequently
produced by fungi. Dense masses of hyphae arise and the short cells
enlarge laterally until a compact pseudoparenchymatous tissue is formed
whose cells become polyhedral by mutual pressure. These are filled
with food materials and the walls may remain thin or become somewhat
thickened, especially the outermost layers of cells which thus form a
protective cortex which may be light in color or more often brown or
black. These sclerotia if small, as in Sclerotium rolfsii Sacc, may be trans-
ported easily by surface water during heavy rains. Larger sclerotia may

FUNGI

remain near where they are produced and then develop typical repro-
ductive organs for the fungus when favorable conditions arrive. This is
the case with sclerotia of Claviceps, Sclerotinia, and some species of
Polyporus. From the sclerotia themselves under certain conditions new
mycehum may grow out instead of reproductive organs. This is the case
with the overwintering stage of a species of Pelliculana, the common black
scurf {" Rhizoctonia") of potatoes and other plants.

Reproduction. Asexual reproduction in the true fungi may occur by
the formation within a zoosporangium of naked cells, zoospores, which
upon their release swim away by means of anteriorly, laterally, or pos-
teriorly attached flagella, either one or two in number depending upon
the order of fungi concerned. This production of zoospores is confined to
some orders of the Phycomyceteae. The zoospores eventually settle down
and encyst, and the encysted cell becomes the start of the new plant. In
the majority of fungi, including many of the Phycomyceteae and all of
the Higher Fungi in which asexual reproduction occurs, no motile spores
are produced but the spores are provided with a wall and are distributed
by air currents, by water, by insects, etc. These spores are of several
types of origin. In the Mucoraceae they are produced internally in a
sporangium; upon the rupture or dissolution of its walls, they are set free
and distributed by air or water currents. Such spores are sometimes called
aplanospores in contradistinction to the motile naked zoospores which
may be called planospores. Conidia arise as single separable cells of the
mycelium. They may arise by the fragmentation of the whole mycelium
or of special hyphae into cylindrical, ovoid, or spherical cells (oidial mode
of conidium formation) or by the cutting off of terminal or lateral cells
from special hyphae or conidiophores. In a number of genera the conidia
are pushed out one by one from the neck of a flask-like cell (or phialid).
The conidia, by whatever means they arise, may be hyaline or colored,
and may remain one-celled or by formation of transverse or longitudinal
septa may become two-celled to many-celled. They may be released
singly or remain attached in a chain. In some cases, instead of producing
conidia, a tangled mass of hyphae may form a rounded ball as in the
genus Papulospora. For the purpose of enabling the fungus to survive
unfavorable conditions such as cold, lack of water, etc., chlamydospores
are produced by many fungi. These are terminal or intercalary cells of a
hypha (or even single cells in a conidium made up of a row of cells) which
enlarge and round up, store supplies of food, and form a thick wall. Such
cells may live for years until favorable conditions arrive. They have
nothing to do with the sexual stage of the fungus so that the use of the
term chlamydospore for the reproductive cells (teliospores) of the
Ustilaginales is unwarranted.

Sexual reproduction, or substitutes for it, may be found in most of

6 INTRODUCTION

the groups of fungi except the artificial class called Fungi Imperfecti
which was estabhshed to include those fungi in which sexual reproduction
has not been discovered. The simplest type of sexual reproduction is the
union of two cells of equal size and to all appearances alike in all charac-
teristics. These two gametes may both be nonmotile (aplanogametes) or
motile (planogametes), as in the Yeasts and Chytridiales, respectively.
The zygote formed by their union may become a new one-celled plant
or may produce mycelium of various types. The gametes are more often
unequal in size and the motility may be limited only to the smaller one,
then designated as the male gamete or sperm. Still more often flagella
are lacking in both gametes and fertilization is brought about by the
union of the cells by dissolution of a portion of the intervening walls.
Sometimes the male nucleus is introduced into the female gamete, or egg,
through a tube extending into the latter. When the zygote is the product
of the union of clearly dissimilar gametes, forming a definite zygote cell
which often serves as a resting spore, it may be called an oospore. When
a similar resting spore results from the union of similar gametes it is
called a zygospore. It must be noted that the gradation between iso-
gametes (i.e., equal gametes) and anisogametes (i.e., gametes that are
dissimilar) is gradual.

In some fungi, instead of producing definite gametes, any cell of one
mycelium may unite with any cell of another compatible mycelium, so
that no definite part of the fungus can be distinguished as a male or
female reproductive organ. This is especially true in the Ustilaginaceae.
However, many fungi do produce clearly distinguishable male and female
organs of reproduction. Very often the gametes may develop into new
plants without union, i.e., by parthenogenesis. This phenomenon is ob-
served from some of the simplest fungi up to many of the Higher Fungi.
Sometimes a vegetative cell may be substituted for the normal male
gametangium (antherid), the nucleus of this substituting cell functioning
in place of a normal sperm nucleus. So it comes about that, with partheno-
genesis occurring in many fungi and substitution of vegetative cells for
gametes in others, it is difficult to follow the evolutionary sequence of the
development of sexual reproduction in these organisms.

The occurrence of the union of the sexual nuclei naturally leads to the
production of a diploid nucleus. In very few fungi does this nucleus
multiply in the diploid state (perhaps in a few Yeasts). Usually its first
divisions are meiotic, so that throughout the life history of the fungus
the nuclei are always haploid except immediately after the union of the
gamete nuclei. Yet among the fungi, particularly in the Ascomyceteae
and Basidiomyceteae, we find a contrast of mycelia that are diploid in
nature and haploid in nature although all the nuclei are haploid. These
are respectively the dicaryon and monocaryon phases of mycelia. Whether

FUNGI

the two nuclei (of separate sexual origin) are present in a cell within one
nuclear membrane (i.e., a diploid nucleus) or each in its own separate
membrane (i.e., two haploid nuclei in the cell), the effect on the cytoplasm
is practically the same, so that a dicaryon cell is to all intents and pur-
poses a diploid cell. Thus when Buller (1930) speaks of the "diploidiza-
tion" of a monocaryon mycelium by the introduction of a compatible
haploid nucleus which multiphes and spreads from cell to cell this term
is essentially correct although the nuclei, now in pairs in each cell, are
still haploid. He was speaking of diploidization of the cell as a whole
not of the contained nucleus.

In the simplest fungi the whole plant, consisting of but one cell,
becomes the reproductive unit that produces the asexual or sexual cells.
As we study fungi of greater complexity we find that the vegetative and
reproductive portions of the organism are more and more segregated.
In many fungi the process of sexual reproduction becomes increasingly
complex and leads to the formation of, not a single zygote, but a very
complex structure, the spore fruit, many of whose cells become the ulti-
mate reproductive spores. Some of the puffballs, e.g., Calvatia gigantea
(Batsch ex Pers.) Lloyd, attain a diameter of over one meter. This is the
spore fruit. The vegetative mycelium is subterranean and not noticed
except when hunted for.

Parasitism. Because of their lack of chlorophyll all fungi must obtain
their organic food from sources external to themselves. The whole my-
celium may have the power to absorb these foods or this task may be
relegated to special portions such as rhizoids or to haustoria, the knob-like
or finger-like processes that enter the cells of the host plant. In many of
the Phycomyceteae there is a great difference between the much branched
mycelium within the substratum and that portion outside which bears
the reproductive organs.

Probably the majority of fungi are saprophytic, i.e., feed upon the
organic products or remains of plants or animals but not upon the hving
organisms themselves. The substances utilized by the fungus are often
very varied in nature. Simple sugars, starches, cellulose, or organic acids
may satisfy its needs provided the necessary mineral nutrients are present.
On the other hand some fungi require the presence of various growth
factors, such as biotin and thiamin, and some proteins or their building
blocks, the amino acids. Some saprophytic fungi are much more limited
as to their organic foodstuffs. In general, however, their range of foods
is far wider than that of strict parasites. Some facultative parasites are
able to grow apparently indefinitely as saprophytes. Thus a species of
Fusarium capable of causing the death of Sesamum indicum L. was studied
in culture by the author (1904). When provided with the necessary
mineral nutrients, it developed upon cellulose, simple and complex sugars.

8 INTRODUCTION

various organic acids, asparagin, peptone, gelatin, glycerine, etc., and
under both aerobic and anaerobic conditions.

On the other hand many fungi are obhgatory parasites and methods
have not yet been devised by which they can be grown except upon
suitable living host tissues. This is true of the Rusts (Uredinales), the
White Rusts (Albuginaceae), most of the Downy Mildews (Peronospo-
raceae), the minute insect parasites belonging to the Laboulbeniales, etc.
This would seem to indicate that for such parasites the choice of food is
very strictly limited. Among some of the commoner parasites, e.g., stem
rusts of wheat, Puccinia graminis tritici Erikss. & Henn., and powdery
mildew of various grasses, Erysiphe graminis DC, there have been
found what are called biologic or physiologic forms. These are races of
the fungus that are indistinguishable except for the fact that one will
grow only on certain species or varieties while the other races grow only
on other varieties.

Parasitic fungi may be destructive or balanced parasites. The former
may kill the host cells or tissues by means of some poisonous substances
which may even diffuse out in advance of the fungus hyphae with the
consequence that the latter actually enter dead tissues. Some species of
Botrytis, Sderotinia, Pythium, etc., are of this type. On the other hand a
balanced parasite is so well adapted in its demands upon the host to the
ability of the latter to supply these needs and to continue to live and
grow that both fungus and host develop together until the time comes
when the fungus is ready to produce its spores. Then it too destroys the
surrounding tissues. Many of the Smuts (Ustilaginales) are balanced
parasites. Actually all gradations between these two extremes may be
found. Mostly the destructive parasites are less strictly confined to
definite hosts than are the balanced parasites.

Some fungi belonging to the Polyporaceae attack and destroy only
the dead cells (wood fibers, tracheary tissues, etc.) of the wood of living
trees and from that viewpoint are saprophytes, yet their growth ceases
when the death of the tree occurs. The conditions within the host tissues
that favor the growth of the fungus are evidently sufficiently changed
when the tree dies so that the fungus no longer finds the conditions of
environment that are requisite for its continued growth. Many fungi are
parasitic in certain stages of their growth and saprophytic later on. Thus
the fungus of apple scab {Venturia inaequalis (Cke.) Wint.) grows and
produces its conidia upon the living leaves and fruits of the apple but
overwinters saprophytically and produces its sexual stage of reproduction
within the dead leaves on the giound. This is true of very many of the
leaf spot fungi of economic as well as wild plants.

Just how a parasitic fungus obtains its food from its host is not clear
in all cases. Apparently it may be by the action of some secretion from

HISTORY OF MYCOLOGY 9

the fungus hyphae or haustoria upon the plasma membrane of the host
cells, making this more permeable to the contained solutes so that they
diffuse out and are absorbed by the fungus.

The whole field of fungus physiology offers many interesting lines of
study and cannot be entered upon within the limits of this book. Nor
will space permit the extensive discussion of medical mycology (see
C. W. Dodge, 1935) or technical mycology (Lafar, 1903, 1910). Only
brief mention is made in Chapter 16 of the production of antibiotics. The
genetics of fungi is noted where necessary for the understanding of the
development of various groups of these organisms.

History of Mycology

A very brief sketch of a few steps in the history of mycology should
not be omitted. The larger fungi, or rather their conspicuous fruiting
bodies, were well known to the ancients, but knowledge of their true
nature and their manner of growth had to await the invention of the
microscope. The Romans knew and distinguished various edible and
poisonous mushrooms. The Emperor Nero is reported to have been very
fond of Amamta caesarea (Schaeff.) Fr., which owes its specific epithet to
this association. The word fungus (related to the verb fungor, to flourish)
was applied to mushrooms and to excrescences from the ground or from
trees. The Greek word mykes (hvktjs) was applied to some types of fungi.
From this comes the characteristic part of the word mycology. For untold
centuries the Chinese have known and used certain fungi for food and
others for medicine but, as in the Occident, with httle real knowledge as
to the true nature of these organisms.

After the invention of printing in Europe there began to appear
various "herbals," describing and, in many cases, illustrating more or
less elaborately the plants of southern and western Europe. In some of
these the larger fungi are illustrated. Thus Clusius (Charles de la Cluse,
1529-1609) in 1601 devoted many illustrations and many pages of text
to the discussion of edible and poisonous fungi. No attempt was made to
classify these into genera or families as these terms are now used. In 1623
in his "Pinax Theatri Botanici," Gaspard Bauhin (1560-1624) attempted
to bring together all plants known to him or to his predecessors. He
divided the approximately 100 species of fungi and lichens into groups
to which he gave names. The idea of the genus as a definite category for
the purposes of classification had not yet become firmly established, so
that some of his group names include directly, as the next subordinate
rank, the species, while in other cases there are intermediate categories.
All lichens he included in the group Muscus Saxatilis vel Lichen (9 species) .
Under the name Fungus he included 81 species which are distributed now
among the Agaricaceae, Boletaceae, Polyporaceae, Clavariaceae, Auricu-

10 INTRODUCTION

lariaceae, Lycoperdaceae, Phallaceae, Clathraceae, Pezizaceae, and per-
haps other famihes. Agaricum Fungus corresponds practically to the
laterally attached Polyporaceae, especially Fomes. Tuhera, with 2 species,
was applied to truffles (Tuber) and other subterranean firm fungi.

Tournefort (1656-1708) is the botanist who, more than any other,
brought to general acceptance the concept of the genus as the classifi-
catory category next above the species. He still maintained the cumber-
some method of naming a species with the genus name followed by a
descriptive phrase, now universally abandoned for the binomial manner
of writing a name which was popularized by Linnaeus. Six genera of fungi
and one of lichens were recognized by Tournefort in his "Elemens de
Botanique" in 1694. The generic names used were adopted from his
predecessors. Fungus corresponds to all centrally stipitate Agaricaceae,
Boletaceae, and Polyporaceae. Boletus includes Morchella, Clathrus, and
Phallus. Agaricus was applied to fungi attached laterally to trees, logs,
etc., such as various Polyporaceae, Auricularia, etc. Lycoperdon included
the Lycoperdaceae and also the larger rounded Mycetozoa. Coralloides
included various branched fungi, among others the branching species of
Clavaria. Tuhera was used as by Bauhin.

Dillenius (1687-1747) added a good many species and some new
genera. He also changed the names of some groups from those used by
Tournefort or, retaining the name, changed its application. Thus all
centrally stipitate Agaricaceae were placed in the genus Amanita. Boletus
was entirely changed and made to include the present Boletaceae and
centrally stipitate Polyporaceae. Morchella and Phallus were introduced
to take up the species included in Tournefort's Boletus. Bovista was substi-
tuted for Lycoperdon and Fungoides for Coralloides. For cup- or saucer-
shaped fungi Dillenius used the name Peziza. Mention must be made of
Sebastien Vaillant (1669-1722) whose book "Botanicon Parisiense" in
1727 gave illustrations of fungi and other plants whose accuracy and
beauty were scarcely equalled for over a century. He listed all genera
alphabetically, regardless of their real relationship, hence the fungi are
scattered throughout the work. Agaricus and Boletus are used as they
were by Tournefort. Most of the Agaricaceae are included in the genus
Fungus which is however a very heterogeneous assemblage of organisms.
Fungoides is in part Peziza, and Corallofungus includes some of the species
of Clavaria. Aside from the beautiful illustrations and the descriptions,
Vaillant added little to mycology.

The foremost student of fungi before the time of Linnaeus was the
Italian botanist Pier' Antonio MicheH (1679-1737). He was apparently
the first student of these organisms to use the microscope on them, crude
as was his instrument. His great work "Nova Plantarum Genera" was
completed by 1719 but, for lack of funds, the first part only was pub-

HISTORY OF MYCOLOGY 11

lished, after a delay of ten years, in 1729. His biographer Targioni-
Tozzetti (1858), reports that the second part was completed but never
published for lack of means. There is no doubt that Micheli knew his
fungi far better than any of his forerunners or contemporaries. He gave
usable keys by which genera could be identified and, for the larger genera,
keys to the species. Many of his figures and descriptions were so excellent
that there is no difficulty now in identifying them. He distinguished Fungi
lamellati (Agaricaceae), Fungi porosi (Polyporaceae and Boletaceae),
Fungi pulverentes (Lycoperdaceae and some others), Fungi Ramosi (the
branching Clavariaceae), etc. Among the generic names used by him and
still recognized are Phallus, Clavaria, Clathrus, Lycoperdon, Geaster, and
Tuber. He used Agaricum as did his forerunners for laterally attached
Fomes, Trametes, Fistulina, and Stereum. Polyporus was confined to the
stipitate polypores; Suillus to the present Boletus and its allies; Erinaceus
to the stipitate Hydnaceae; Fungus to the stipitate Agaricaceae. Boletus
as used by him is now known as Morchella; his Puccinia is now called
Gymno sporangium. Coralloides was equivalent to the branched species of
Clavaria, etc. In addition to collecting and studying the larger fungi,
Micheli was perhaps the first botanist to attempt cultures of molds. He
sowed spores of "il/wcor" (evidently Rhizopus nigricans Ehr.) on one side
of pieces of squash and ^^ Aspergillus^^ on the other. Each produced its
own kind of fungus. He inoculated two pieces of squash with " Botrytis,"
covering one with a bell jar and leaving the other exposed. The covered
piece developed only Botrytis while the uncovered piece developed Mucor
as well, thus showing, as Micheli pointed out, that the spores of these
various molds were distributed through the air.

Linnaeus (Carl von Linne, 1707-1778), who is often called the "Father
of Botany," advanced the knowledge of fungi little if at all. In his great
work "Species Plantarum" (1753) he attempted to bring together de-
scriptions of all of the known species of plants. His adoption of the two-
word form of name for species (which we call generic and specific epithets)
marked a very great advance in convenience and simplicity. In his
twenty-fourth class, "Cryptogamia," the fungi are to be found chiefly
under the heading Cryptogamia Fungi but a few are located among the
Cryptogamia Algae. His treatment of the fungi, which he mostly knew
only from the study of botanical literature, and only superficially at first
hand, is far less scientific than that of Micheli or of Dillenius. The lichens
were included in the genus Lichen, among the algae, as was the genus
Tremella. This genus includes the alga Nostoc as well as the rust Gymno-
sporangium, the Basidiomycete Auricularia, several lichens, and probably
one or more species now included in Tremella. All of the Agaricaceae, as
we know the family, were included in the genus Agaricus and all the pore
fungi in the genus Boletus, differing from Micheli and his predecessors.

12 INTRODUCTION

Erinaceus of Micheli and earlier botanists became Hydnum. Phallus was
made to include both Phallus and Morchella, being equivalent to Tourne-
fort's Boletus. Lycoperdon included Lycoperdaceae and some Mycetozoa.
Mucor included all molds such as Mucorales, Fungi Imperfecti, and
Erysiphaceae. Other genera recognized by Linnaeus were Elvela, Peziza,
and C lav aria.

The most significant advance in the classification of fungi after
Linnaeus is to be found in the works of Christiaan Hendrik Persoon
(1755-1837). The number of recognized species had become greatly in-
creased and the great improvements in the microscope made it possible
to study the manner by which the spores were borne, so that the major
groups as now recognized began to appear.

Probably the greatest contribution to the knowledge of the larger
fungi, particularly the "Hymenomycetes," was made by Elias Magnus
Fries (1794-1878), whose active mycological work extended over a period
of more than half a century. The impetus given to mycology by these
two great botanists was felt over the whole world and fungi unknown to
science were discovered by the thousands.

In the first third of the nineteenth century the smaller Ascomyceteae,
especially Sphaeriales, and the pycnidial Fungi Imperfecti, were mostly
described superficially, often being thrown together in the same genus.
Little was known of the Rusts, Smuts, molds of all sorts, various Monili-
ales, Melanconiales, etc. Only when these were studied carefully with the
compound microscope did order begin to arise out of chaos. Of the many
workers in that period, mention may be made of August Carl Joseph
Corda (1809-1849) whose "Icones Fungorum," a six-volume work, pub-
lished from 1837-1854, showed the detailed structure of many of the
larger fungi but also threw light on hundreds of the microscopic forms.
Soon following this came the beautifully illustrated three-volume work of
the Tulasne brothers, "Selecta Fungorum Carpologia," 1861-1865 (Louis
Rene Tulasne, 1815-1885, being the chief author).

In the United States the first extensive study of fungi was undertaken
by Lewis David von Schweinitz (1780-1834), a minister in the United
Brethren Church, who collected extensively in North Carolina and
Pennsylvania. His publications on American fungi (1822 and 1832) were
the first noteworthy ones that appeared. A third of a century later Charles
Horton Peck (1833-1917) began his work at Albany, New York, as state
botanist, a position which he held from 1867-1915. His chief interest was
in the fungi of which he described about 2500 species previously un-
recognized. His collections and descriptions formed the foundation for
many monographic studies of various genera by later students, especially
of the Agaricaceae. Other nations had similar lovers of fungi who added
greatly to the knowledge of these organisms, but space does not permit

HISTORY OF MYCOLOGY 13

the mention of their names, except Carlos Spegazzini (1858-1926), of
Argentina, who explored the mycologically almost unknown territory of
southern South America and gave the first descriptions of several thousand
species.

As new species of fungi were recognized in all parts of the world their
descriptions appeared in all sorts of scientific journals, reports of learned
societies, and even in textbooks, so that it became increasingly difficult
for a student of fungi to know whether a fungus under study by him
was new to science or already described. Many fungi were unavoidably
named several times by different investigators. In Germany and other
European countries floras were published in which were described all
species of fungi known to occur in those regions, such, for example, as
Rabenhorst's "Kryptogamen-Flora" (1844, 1845). Valuable as were such
works, they did not include fungi from other regions, and since fungi are
much more cosmopolitan in their distribution than higher plants the
probability always existed that many fungi described from elsewhere
would be found to occur within the area covered by the work. This un-
certainty deterred many mycologists from describing supposedly new
species for fear of duplication. This was especially true of students of
fungi who lived away from the great European centers of mycological
activity. This condition became so bad that the great Italian mycologist
Pier' Andrea Saccardo (1845-1920) decided, before 1880, to bring to-
gether in one work the descriptions of all fungi hitherto recognized. Thus
began the monumental "Sylloge Fungorum," the first volume of which
appeared in 1882 and the twenty-fifth in 1931. With the appearance of
the first volume of this work systematic mycology again took a great
leap forward. In the meantime various periodicals were established which
were devoted partly or entirely to fungi. These are found in many
countries and in many languages, for the science of mycology is bounded
by no political or linguistic boundaries.

With the greatly increased knowledge of the structures of fungi, for
which the Tulasne brothers were in great degree responsible, there began
a new phase of mycological work about the middle of the nineteenth
century. This was the study of the life histories of fungi. The earlier
mycologists had been, in the main, satisfied to describe the different forms
as they found them, often not dreaming that in many cases they were
treating of different stages of the same organism. To be sure some of the
earlier describers of fungi suggested that different forms found in close
association might be different stages of the same fungus, suggestions
which in many cases were found by later investigators to be correct. The
work of Anton de Bary began at about this period and was carried on
with such enthusiasm and skill that a great series of life history investi-
gations followed. His first outstanding work was his investigation of the

14 INTRODUCTION

life history of the Mycetezoa (1859). He maintained his interest in this
group of organisms for many years and inspired the work of many
outstanding students in this field. Probably the most outstanding of
de Bary's earlier studies was that in which he determined the life cycle
of the rusts (Uredinales) and proved the heteroecious nature of black
stem rust of small grains {Puccinia graminis Pers.). In the two decades
after the appearance of this work (1865) we find many of de Bary's pupils
following and extending life history studies in all groups of fungi. Pre-
eminent among these was Oscar Brefeld, who employed, not for the first
time but probably most extensively up to then, the method of growing
the fungi under study in pure culture on various types of culture media
and under various external conditions. In this way he was enabled to
study the developmental stages of many fungi and to learn much of their
physiology as well. His chief series of contributions began in 1872 and
the last volume appeared in 1912. One of de Bary's early students and
collaborators was the Russian, M. S. Woronin (1838-1903), who returned
to Russia after several years of association with his great teacher, there
becoming in his turn the center of a group of very able mycologists and
plant pathologists. The life history studies thus stimulated by de Bary
and his students have continued up to the present with so great a number
of investigators that mention of their names, even, must be omitted.

The introduction of the recently developed cytological methods to
the study of fungus life histories began on a large scale with the investi-
gations by P. A. Dangeard (1894) in France, and of R. A. Harper (1896-
1897).. Their first publications along these lines appeared between 1894
and 1897 and were soon followed by the contributions of a host of other
eager students in all parts of the w^orld. The correction or confirmation
of previously held ideas — particularly with reference to the nature of the
sexual act in fungi — thus made possible, has proved to be of the utmost
value in assisting the determination of relationships among the fungi.

In 1904 and succeeding years A. F. Blakeslee made known the occur-
rence of those sexual phenomena in the Mucorales to which he gave the
names heterothallism and homothallism. The study of these types of
sexual reaction has been extended to other groups of fungi: In the
Basidiomyceteae by Mile. Bensaude (1918), Hans Kniep (1913-1917),
Miss Mounce (1922), Vandendries (1923), Hanna (1925), and many
others from 1915 to the present; in the Ustilaginales by Bauch (1922),
Hanna (1929), Kniep, Stakman (1927), and others in the last twenty
years; in the Uredinales by Craigie (1927, 1931), Andrus (1931), Miss
Allen (1930), etc., since 1927; in the Ascomyceteae by B. O. Dodge
(1927), Ames (1932), Drayton (1932), and several others, mainly since
1927. Dodge (1928), Lindegren (1933), and others have made intensive
studies on the genetics of fungi in the last fifteen or more years, particu-

RULES FOR NOMENCLATURE 15

larly in various species of Neurospora, while many others have studied
the smuts from the genetic standpoint. Hybrids have been produced in
both these groups as well as in the rusts and their structure and their
behavior studied. So much has been published in these fields in the last
fifteen to twenty years that the names of the investigators cannot be
listed here. Of those mentioned above only a few of their earlier papers
are noted, though in many cases they produced many later contributions.

The present day finds systematic mycologists active all over the
world. Life histories are being studied in all groups. The sexual relations
are being scrutinized from the lowest to the highest fungi and genetic
studies are revealing results somewhat parallel, but on a smaller scale as
yet, to those attained by the study of Zea mays and Drosophila. Now, as
never before, a knowledge of the fungi themselves is necessary.

The discovery of the production of antibiotic substances by various
fungi has encouraged intense research in that field, the results of which
have only recently become of great importance in the medical field.

Rules for Nomenclature

The rules for botanical nomenclature, especially as applicable to fungi,
have been given an extended discussion by Bisby (1945). Only the more
fundamental points will be taken up here. These rules have been formu-
lated and added to and modified at a series of International Botanical
Congresses in 1867, 1905, 1910, 1930, and 1935. The aims are expressed in
Articles 2 and 4 of the Rules, from which the following sentences are
quoted :

The object of the rules is to put the nomenclature of the past into order and
to provide for that of the future. They are always retroactive: names and forms
of nomenclature contrary to a rule {illegitimale names or forms) cannot be main-
tained. . . . The essential points in nomenclature are: (1) to aim at fixity of
names; (2) to avoid or to reject the use of forms and names which may cause
error or ambiguity or throw science into confusion. . . . Next in importance
is the avoidance of all useless creation of names.

Art. 7. Scientific names of all groups are usually taken from Latin or Greek.
When taken from any language other than Latin, or formed in an arbitrary
manner, they are treated as if they were Latin. Latin terminations should be
used so far as possible for new names.

The earliest name properly applied to a plant shall be retained, pro-
vided it is a binomial, i.e., consists of the generic name and specific
epithet. Since, however, it is impractical to go back to the classical
authors or those of the Middle Ages it has been agreed that names applied
before the appearance of Linnaeus' "Species Plantarum" (1753) shall
not be considered, nor those subsequent to that date which do not use
the binomial nomenclature. Because many of the fungi were not well
known to Linnaeus the basic dates for the earliest authoritative names

16 INTRODUCTION

have been assigned variously in accordance with the groups of fungi con-
cerned. Thus for the Mycetozoa and Lichens the date remains 1753; for
the Uredinales, Ustilaginales, and Gasteromycetes it is 1801, based on
Persoon's "Synopsis Methodica Fungorum"; and for all other fungi
1821-1832, based on the appearance of the various volumes of Fries'
"Systema Mycologicum." So the beginning date for the Hymenomycetes
is based upon Volume I of this work, part of which is said to have appeared
late in 1820 and the remainder in 1821. Any names given by later authors
are valid only if the species concerned were not included in that volume.
The first section of Volume II appeared in 1822 and included, and there-
fore is authoritative for their nomenclature, the Discomycetes and many
of the larger Heterobasidiae, and the sclerotioid Fungi Imperfecti. Part II
of this volume appeared in 1823 and contained some of the Gastero-
mycetes, whose basic date is Persoon's work in 1801, and the Pyreno-
mycetes (and Sphaeropsidales). Volume III, Section I (1829) contains
(in addition to the remainder of the Gasteromycetes and the Mycetozoa)
the Erysiphales. For this last group it is the basic work. Section II (1832)
consists mainly of the Fungi Imperfecti, but also Hypodermii (i.e.,
Uredinales and Ustilaginales) for which the work of Persoon is basic.

A generic name is always a noun in the singular number. It is always
written with an initial capital. The specific epithet is mostly an adjective
in Latin form, which must agree in gender with the generic name (e.g.,
Lepiota procera), or it may be the genitive case of some noun (e.g.,
Mycosphaerella fragariae), or it may be a noun in the nominative case
(e.g., Xylaria hypoxylon or Fomes pinicola). Mostly the specific epithet is
written without an initial capital, except where it is a noun in the nomi-
native case that was an old generic name, or where it is based upon a
personal name or a generic name. INIany authors prefer to decapitalize
all specific epithets, even when based upon personal or generic names.
The author attempts to follow this procedure in this book except where
quoting directly from another author.

Valid publication of a hitherto undescribed plant consists of the
assignment of a name and the description of the organism, in proper
manner. After 1935 all descriptions to be valid must be in Latin aside
from any descriptive text in any other language. The description of a
species must be based upon a definite collection, specimen, or culture,
which is designated as the "type specimen." This should be preserved
and the place where it is deposited should be indicated. A genus must be
based upon some particular species, the "type species." A family must
be based upon a "type genus" and an order upon a "type family."
Ordinarily the familial and ordinal names are based upon the stem of the
name of the type genus, with the appropriate endings, -aceae or -ales,
respectively. If a genus, family, or order is divided into two or more parts

RULKS FOR NOMENCLATURE 17

the original name must be retained for the part in which the type species,
genus, or family remains.

When a species is transferred to another genus than that under which
it was first described, or when for any other valid reason the generic
name is changed, the specific epithet must be retained, subject to change
of gender in case it is an adjective, if the new genus name is of different
gender. Exceptions are as follows: If the new genus name is the same as
the specific epithet or if in the new genus a similar combination already
exists (e.g., when Leontodon taraxacum was split off from the genus
Leontodon and placed in the new genus Taraxacum, the epithet had to be
replaced; Uredo sorghi (1897) was transferred to the genus Puccinia, but
as there was already a Puccinia sorghi (1832), another epithet had to be
chosen). In this case the oldest available epithet of a synonym must be
taken or in lack of such a name a new epithet must be provided by the
author making the transfer.

In describing a species de novo the describer's (abbreviated) name
follows the specific epithet, but if an author changes the genus to which
the species must be assigned the name of the person who first gave the
epithet must be placed in parentheses followed by that of the author of
the new combination. For example, Bessey and Thompson described a
new species of fungus under the name Genea cuhispora Bessey & Thomp-
son. When Miss Gilkey determined that this belonged to the genus
Hydnotria, the name became Hydnotria cuhispora (Bessey & Thompson)
Gilkey. If the epithet was given to a plant accompanied by an adeqviate
description before the basic date for that group of plants it is not neces-
sary to refer to the author of that epithet but only to the person who
first used it on or after the basic date. However, this later author could
indicate that he took this epithet on the authority of the earlier author
by placing, as authority for the combination, the abbreviated name of the
earlier author followed by "ex" and the name of the later author. Thus
Persoon in 1801 described a puffball under the name Lycoperdon giganteum,
ascribing this combination to Batsch who published the name before
1790. It is permissible simply to give "Pers." as the authority, but it is
preferable to write "Batsch ex Pers."

To avoid confusion a genus name once applied to any plant may not
ever thereafter be used for another genus, even though the name is
invalid for the first genus. Thus, unless an exception is made at some
future Botanical Congress, the generic name Empusa, given about 100
years ago to an orchid, is not available for a genus of the Entomophtho-
rales. A specific epithet once applied in a given genus may never be used
for another species in the same genus.

In fungi with several stages of development to which different names
have been given, the species epithet that is to be retained is the one

18 INTRODUCTION

applied to the ''perfect" stage of the fungus, regardless of the fact that
names may have been given earlier to some of the other stages. Thus
Persoon (1801) recognized the following three species of rusts which are
now known to be different stages of the same rust: Aecidium herheridis
for the aecial stage occurring on Berberis vulgaris L.; Uredo linearis for
the uredial stage on small grain; and Puccinia graminis, for the telial
stage on the same host. Only the last name may be used for the species,
for it is the name of the "perfect" stage. The name Lycoperdori poculiforme
Jacq., given 10 or 12 years earher may not be used for two reasons: it
was given before the basic date 1801 and was applied to the aecial stage
of the rust.

The fungi, including the Mycetozoa, which probably do not belong
at all in this group of organisms, may be divided as follows :

Key to the Major Groups of Fungi

Vegetative stage permanently naked and either flagellate or amoeboid. Encysted
spores produced to serve as organs of distribution or to carry the organisms
over unfavorable conditions.

Subclass Mycetozoa (Chap. 2)
Vegetative stage for all or part of its course of development with cell walls.

True Fungi
Plant one-celled and then giving rise to planocytes, or producing a coenocytic
mycelium. Sexual reproduction resulting in the formation of a zygospore
or an oospore. Class Phycomyceteae (Chaps. 3-7)

Plant one-celled, not producing planocytes, or producing a cellular mycelium.
Sexual reproduction resulting in the formation of a spore fruit.

Higher Fungi (Phylum Carpomyceteae, Chaps. 8-16)
Ultimate reproductive spores of the spore fruit produced internally in an

ascus. Class Ascomyceteae (Chaps. 9-11)

Ultimate reproductive spores of the spore fruit produced externally upon
a basidium or its equivalent.

Class Basidiomyceteae (Chaps. 12-15)
Sexual reproduction stage not known.

Class Fungi Imperfect! (Chap, 16)

At the close of each chapter is a key to the families and more impor-
tant genera of the orders considered there.

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thallism in the Ascomycete Neurospora, /. Agr. Research, 35(4):289-305.

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36(1):1-14. P/s. 1-4. 1928.
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296. 1926.

20 INTROBUCTION

Drayton, F. L. : The sexual function of the microconidia in certain Discomycetes,

Mycologia, 24(3) :345-348. 1932.
Fries, Elias Magnus: Systema mycologicum, sistens fungorum ordines, genera

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Figs. 1-6, 1933.
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: Mycologia Europaea seu completa omnium fungorum in variis Europaeae

regionibus detectorum enumeratio, methodo naturali disposita, descriptione

succincta, synonymia selecta et observationibus criticis additis, 3 sections,

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LITERATURE CITED 21

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Chapter 18 of this book.)
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plantes. Tome I, text, xviii + 562 pp. + index, 18 pp. Tomes II and III,

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Wiss. Wien, Math, naturw. Klasse, Abt. I, 130(1) :3-20. 1921.

MYCETOZOA AND RELATED ORGANISMS

UNDER this title are brought together several groups of organisms
which are probably interrelated but whose relationship to the true
fungi is very doubtful. As mentioned in the preceding chapter the author
follows de Bary (1887) in considering them to be more nearly related to
members of the Animal Kingdom. They are discussed in this chapter
because, for several hundred years before their hfe histories were known,
botanists looked upon their mature fruiting bodies as fungi and gave
them names, many of which are still maintained. Because they are more
usually considered, even at this date, as fungi, they are given a place in
this text.

These organisms all agree in the fact that they are naked during all
stages of development except the culminating spore stage. Mostly for a
portion or for the whole of this naked stage they are more or less amoeboid
and ingest particles of food, rejecting the portions remaining after diges-
tion is accomplished. At germination of the spore the escaping naked
protoplast may be a simple uninucleate amoeboid cell, or myxamoeba,
or it may be provided with one or two anterior flagella which eventually
are retracted, leaving the cell a myxamoeba. To a greater or less degree
both the planocytes and the myxamoebae are capable of multiplication
by fission. Usually after a while the myxamoebae, through the division
of the protoplast, become multinuclear plasmodia, or these may arise
by the fusion of separate myxamoebae as well as by growth and nuclear
division. In the better known groups there is usually a sexual stage, and
eventually, just before spore formation, a meiotic division of the nuclei
of the Plasmodium so that the encysted spores contain haploid nuclei.

These organisms are usually classified by the zoologists as belonging
to Phylum Protozoa, Class Sarcodina (or Rhizopoda). This classification
is quite different from that mostly used by botanists. In the following
pages an attempt will be made to reconcile these two viewpoints, yet
maintaining so far as possible the nomenclature with which botanists, in
particular mycologists, are more or less familiar. Four orders will be

ORDER MYXOGASTRALES OR SLIME MOLDS

considered. The ordinal limits as here treated are far wider than those
found in the monographs of the Slime Molds by Lister (1925), or Macbride
and Martin (1934), and others. These four orders may be included in the
Subclass Mycetozoa of Class Sarcodina, Phylum Protozoa. The four
orders are Myxogast rales, the true Slime IMolds, Acrasiales, Plasmo-
diophorales and Labyrinthulales. There are other organisms probably
closely related to these that have been studied by zoologists but which
have usually been neglected by botanists and are not considered here.

Order Myxogastrales or Slime Molds. Often called Myxomycetes or
Myxogastres, the Myxogastrales or Slime Molds compose the most
numerous group of this subclass. They are terrestial organisms or inhabi-
tants of manure, decaying wood, decaying fungi, etc. Their spores are
produced upon or within aerial sporangia and are more or less dependent
upon wind for their distribution. In general the life history (with some
modifications) is as follows: The spore, upon absorbing water, cracks open
its cell wall and escapes as a single (in some species by division of the
nucleus before germination, two) naked uninucleate swarm spore or
planocyte, rounded posteriorly and tapering to the anterior end from
which arises a single flagellum or in many cases two flagella. De Bary
(1859) described and figured occasional biflagellate zoospores of Fuligo

Fig. 1. Myxogastrales. (A-H) Physarum polycephalum Schw, (A) Spore. (B) Ger-
minating spore. (C, D) Swarm spores. (E) Uniting swarm spores. (F, G) Amoeboid
zygotes. (H) Portion of Plasmodium. (I-J) Physarella oblonga (B. & C.) Morg. (I)
Uniflagellate swarm spore. (J) Biflagellate swarm spore. (All figures much mag-
nified.) (A-H, after Howard: Am. J. Botany, 18(2). I-J, after Sinoto and Yuasa: The
Botanical Magazine [Tokyo], 48(574) :722.)

24 MYCETOZOA AND RELATED ORGANISMS

septica (L.) Gmel. and Trichia varia Pers., as well as the normal uni-
flagellate zoospores. Gilbert (1927) reports that in one collection of
Stemonitis fusca Roth he studied about one-fourth of the swarm spores
bore two flagella and three-fourths bore only one. Sinoto and Yuasa (1934)
and Yuasa (1935) showed that whether one or two flagella are produced
the cell is uninucleate and possesses two blepharoplasts usually connected
by a slender rod and joined to the nucleus by means of a single rhizoplast.
In the case of two flagella each blepharoplast bears one, in the case of one
flagellum one of the blepharoplasts remains without a flagellum, Ellison
(1945) confirmed Yuasa's findings and demonstrated that the flagella
whether single or two in number were all of the whiplash type, i.e., have
a firmer, outer tubular portion beyond which the interior portion projects
more or less or may be gathered into a ball. No flagellum of the tinsel
type is produced, i.e., with numerous minute cilium-like lateral out-
growths. Elhott (1948) made intensive studies of the germinating spores
of 1 1 species of this order and found that biflagellate swarm spores were
present in all these species, up to nearly 100 per cent in Fuligo septica (L.)
Gmel. In all cases except in the genus Stemonitis one flagellum is long and
the other very short and often somewhat recurved. Unless seen in profile
the shorter flagellum is difficult of demonstration. This probably accounts
for the earlier belief that these organisms possess but one, anteriorly
directed, flagellum. (Fig. 1.)

These motile swarm cells ingest food in the manner of Amoeba, leaving
behind the undigested debris. This food is in many cases bacterial cells
but various other objects of an organic nature may be consumed. The
swarm cells may divide by fission several times and then change their
form, retracting their flagella and becoming more rounded, with usually
more conspicuous pseudopodia. These myxamoebae usually enlarge and
divide several times. Eventually they begin to unite by twos, with
nuclear fusion, to form zygotes. In a number of genera the myxamoebal
stage does not occur and the sexual fusion takes place between two swarm
cells {Reticularia, according to Wilson and Cadman, 1928; Didymium
difforme (Pers.) Duby, according to Miss Cayley, 1929; Physarum
polycephalum Schw., according to Howard, 1931; and others according
to Abe, 1934). Skupienski (1928) claims that the nuclear fusion does not
occur in Didymium difforme until just before spore formation. In general
the zygote formed by the union of two myxamoebae or of two swarm
cells is nonflagellate but continues its existence as a naked amoeboid cell
which ingests its food and grows in size, with accompanying mitotic
division of the nucleus. This multinucleate structure is cahed a Plas-
modium. Zygotes or small plusmodia may fuse with other zygotes and
Plasmodia so that growth is both internal and by accretion, Plasmodia
may ingest and feed upon swarm spores and myxamoebae. The plas-

ORDER MYXOGASTRALES OR SLIME MOLDS

A B

Fig. 2. Myxogastrales. Didymium squamulosum (A. & S.) Fr. (A) Plasmodium. (B)
(B) Sporangium. (Courtesy, D. M, Cayley.)

modium creeps through the soil or rotten wood or decaying vegetable
matter, fruiting bodies of fungi, etc., digesting the food suitable for it
and increasing in size and in the number of contained nuclei. Howard and
Currie (1932) showed that the plasmodia of various species of slime molds
destroy the mycehum and sporophores of many types of Hymeno-
myceteae with great rapidity. On the other hand some species can be
grown very successfully upon various standard culture media. Eventually
it emerges as a fine or coarse network upon the surface of the substratum.
It may be lobed and exhibit a continual creeping motion in various direc-
tions. In diameter it may vary from a few millimeters up to 15 or 20
centimeters, and in color from white to yellow, orange, red, brown,
violet, and other colors. Camp (1937) showed that it contains true con-
tractile vacuoles. Food is obtained by invagination, the lining of the food
vacuole being a portion of the original external surface of the Plasmodium.
Eventually the Plasmodium heaps itself up somewhat, on the exterior
of its substratum, or even creeps up adjacent objects, and there undergoes
the changes which lead to the production of the fructifications. These
may be separate from one another or may be crowded together into a
compound structure. In the Suborder Endosporeae there appears ex-
ternally, a noncellular peridium secreted by the Plasmodium. This may
be of various thicknesses according to the species, and may or may not
be encrusted with lime. Within the fructification are secreted numerous
noncellular threads and beams which form a sort of framework. This is
the capillitium. Its structure and arrangement are of great value in
classifying this difficult group of organisms. Between the threads of the
capillitium, usually following simultaneous meiotic divisions of all nuclei
of the Plasmodium, the protoplasm rounds up into innumerable small
uninucleate cells which secrete cell walls. In Physarum polycephalum it

26 MYCETOZOA AND RELATED ORGANISMS

has been shown by Howard (1931) that the cell walls are secreted before
the spores round up, so that at first a continuous mass of polyhedral cells
is produced. The cells then begin to round up and the cell walls split,
thus forming the spores. The walls of some species are said to contain
cellulose, but some investigators deny this. C. van Wissehngh (1898)
claimed the presence of cellulose in Didymium squamulosum (A. & S.) Fr.,
but denied its presence in Fuligo septica (L.) Gmel. F. von Wettstein (1921)
studied the composition of the spore wall in seven other genera and could
not demonstrate the presence of cellulose except, doubtfully, in Stemonitis
and Reticularia. Neither author found chitin in any of these. The spores
are most often violet, purple, or brown. (Fig. 2.)

By the rupture or dissolution of the peridium the spores are permitted
to escape, the capillitial threads preventing all the spores from escaping
at once. The fructifications may be sessile or stalked, round or elongated,
scattered or crowded, almost microscopic or, in the case of some of the
compound fructifications, up to 10 cm. in length and 4-5 cm. in width
and thickness.

In the genus Ceratiomyxa, the only genus of the Suborder Exosporeae,
the spores are extruded externally from the fructification instead of being
produced internally. The studies of H. C. Gilbert (1935) seem to indicate
that this genus may be looked upon as an extreme modification of the
condition in Suborder Endosporeae. As in that suborder the sexual fusion
occurs by the union of two or more swarm spores followed after some time
by the union of pairs of nuclei. The zygotes then enter decaying wood
within which they feed, perhaps upon the mycelium of the fungi causing
the decay. Eventually the plasmodium creeps out of the wood in which
it has been growing and secretes a massive central core of mucilaginous
nature, on the outside of which the protoplasm creeps as a thin, some-
what reticulate, sheet. In this sheet a further division of the nuclei is
followed by a cleavage of the protoplasmic layer into uninucleate naked
cells, flattened at the contact surfaces. Each of these secretes a stalk of
material similar to that of the main central core, on whose tip the thin-
walled spore takes its position. Within this the nucleus divides meiotically
to form four haploid nuclei, the spore falling off before or after this stage.
It cleaves into four cells in each of which mitotic nuclear division and
cleavage occur, so that 8 naked flagellate cells arise from each spore.
Gilbert homologizes the stalk upon which the spore sits and the spore
with the stalk and sporangium respectively in the Endosporeae. Olive
(1907) disagrees with Gilbert, claiming that the nuclear fusions do not
occur until just before the formation of the stalks for the external spores.
Jahn (1936) agrees with Gilbert that sexual union occurs by the fusion of
the swarm cells and their nuclei but locates the meiotic division of the
nuclei, as Olive did, in the plasmodium before the stalked spores are pro-

OKDER MYXOGASTRALES OR SLIME MOLDS

Fig. 3. Myxogastrales. Ceratiomyxa fruticulosa (Muell.) Macbr. (A) Habit sketch.
(B) Development of young fruiting body from Plasmodium. (C) Portion of mature
sporophore, showing production of external spores. (After Famintzin and Woronin.
From Engler and Prantl: Die natlirlichen Pflanzenfamilien, Leipzig, W. Engelmann.)

duced. In the Endosporeae, Skupienski (1928) claims that the nuclear
fusion does not occur in Didymium difforme until the time for spore forma-
tion is at hand. Then the hundreds of nuclei in the Plasmodium unite
simultaneously in pairs, this being followed closely by the meiotic divi-
sions which give rise to the nuclei of the spores. His interpretation of the
nuclear cycle in Didymium thus agrees with Olive's for Ceratiomyxa.

The Suborder Exosporeae contains but one genus, Ceratiomyxa, with
two or three species. Their fructifications appear as simple or branched
columns or even as poroid structures 2-6 mm. tall, light in color, growing
on decayed logs, stumps, etc. (Fig. 3.)

The Suborder Endosporeae is recognized by Macbride and Martin
(1934) as possessing 59 genera and 380 species. The classification of the
numerous genera and species is based upon the type of the peridium
and the presence or absence of lime in it and in the elements of the
capillitium, the microscopic structure of the latter, the formation of
simple or compound fructifications and their size, shape, and color.
Further characters are the color, shape, size, and markings of the spores
as well as the characters of the Plasmodium,

MYCETOZOA AND BELATED ORGANISMS

Fig. 4. Myxogastrales. (A, B) Fuligo septica (L.) Gmel. (A) Habit drawing. (B)
Capillitium and spores. (C, D) Physarum nutans Pers. (C) Sporangia. (D) Capillitium.
(E) Dictydium cancellatum (Batsch) Macbr., showing persistent ribs of sporangium
wall. (After Lister: A Monograph of the Mycetozoa, London, The British Museum.)

Mention should be made of Stemoniiis, with purple or rust-brown
spores, clustered, stalked fruit bodies with the stalk extending upward
as an axial strand (columella) from which branch off the capillitial threads
which are combined into a loose network that is surrounded by an eva-
nescent peridium. Physarum has clustered, separate or compacted, sessile
or stalked, lime-encrusted sporangia whose capillitial threads are ex-
panded here and there and filled with lime granules. Fuligo has a similar
capillitium but the spore fruits are united into a single large convolute
"aethalium." This is one of the largest of the Slime Molds. In Dictydium
the stalked sporangium has no internal capillitium but when the peridium
disappears it leaves numerous longitudinal ribs that run from base to
apex like the Hues of the meridians on the globe. Arcyria and Trichia have
sessile or stalked fruits. The capillitium lacks a columella and consists of
a tangle of tubular threads with characteristic thickenings such as spines,
warts, rings, etc. (Fig. 4.)

The fructifications of Slime Molds may be found on rotten logs or
stumps, on sawdust, leaves, beams in moist cellars, and frequently on
blades of grass or other vegetation in lawns. Very dry habitats do not
favor their occurrence. In moist weather the plasmodia may be found
creeping about on the surface of, or emerging from, the various substrata
within which they develop. It is possible to grow some species in pure
cultures, from the spores to the maturity of the fructifications.

ORDERS ACRASIALES AND LABYRINTHULALES

Orders Acrasiales and Labyrinthulales. Sometimes associated with
the Subclass Mycetozoa are the two orders Acrasiales and Labyrinthulales.
Whether they should be included in this subclass or in distinct subclasses
the author will not seek to decide. In the first of these two orders the
swarm spores do not possess flagella although they are amoeboid upon
emerging from the spore wall. They consist of saprophytic or parasitic
organisms occurring on dung, decaying wood, leaf mold or other organic
matter. The spores upon germination give rise to naked amoeboid cells
(myxamoebae) with or without conspicuous pseudopodia and containing
a single nucleus and one or more food vacuoles. Within these vacuoles are
digested the bacteria and other bits of organic matter that serve as food
for the organism. Raper (1937) has shown that the myxamoebae of
Dictyostelium discoideum Raper are strictly parasitic upon bacteria, many
kinds of which may serve as their food. There is no indication of a
symbiotic relation between the bacteria and the myxamoebae. The latter
enlarge and divide several times and in D. mucoroides Bref., according to
Skupienski (1920), then unite by twos. The resulting zygotes seem now
to be mutually attracted to one another and draw together into heaps of
naked cells that maintain their individuality. These heaps of separate
cells are called pseudoplasmodia in contrast to the true plasmodia that
are found in the Myxogastrales. At this stage pressure or, in some cases,
exposure to bright light will cause the pseudoplasmodium to separate into
its individual cells which reassemble again elsewhere. Skupienski claims
that in Dictyostelium the cells eventually unite into a true Plasmodium
within which the nuclei undergo two more divisions (probably meiotic).

Fig. 5. Acrasiales. (A) Pseudoplasmodium of Polysphondtjlium violaceum Bref.
(B, C) Dictyostelium mucoroides Bref. (B) Stalk and terminal ball of spores. (C)
Details of cells of stalk. (After Olive: Proc. Boston Soc. Natural History, 30:451-513.)

30 MYCETOZOA AND BELATED OKGANISMS

In this genus the plasmodium heaps itself up, at first, into a conical
structure in the basal part of which cellulose walls are produced, sepa-
rating it into the more or less hexagonal cells of the stalk. In the upper,
head-like part it breaks into separate, rounded cells, which also develop
cellulose walls and become the spores. These are embedded in a slimy
drop. Sexual reproduction and the eventual formation of a true Plas-
modium have not been reported but possibly occur in the other genera of
the order. The myxamoebae of Copromyxa and Guttulina lack con-
spicuous pseudopodia. The fruiting bodies of the former are sessile, of
the latter short-stalked. Dichjostelium, Acrasis, and Polysphondylium pro-
duce myxamoebae with well-developed pseudopodia. All produce stalked
fruiting bodies (branched in the last named) with ovoid or spherical
heads of spores. Olive (1902) gives a monograph of this order, describing
all known species. (Fig. 5.)

The Acrasiales differ from the Myxogastrales in the absence of a
flagellate swarm-spore stage preceding the formation of the myxamoebae,
and in the fact that the naked cells remain distinct for a long time in the
pseudoplasmodium, forming a true plasmodium only a short time before
the fruiting body is produced, if at all. Furthermore no capillitium is
formed nor a peridium, although the slime that lies between and around
the spores may represent these structures.

The Labyrinthulales include parasitic forms attacking algae and other
aquatic plants in both marine and fresh-water habitats, and possibly
one genus of dung-inhabiting saprophytes. There are several species in
the genus Lahyrinthula, of which L. macrocystis Cien. was shown by
Renn (1935) to be the cause of the destruction of most of the eel grass
(Zostera marina L.) of the shallow waters of the North Atlantic Ocean
in recent years, both on European and on North American shores. This
genus, according to Young (1943) consists of naked, spindle-shaped cells
with a single nucleus and a vacuole which may contract at intervals.
These cells divide transversely or obliquely and form a rope-hke mass.
Those near the tips of this mass send out, apically, thin colorless fila-
ments, one to each cell, eight to ten times its length. These filaments fuse
to form a net-like track along which the cells glide— externally to the
track, not in it as in a tube as Valkanov (1929) claimed. Eventually the
cells may assemble into a pseudoplasmodial mass embedded in a gelati-
nous matrix. Some of these cells may encyst or encystment of individual
cells may occur without the formation of a pseudoplasmodium. The
encysted cells apparently give rise to four naked cells which penetrate
through the cell walls of the host plant and in their turn become spindle-
shaped and start the development of new "net-plasmodia." Dangeard
(1932) suggested that possibly there is a sexual stage somewhere in the
life cycle but did not demonstrate where it occurred. The naked cells

OKDER PLASMODIOPHORALES

emerging from the cysts are nonflagellate in Lahyrinthula and anteriorly
uniflagellate in the possibly related Lahyrinthomyxa. (Fig. 6.)

Order Plasmodiophorales. This order to which eight or more genera
have been ascribed is not definite as to its hmits or relationships. The
type genus Plasmodiophora differs in so many points from many of the
genera assigned to the order that it may be necessary to limit the order

Fig. 6. Labyrinthulales. Lahyrinthula 7nacro-
cystis Cien. (A) "Net Plasmodium," showing
clumps of cells on the "tracks" and cells sending
out threads which are the beginnings of new tracks.
(B) Highly magnified single cell lying on track.
(Courtesy, Young: Am. J. Botany, 30(8) :586-593.)

to one genus and to place the others in another order, perhaps not at all
closely related. The type species P. hrassicae Wor., causing swellings and
malformations of the roots of Brassicaceae (Cruciferae), will be discussed
first and then some of the others with remarks as to their differences and
possible relationship. Woroni7ia, placed here by Sparrow (1943), is re-
turned to a position in the Order Lagenidiales because of its cellulose
walls and the possession of flagella of two types. Possibly other genera
may have to be relegated to that position also. In the following account
many points still are disputed or assumed without adequate confirmation.
The spores of Plasmodiophora hrassicae possess a dark wall which,
according to van Wissenlingh (1898), contains chitin but not cellulose.
Upon germination, usually a single zoospore emerges, although Honig
(1931) and Rochlin (1933) claim that only a nonflagellate amoeba is

MYCETOZOA AND RELATED ORGANISMS

Fig. 7. Plasmodiophorales. Life cycle of Plasmodiophora brassicae Wor. (After Cook
and Schwartz: Trans. Roy. Soc. London, B, 218:283-314.)

formed. The zoospore possesses two anterior flagella, as demonstrated by
Ledingham (1934), the longer one pointing forward, the shorter directed
almost at right angles. Ellison (1945) showed that both of these flagella
are of the truncated whiplash type, neither showing any tinsel structures.
This is a character that indicates that Plasmodiophora is not related to
the Olpidiopsidaceae, whose anteriorly biflagellate zoospores possess one
flagellum of each of these two types. The zoospore is more or less amoe-
boid. When it comes into contact with a root hair or epidermal cell of the
root of a suitable host, the flagella disappear and a hole is dissolved in the
host cell wall through which the amoeba enters, then this hole is closed,
presumably by host action. Cook and Schwartz (1930) showed that within
the root hair this amoeba enlarges and mitotic divisions of the nucleus
occur until a small Plasmodium is formed, containing from a few up to
one hundred nuclei according to Fedorintschik (1935). This Plasmodium
cleaves into uninucleate cells around each of which a thin wall is formed.
Whether this wall contains cellulose or chitin has not been determined.
They give rise to structures called sporangia or gametangia. The nucleus
divides mitotically two or three times and thus there are formed four to
eight uninucleate, anteriorly flagellate swarm cells, smaller than those
that emerge from the resting spores. The number and character of the

ORDER PLASMODIOPHORALES

Fig. 8. Plasmodiophorales. (A) Spore balls of Spongospora subterranea (Wall.)
Lagerheim. (B-E) Plasmodiophora brassicae Wor. (B) Biflagellate zoospores. (C)
Amoebae in root hairs. (D) Young plasmodia in root cortex. (E) Host cell filled with
spores. (F) Sorosphaera veronicae Schroet. (A, after Osborn: Ann. Botany, 25(98) :327-
341. B, after Ledingham: Nature, 133(3362) :534. C-E, after Chupp: Cornell Univ.
Agr. Sta. Bull, 387 :421-452. F, after Palm and Burk: Arch. Protistenk., 79(3) :263-276.)

flagella have not been determined in this species. The foregoing authors
beheve that the swarm cells unite by twos and that the resulting amoeboid
zygotes are the origins of the large plasmodia which give rise to the resting
spores. The zygotes and young plasmodia arising from them may possibly
unite to form larger plasmodia. These are slowly amoeboid and at least
in their younger stages seem to be able to pass from cell to cell of the host.
The young plasmodia apparently may undergo division. As growth of the
parasite progresses, the host cells multiply hyperplastically and the
infected cells undergo hypertrophy. Eventually the cell contents of the
invaded cells are almost completely exhausted and the cell is practically
filled by the Plasmodium. The nuclei of the latter then undergo two
rapidly succeeding divisions which are believed to be meiotic, although
cytologic studies have not definitely proved this. Then the protoplasm
rounds up into uninucleate spores around each of which a dark chitin
wall is secreted. There is no enclosing membrane around the mass of
spores which lie free in the host cell. Upon decay of the root the spores
are set free. In the laboratory they are brought to germination only with
difficulty. The wall cracks open and the zoospore emerges with the two
flagella in advance of the cell body. At this stage, the author has observed
the two flagella on the living zoospore before it has escaped from the
spore wall, so that there is no doubt as to the correctness of Ledingham's

34 MYCETOZOA AND RELATED ORGANISMS

report. Webb (1949) reports the occurrence of sporangia of this fungus in
root hairs of Rumex sp. and Holcus lanatus L., growing in infested soil.
These roots were carefully washed and planted in sterile soil. Seedlings of
Brussels sprouts (Brassica sp.) planted in the pots along with these washed
roots developed the disease. In spite of numerous extended studies on this
parasite, many contradictory reports have been published. Thus P. M
Jones (1928) reports eight swarm spores emerging from the resting spore
and their fusion by twos so that infection of the root hairs is by means of
amoeboid zygotes. Fedorintschik believes that the nuclei of the zoospor-
angia formed in the root hairs are diploid and that two meiotic divisions of
the nucleus occur as the swarm cells are formed, these last fusing by twos,
as reported by Cook and Schwartz. Karling (1942) doubts the sexual
nature of these swarm cells and suggests that they are only secondary
zoospores, such as are reported in some other so-called Plasmodiophorales.
Several other species have been ascribed to the genus Plasmodiophora but
their life histories have not been studied and it is not at all certain that
they belong here. (Figs. 7, 8.)

The other genera which Karling admits to this order, which has but
one family, are as follows:

Tetramyxa: causing pronounced hypertrophy of host tissues, developing spores
in twos or fours, and with sporangia and sexual stage unknown, and
zoospores not seen.
Two or possibly three species in stems or roots of seed-plants.

Odomyxa: spores usually develop in eights, sporangia numerous, zoospores
anteriorly biflagellate, one flagellum directed forward, a longer one pos-
teriorly, when swimming. No sexual stage known. Cell walls not of cellu-
lose. Causes gall-like enlargements in Achlya glomerata Coker.

Sorosphaera: spores compacted into a hollow sphere, zoosporangia are formed
(Ledingham, 1939) and their zoospores anteriorly biflagellate and hetero-
cont. Spore walls do not contain cellulose.
Two species: S. veronicae Schroeter forms galls in the stems of Veronica;
S. radicalis Cook & Schwartz forms galls in the roots of grasses. In the
former zoosporangia are unknown, in the latter thin-walled zoosporangia
are formed. Sexual reproduction not demonstrated.

Sorodiscus: spores formed in a disk-shaped sorus usually in two layers, much
like a flattened spherical sorus of Sorosphaera. Zoosporangia and character
and number of flagella unknown. Sexual reproduction not observed.
Two species : one in stems of Callitriche and one in Chara.

Spongospora: spores in a hollow sphere with several openings, zoosporangia are
formed according to L(Mli,igham (1935), zoospores anteriorly biflagellate
and heterocont, similai' in size whether from sporangia or from resting
spores. Sexual fusion of myxamoebae reported by Cook (1933).
Best known species is S. subtcrranea (Wall.) Lagerheim causing the powdery
scab of the tubers of potato {Solanum tuberosum L.). It also attacks the
stems and roots of this host and of related plants.

Ligniera: spores in irregular clusters, usually not causing hypertrophy of
tissues except of root hairs in one species. This, according to Palm and

ORDER PLASMODIOPHORALES 35

Burk (1933), is a very doubtful genus. Zoosporangia are known but
further study is needed to determine whether the zoospores are truly
uniflagellate anteriorly as reported or biflagellate.
Polymyxa: spores in irregular clusters in the roots of grasses, zoosporangia with
long necks are abundant, zoospores from the resting spores as well as
those from the zoosporangia are anteriorly biflagellate and heterocont
(Ledingham, 1939). Sexual reproduction has not been observed. Hyper-
trophy of the host cells is not brought about.

Several other genera have been ascribed to this family but Karling
(1942) is doubtful as to the correctness of their assignment here {Sorol-
pidium, Anisomyxa, Trematophlyctis, Sporomyxa, Peltomyces, Cystospora).

The relationship of the four orders — -Myxogastrales, Acrasiales,
Labyrinthulales, and Plasmodiophorales — described in the foregoing, is
not admitted by all. Sparrow (1943) places the last named order in his
group Biflagellatae, in which he also includes the Saprolegniales, Lepto-
mitales, Lagenidiales, and Peronosporales. The author admits the simi-
larity of Octomyxa and Polymyxa to Woronina, which he includes in the
Lagenidiales. It is possible that with further study of these and some
other genera now assigned to the Plasmodiophorales they may be found
to have cell-wall composition and flagellar structure that will compel
their removal from their present position. Be that as it may, the order
stands or falls by its type species Plasmodiophora hrassicae. Therefore it is
with this species that comparisons must be made.

In these four orders the resting spores possess a cell wall whose chief
component is chitin in Plasmodiophora but is not definitely determined
in the other groups. Upon germination a naked amoeboid uninucleate cell
is set free, which may lack flagella (Acrasiales, Lahyrinthula of the
Labyrinthulales) or which may have one or two anteriorly attached
flagella (Myxogastrales, Labyrinthomyxa of the Labyrinthulales, and
Plasmodiophora). Where two flagella are present they are both of the
w^hiplash type, neither being of the tinsel type. Sexual reproduction where
reported (Myxogastrales and Plasmodiophora) is by the union of two
myxamoebae or two flagellate swarm spores. The vegetative body of the
organism is a Plasmodium or a more or less loose aggregation of myxa-
moebae. Eventually this separates into naked uninucleate spores around
which a spore wall is produced. Because of the naked vegetative body of
Plasmodia! nat^ure the zoologists have included these four groups among
the Protozoa. The protozoologist Kudo (1946) places these four groups
in Phylum Protozoa, Class Sarcodina, including the Labyrinthulales in
his Order Proteomyxa, and the other three in his Order Mycetozoa.

The genus Reiiculomyxa has recently been described by Miss Nauss
(1949). The organism consists of a central multinucleate Plasmodium
with radiating, forking, and anastomosing branches which appear to be
the chief organs for the capture of the food particles upon which it lives.

36 MYCETOZOA AND BELATED ORGANISMS

In some regards this seems to lie in a position intermediate between the
Myxogastrales and the Proteomyxa, while in some structural features it
shows similarity to the Labyrinthulales. No definite formation of spo-
rangial structures has been observed.

In the author's opinion the primitive ancestors of these four groups
were more or less colonial amoeboid organisms consisting of encysted
cells at one stage of their life history. From these emerged biflagellate
swarm cells, both fiagella being of the whiplash type and nearly or quite
equal. This biflagellate condition has persisted in Plasmodiophora and is
found in a considerable number of the swarm cells of the Myxogastrales,
the presence of two blepharoplasts remaining even when one flagellum is
missing. In the Labyrinthulales in one genus no flagella are known and
in one genus the swarm spores are described as anteriorly uniflagellate.
The questions as to blepharoplast number and type of flagellum are not
solved. In the Acrasiales the flagella are entirely lacking. The Myxo-
gastrales are undoubtedly the furthest developed from the evolutionary
standpoint, in the development of sporangia and capillitium and adapta-
tion to aerial dispersal of the encysted spores. The Acrasiales are probably
closely related. The Labyrinthulales and Plasmodiophora are water or soil
organisms and lack the complicated structure of the Myxogastrales.

Key to the More Important Orders of Mycetozoa

Saprophytes or surrounding and ingesting fungi, bacteria, etc. Sporangia aerial.
Sporangia with thin or thick peridium, and mostly with a capillitium.

Spores upon germination producing an anteriorly uni- or biflagellate swarm
cell (rarely nonflagellate myxamoeba). Order Myxogastrales

Sporangia without peridium and capillitium, spores embedded in a mass of
slime.
Spores producing nonflagellate myxamoebae. Order Acrasiales

Parasites in the cells of algae and of submerged aquatic. plants, forming net-
plasmodia.
No aerial sporangia. Spores upon germination producing myxamoebae or
anteriorly uniflagellate zoospores. Order Labyrinthulales

Parasites producing plasmodia within the cells of roots and stems of higher plants,
a few in algae and aquatic fungi. Swarm spores anteriorly l>iflagellate.

Order Plasmodiophorales

Key to the More Important Families and Genera of Myxogastrales

(Based in Part Upon Macbride and Martin)

Spores produced externally. Family Ceratiomyxaceae

Only genus. Ceratiomyxa

Spores produced internally.

Spores violet, brown, or purplisli gray (rarely ferruginous or colorless).
Capillitium always present.
Sporangia with lime granules (calcium carbonate) in capillitium and often
in peridium also. Family Physaraceae

KEY TO THE MORE IMPORTANT GENERA OF MYXOGASTRALES 37

Fructification aethalioid, capillitium with lime knots. Fuligo
Fructification of plasmodiocarps or of separate sporangia.

Capillitium of nearly uniform anastomosing tubules, containing lime

granules throughout. Badhaniia

Capillitium of threads containing lime knots.
Peridium incrusted with hme.

Sporangium dehiscent circumscissilely. Craterium

Sporangium by introversion thimble-like or vase-like, dehiscence

by petal-like lobes. Physarella

Sporangium not introverted, dehiscence irregular. Physarum
Peridium smooth and shining. Leocarpus

Sporangia with lime on or in peridium, none in the capillitium.

Family Didymiaceae
Calcareous deposits in form of stellate crystals.

Aethalioid. Mucilago

Plasmodiocarpous or of separate sporangia. Didymium

Calcareous deposits not stellate.

Lime in form of closely adjacent peg-like processes. Physarina
Lime in scattered flattened scales. Lepidoderma

Lime forming a continuous shell, peridium mostly double.

Diderma
Sporangium without lime in capillitium and peridium (in stipe and columella
in some species of Diachea). Capillitium of more or less reticu-
lately anastomosing threads.
Columella usually well developed, capillitium arising along its whole
length. Family Stemonitaceae

Stipe and columella calcareous or waxy. Diachea

Stipe and columella never calcareous or waxy.

Capillitial branches forming definite outer network. Stemonitis
Capillitial branches not forming surface network. Comatricha
Columella short or well developed, capillitium arising at apex or in apical
portion. Family Lamprodermaceae

Columella reaching apex of sporangium, capillitium arising from a disk

at its top. Enerthenema

Columella one-third to half the height of sporangium, peridium iri-
descent, capillitium dense, bushy, branches tapering.

Lamproderma
Columella short, capillitium bushy, the tips of the branches with disk-
like fragments of peridium. Clastoderma
Spores violet to ochraceous or pale, columella and true capillitium lacking.
Outer layer of peridium flaking off leaving the inner layer of reticulate thick-
enings which surround the spore mass.

Family Cribrariaceae
Meshes of the reticulum more or less isodiametric in the upper portion of

the sporangium. Cribraria

Thickenings of the peridium wall like the meridians on a globe with very
delicate cross connections. Dictydium

Peridium wall not reticulately thickened.

Sporangia separate, sometimes plasmodiocarpous.

Family Liceaceae
Sporangia mostly sessile, not dehiscing by a lid. Licea

Sporangia mostly stalked, opening by a lid. Orcadella

38 MTCETOZOA AND RELATED ORGANISMS

Sporangia closely appressed, retaining their walls, dehiscent at apex.

Family Tubiferaceae
Sporangia cylindrical, densely clustered. Tubifera

Sporangia ovate in a loose cluster on a common stalk.

Alwisia
Sporangia sessile, flattened, closely clustered into a pseudoaethalium,
rarely scattered. Liceopsis

Forming an aethalium, no sporangial walls remaining at maturity.

Pseudocapillitium thread-like or of perforate or frayed sheets, spores
ochraceous. Family Reticulariaceae

Surface alveolar, made up of the caps of the sporangial units.

Dictydiaethalium

Surface not alveolar.

Pseudocapillitium of flat irregular plates fraying out into threads.

Reticularia
Pseudocapillitium of broad perforated plates. Enteridium
Pseudocapillitium of colorless branched tubes, spores pale.

Family Lycogalaceae
Only genus. Lycogala

Spores yellow to ochraceous, capillitial threads with characteristic markings:
spirals, rings, spines, cogs, etc., sometimes faint or wanting.
Capillitium a network or of separate threads, marked with spiral bands,
threads coarse. Family Trichiaceae

Capillitial threads separate, spirals irregular or faint. Oligonema
Capillitial threads separate, spirals distinct, regular. Trichia

Capillitial threads forming a network.
Spirals regular. Hemitrichia

Spirals irregular or obscured by reticulations. Calonema

CapilUtium a network of coarse threads attached to lower part of peridium,
markings various, never of spirals. Family Arcyriaceae
Capillitium elastic, pushing out far beyond the cup-like base of the

peridium. Arajria

Capillitium not elastic. Lachnobolus

CapiUitial threads slender, warted or spinulose or smooth.
Capillitial threads soUd, peridium usually single.

Family Dianemaceae

Capillitial threads hair-like, coiled. Margarita

Capillitial threads nearly straight. Dianerm

Capillitial threads hollow, peridium double. Family Perichaenaceae
Capillitium warty or spiny, dehiscence irregular. Ophiotheca

Capillitium as above, dehiscence circumscissile. Perichaena

Key to the Genera of Order Acrasiales

Myxamoebae with inconspicuous, rounded pseudopodia.

Fruiting bodies sessile. Copromyxa

Fruiting bodies short-stalked. Guttulina

Myxamoebae with well-developed, more or less acute, pseudopodia. Fruiting
bodies stalked.
Spores in rounded slime-covered heads.

Stalks not branched. Didyostelium

Stalks branched. Pohjsphondylium

Spores in chains, stalks not branched. Acrasis

LITEBATUEE CITED 39

Key to the Genera of Order Labyrinthulales

Germinating resting spores producing nonflagellate cells. Ldbyrinthula

Germinating resting spores producing anteriorly uniflagellate cells.

Lahyrinthomyxa

Key to the Genera of Plasmodiophorales

(Based upon Karling, 1942)

Resting spores not united, mostly nearly filling the host cell. Zoosporangia small,

producing few zoospores. Plasmodiophora

Resting spores in small clusters or united in more or less compact cystosori. Except

Odoniyxa parasitic in tissues of Higher Plants.
Spores mostly in 4's or 2's. Zoosporangia unknown. Tetramyxa

Spores mostly in 8's. Zoosporangia small. In Achlya. Octomyxa

Spores united to form a hollow sphere. Zoosporangia small. Sorosphaera
Spores forming a two-layered flattened disk. Zoosporangia unknown.

Sorodiscus
Spores in a rounded sponge-like mass perforated by large canals. Zoosporangia

mostly small. Spongospora

Spores in variable-sized masses, sometimes loose in the cell which they usually

do not fill. Zoosporangia small. Ligniera

Spore masses variable in size. Zoosporangia large, elongated, with prominent

exit tubes. Polymyxa

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: Occurrence of zoosporangia in Spongospora subterranea (Wallroth)

Lagerheim, ibid., 135:394. Figs. 1-4. 1935.
-: Studies on Polymyxa graminis, n. gen. n. sp., a plasmodiophoraceous

root parasite of wheat, Can. J. Research, C, 17:38-51. Pis. 1-5. Figs. 1-3.

1939.
Lister, Arthur: A Monograph of the Mycetozoa. A descriptive catalogue of the

species in the herbarium of the British Museum. Ed. 3, revised by Gulielma

Lister, xxxiii + 296 pp. 223 pis. 56 figs. London, Trustees of the British

Museum, 1925.
Macbride, Thomas H., and G. W. Martin: The Myxomycetes. A descriptive

list of the known species with special reference to those occurring in North

America, 339 pp. 21 pis. New York, Macmillan Co., 1934.
Nauss, Ruth W.: Reticulomyxa filosa Gen. et Spec, nov., a new primitive

Plasmodium, Bull. Torrey Botan. Club, 76(3):161-173. Figs. 1-13. 1949.
Olh^e,' Edgar W.: Monograph of the Acrasiae, Proc. Boston Soc. Natxmil History,

30:451-513. Pis. 5-8. 1902.
: Cytological studies on Ceratiomyxa, Trans. Wisconsin Acad. Sci., 15(2):

753-774. P/. 47. 1907.
Palm, B. T., and Myrle Burk: The taxonomy of the Plasmodiophoraceae,

Arch. Protistenk., 79(2) :263-276. Figs. 1-15. 1933.

LITERATURE CITED 41

Raper, Kenneth B. : Growth and development of Dictyostelium discoideum with

different bacterial associates, J. Agr. Research, 55(4):289-316. Figs. 1-4.

1937.
Renn, Charles E.: Wasting disease of Zostera in American waters, Nature,

134(3385) :416. FiV. 1. 1934.
: A Mycetozoan parasite of Zostera marina, ibid., 135(3414) :544-545.

1935.
: The wasting disease of Zostera marina, Biol. Bull., 70(1):148-158. Figs.

1-5. 1936.
RocHLiN, Emilia: Zur Frage der Widerstandsfahigkeit der Cruciferen gegen die

Kohlhernie (Plasmodiophora brassicae Wor.), Phytopath. Z., 5(4) :381-406.

Figs. 1-7. 1933.
SiNOTO, Y., AND A. Yuasa: Studies in the cytology of reproductive cells: I. On

the planocytes in five forms of Myxomycetes, The Botanical Magazine

(Tokyo), 48(574) :720-729. Figs. 1-19. 1934.
Skupienski, F. X.: Recherches sur le cycle ^volutif de certains Myxomycetes,

83 pp. 2 pis. 2 figs. Paris, Imprimerie M. Flinikowski, 1920.
: Badania bio-cytologiczne nad Didymium difforme. Czesc pierwsza (Bio-

cytological study of D. difforme. First part. Polish with French summary),

Acta Sac. Botan. Polon., 5(3):255-336. 7 pis. (2 colored). IS figs. 1928.
Sparrow Jr., Frederick K.: Aquatic Phycomycetes, Exclusive of the Sapro-

legniaceae and Pythium, xix + 785 pp. 634: figs. Ann Arbor, Univ. Michigan

Press, 1943.
Valkanov, Alexander: Protistenstudien: 4. Die Natur und die systematische

Stellung der Labyrinthuleen, Arch. Protistenk., 67:110-121. Figs. 1-10. 1929.
Webb, P. C. R.: Zoosporangia believed to be those of Plasmodiophora brassicae

in the root hairs of non-cruciferous plants, Nature, 163(4146) :608. 1949.
VON Wettstein, Fritz: Das Vorkommen von Chitin und seine Verwertung als

systematisch-phylogenetisches Merkmal im Pflanzenreich, Sitz. her. Akad.

Wiss. Wien, Math, naturw. Klasse, Abt. I, 130(1) :3-20. 1921.
Wilson, Malcolm, and Elsie J. Cadman: The Ufe history and cytology of

Reticularia Lycoperdon Bull., Trans. Roy. Soc. Edinburgh, 55:555-608. Pis.

1-6. Figs. 1-4. 1928.
van Wisselingh, C: Mikrochemische Untersuchungen iiber die Zellwande der

Fungi, Jahrb. wiss. Botan., 31(4):619-687. Pis. 17-18. 1898.
WoRONiN, M.: Plasmodiophora brassicae, Urheber der Kohlpflanzen-Hernie,

Jahrb. wiss. Botan., 11:548-574. Pis. 29-34. 1878.
Young III, Edward Lorraine: Studies on Labyrinthula. The etiologic agent

of the wasting disease of eel-grass. Am. J. Botany, 30(8) :586-593. Figs. 1-2.

1943.
Yuasa, Akira: Studies in the cytology of reproductive cells :III. The genesis of

the flagellum in the planocytes of Fuligo septica Gmelin, The Botanical

Magazine (Tokyo), 49(584) :538-545. Figs. 1-27. 1935.

PHYCOMYCETEAE: CHYTRIDIALES AND
HYPHOCHYTRIALES

THE first class of true fungi, for the Mycetozoa discussed in Chapter 2
are not considered to be fungi or even plants, is that of the Phyco-
myceteae. In this class the fungus body may consist of

1. One cell — with a cell wall and with or without non-nucleate
rhizoids — which either becomes directly a zoosporangium or gametangium
or else empties its contents into an external zoosporangium or
gametangium.

2. Instead of a single cell with non-nucleate rhizoids, nuclei may pass
from the first cell through modified rhizoids and take their positions in
swellings in the latter, there forming secondary centers. Such fungi are
polycentric in contrast to the monocentric forms with but a single cell.
When the fungus is entirely contained in the cell of its host and com-
pletely lacks rhizoids or haustoria, it is called holocarpic; when it is
external to its substratum, or internal and gains its nourishment by
means of rhizoids or haustoria, it is called eucarpic.

3. A still further advance is the development of a true coenocytic
mycelium containing many nuclei, usually with rhizoids or nucleate
trophic hyphae within the substratum. In general this type of mycelium
is not provided with true septa (although greatly perforate pseudosepta
occur in Allomyces) except to fence off sporangia and gametangia, injured
regions, or portions of the mycelium that have exhausted their proto-
plasmic contents.

In seven of the twelve orders treated here as belonging to the Phyco-
myceteae asexual reproduction and, to a much lesser degree, sexual repro-
duction is by means of planocytes — i.e., flagellate, naked cells. These are
of three types which appear to characterize three series of orders. The
types are as follows: flagellum single, posterior; flagellum single, anterior;
flagella two, anterior or lateral. In the other five orders (the relationship
of two of which is greatly in doubt), asexual reproduction is by means of

KEY TO THE ORDERS OF BIFLAGELLATE FUNGI 43

nonflagellate cells and sexual reproduction is by the union of two gametes
or two gametangia. The Keys given below will serve to distinguish the
orders of this class.

Key to the Orders of Posteriorly Uniflagellate Fungi

Without true mycelium, at most a rhizomycelium. Alternation of sporophytic
and gametophytic generations wanting. Zoospores mostly without large
nuclear cap or "side body." Order Chytridiales (Chap. 3)

Rarely without mycelium, mostly with multinucleate, coenocytic mycelium.
Sexual reproduction, where known, by the union of isoplanogametes or
anisoplanogametes. Alternation of sporophytic and gametophytic genera-
tions often present. Zoospores with large nuclear cap and "side body."

Order Blastocladiales (Chap. 4)

With multinucleate, coenocytic mycelium. Sexual reproduction by the union of
motile sperm with non-motile egg, to form thick-walled oospore. Alternation
of sporophytic and gametophytic generations wanting. Zoospores with
numerous small granules at the apex. Order Monoblepharidales (Chap. 4)

Key to the Order of Anteriorly Uniflagellate Fungi
Only one order recognized. Order Hyphochytriales (Chap. 3)

Key to the Orders of Anteriorly or Laterally Bifiagellate Fungi

Holocarpic fungi, parasitic within animal or plant hosts (rarely saprophytic),
the whole organism being converted into a single reproductive organ (game-
tangium or zoosporangium) or a series of such organs, separated by septa.
Zoospores of primary type or much oftener of secondary type, preformed in
the zoosporangium or becoming organized in a vesicle formed at the apex
of the exit papilla or tube. Sexual reproduction by the union, sometimes
through a short conjugation tube, of two gametangia of equal size or more
often of an antherid and oogone. Oospore usually not surrounded by peri-
plasm. Order Lagenidiales (Chap. 5)

Eucarpic fungi usually (but not always) with well-developed branched and
tapering holdfast system. The external fungus body [a system of usually
stout cylindrical, branched coenocytic hyphae, nearly uniform in thickness
or with large basal segments and slender branches. Reproductive organs
borne on the branches and making up only a small part of the whole fungus.
Asexual reproduction by bifiagellate zoospores, often dimorphic, rarely of
primary type only, often only of secondary type. Oogones with one or more
eggs, with or without periplasm. Fertilization by antherids to form thick-
walled oospores. Mostly saprophytic in soil and fresh water, in some cases
parasitic in roots, algae, fungi, or fresh-water animals.

Order Saprolegniales (Chap. 5)

Eucarpic fungi consisting of slender, cylindrical hyphae with or without haus-
toria, mostly growing within the tissues of living hosts (mostly plants), or in
dead organic matter in soil or water. Zoospores of secondary type only,
produced under water in slender or rounded zoosporangia and organized in
a vesicle at the mouth of the exit tube, or the zoosporangia (coiiidia) are
borne aerially and distributed by air currents, the formation of the zoospores
occurring in these conidia after they fall into water. In some genera zoospore
formation is omitted and the conidia germinate by germ tubes. Sexual

44 PHYCOMYCETEAE

reproduction by oogones with usually one egg and periplasm, fertilized by an
antherid through a conjugation tube. Order Peronosporales (Chap. 6)

Key to the Orders of Nonflagellate Phycomyceteae

Walls of the mycelium readily responding to the test for cellulose by chloriodide
of zinc.

Parasites entirely within the tissues of the aerial parts of higher plants and
forming on their branched, occasionally septate, coenocytic hyphae large,
thick-walled intercalary or terminal resting spores within which, upon ger-
mination, are produced numerous small spores which upon escaping may
unite by twos and then infect other host plants. Relationship very doubtful.

Order Protomycetales (Chap. 6)

Parasites in the alimentary canals of Arthropoda, forming slender, at first non-
septate, coenocytic hyphae which very rarely branch. The contents of the
hyphae are transformed by formation of septa into endospores which escape
through basal or apical openings. Larger endospores may be formed in which
by a sexual (?) process there is a union of nuclei. Relationship very doubtful.

Order Eccrinales (Chap. 7)
Walls of the mycelium not readily responding to tests for cellulose. Chitin-like
substances present.

Saprophytes or more rarely parasites (mainly in fungi). Mycelium relatively
large and abundant, more often nonseptate, at least when young. Asexual
reproduction by aplanospores produced in sporangia or the latter reduced
to conidium-like sporangioles. Sexual reproduction by the union of unequal
or almost equal gametangia to form "zygospores."

Order Mucorales (Chap. 7)

Parasites in insects or in desmids or fern-prothallia or fungi or saprophytes on
dung of lizards and frogs. Mycelium at first coenocytic but early becoming
septate and very often breaking up into short "hyphal bodies." Asexual
reproduction mostly by formation on external conidiophores of terminal
conidia which are shot off with violence. Sexual reproduction by union of
apparently equal gametangia to form thick-walled zygospores.

Order Entomophthorales (Chap. 7)

Parasites in soil-inhabiting amoebae and nematodes or in the alimentary canal
of aquatic insects. Mycelium (except where thickened haustoria are pro-
duced) slender, at first coenocytic. Asexual reproduction by the formation
of terminal, ellipsoidal or fusiform conidia, single or in chains, or of similar
lateral conidia. Sexual reproduction by the union of apparently equal game-
tangia, adjacent in the same hypha or in separate hyphae lying in close
proximity, or at the ends of long, slender filaments.

Order Zoopagales (Chap. 7)

Order Chytridiales. The Chytridiales are comparatively simple in
their structure and for that reason are considered first. Whether this
simplicity is due to their primitive nature or whether it is due to simplifi-
cation from more complex forms is a matter of dispute and will be dis-
cussed at more length in Chapter 17 on the Phylogeny of the Fungi.

This order includes a number of families of fungi that are either largely
aquatic or depend upon the presence of water for their dispersal. They
are either parasitic in the roots, stems, or leaves of higher plants; in algae;

ORDER CHYTRIDIALES 45

in fungi; in the eggs or larvae of worms, Arthropoda, or simpler animals;
or, perhaps more often, saprophytic in dead plant or animal material.
They are abundant in some types of soils and in fresh and salt water.
Their life histories are poorly known in many cases but have been worked
out very well in some species. In general the life history is as follows : The
posteriorly uniflagellate zoospore after a period of swimming settles down
on the substratum and penetrates it or encysts on the outside and then
enters it wholly or merely as a more or less extensive haustorium or
system of rhizoids. Unlike the Mycetozoa it produces a cell wall of its
own, usually at an early stage. The single nucleus divides repeatedly as
the cell enlarges, and eventually the cytoplasm fragments into very many
uninucleate zoospores or naked gametes or into uninucleate cells which
produce cell walls and sooner or later divide internally into zoospores or
naked gametes. Sexual reproduction where known may be accomplished
by the union of two motile gametes before entry into the substratum,
by the union of two cells attached to or residing within the substratum,
or by the union of two cells by means of a rhizoid through which a gamete
nucleus passes.

In this order the zoospores or motile gametes escape through an exit
papilla or tube whose apex softens and permits the motile cells to push
out (the inoperculate series), or they escape through a sort of cap that
opens like a trap door, the so-called operculum (the operculate series).
This operculum is more generally at the apex of the papilla or tube, but in
Karlingia (Johanson, 1944) and Catenomyces (Hanson, 1945) it may be
formed within the tube some distance from the apex, which softens and
deliquesces much as in the inoperculate forms. The single posterior
flagellum is of the "whiplash" type, i.e., the basal portion consists of a
firmer outer tube whose more fluid contents extend beyond its tip as a
more slender, very flexible structure resembling the lash of an old-
fashioned buggy whip. This lash is sometimes very short or scarcely
distinguishable.

The cell wall of the Chytridiales does not ordinarily give the charac-
teristic blue or violet coloration of cellulose when chloriodide of zinc is
applied, but there are a number of exceptions in which this reaction is
distinct. Whether chitin is actually present in all cell walls that do not
show the cellulose reaction is not certain. Harder (1937) reported that in
one species of the Family Rhizidiaceae different individuals possess
cellulose or chitin, perhaps in cells of different ages. Nabel (1939) demon-
strated the presence of chitin but not of cellulose in some species of
Rozella and Synchytriuni. Schwartz and Cook (1928) report the presence
of cellulose in the cell walls of Olpidium radicale while Scherffel (1925)
states that in certain structures of Micromycopsis cristata Scherffel and
Synchytrium mercurialis (Lib.) Fuckel cellulose is present, Tischler (1927)

46 PHYCOMTCETEAE

reports the haploid chromosome number in the Chytridiales to be usually
4 or 5 but 10 to 12 in Polyphagus euglenae Now.

A plant body which consists of but a single rounded cell without any
rhizoids or haustoria is said to be holocarpic, but if such structures are
present it is eucarpic.

In the majority of the eucarpic genera (the monocentric series) the
zoospores produce but one enlargement from which, directly or indirectly,
the /oosporangium arises. In several genera (the polycentric series) the
germinating zoospore produces an enlarged cell from which arise rhizoids
which here and there in their course produce other enlargements into
which enter nuclei derived from the original cell. These secondary en-
largements may become zoosporangia themselves or may give rise to
zoosporangia as well as to other rhizoids with further swellings.

Until recently it has been customary (Fitzpatrick, 1930) to include in
the Order Chytridiales all those organisms with structures and life
histories similar to those outlined above. The differences in the types of
flagellation of the motile cells and of the composition of the cell wall
have led mycologists recently to exclude from this order those genera
with a single anterior fiagellum of the tinsel type (Order Hyphochytriales)
and those with two anterior or lateral flagella, one of the tinsel and one of
the whiplash type (Family Olpidiopsidaceae).

Sparrow (1942) makes the primary division of this order, on the basis
of mode of escape of the zoospores, into two series: Inoperculatae and
Operculatae. Miss Whiff en (1944) on the contrary considers the mono-
centric or polycentric structure to be of prime importance, using the
method of zoospore escape as a subordinate character.

In the treatment of this order the author follows mainly the arrange-
ment of Miss Whiffen, with constant reference to the work of Sparrow.
First, the holocarpic forms are considered. Whether these are more
primitive than the eucarpic monocentric forms, or have arisen from some
of these by loss of the rhizoidal or haustorial apparatus in connection
with their habitat entirely within the host cell, must remain undecided
at present. Sparrow recognizes three families, the relationship of the
second of which is somewhat doubtful. They are distinguished as follows:

Olpidiaceae: vegetative cell enlarging to form a single sporangium.
Achlyogetonaceae: vegetative cell elongated and by septation forming a linear

'series of sporangia.
Synchytriaceae : vegetative cell dividing internally into numerous sporangia, or

the latter formed within an outgrowth from the vegetative cell.

Family Olpidiaceae. This family contains six or more genera,
totaling forty or more well-substantiated species and many more de-
scribed species whose status, according to Sparrow, is uncertain. They
are entirely endobiotic, i.e., living entirely within the host cells or tissues.

ORDER CHYTRIDIALES 47

They are more frequently parasitic but may occur as saprophytes within
the dead host cells. The posteriorly uniflagellate zoospore settles on the
exterior of the host and the flagellum disappears, probably in most cases
being withdrawn into the body of the cell which then becomes covered
with a thin wall. A slender infection tube grows through the host cell wall
and the contents of the encysted zoospore pass into the host. The empty
cyst soon disappears. Within the host cell the uninucleate fungus enlarges
and usually early produces a cell wall. In some genera, e.g., Olpidium, the
fungus cell does not completely fill that of the host, but in Rozella, as it
expands, the fungus eventually completely fills the host cell, the walls
of the fungus and host coming into close apposition. The most important
genus of the family is Olpidium which is parasitic in algae in fresh or salt
water, in pollen grains or fungus spores that have fallen into the water,
in small aquatic animals or their eggs, as well as in the roots of various
land plants. 0. hrassicae (Wor.) Dang, is parasitic in the roots of cabbage
and other plants of the genus Brassica, and 0. viciae Kus. inhabits the
roots of vetch (Vicia). In these the posteriorly uniflagellate zoospore
becomes attached to the root of the host plant and encysts there. The
contents of the cyst dissolve a small hole through the host cell wall and
enter the cell. It forms at first a naked uninucleate mass near the center
of the host cell. It may remain there or may dissolve its way into the next
underlying cell or even further. Eventually the fungus cell encysts and
begins to grow. Alore than one fungus cell may infect the same host cell,
up to 15 or 20 in 0. viciae. In this case they do not attain so great a size
as when single and become flattened where they press against one another.
As the fungus cell enlarges the nucleus divides repeatedly. Eventually the
organism consists at maturity of a smooth, thin-walled, more or less
round or ellipsoid zoosporangium, within which cleavage into uniflagellate
zoospores occurs. These escape through one (rarely more) exit tube which
grows to the outside of the host and then permits the zoospores to escape
by the softening and giving way of the apex of the tube. Sexual repro-
duction in 0. viciae and in 0. trifolii Schroet. occurs, according to Kusano
(1912, 1929), by the union of two zoospores (in this case functioning as
gametes) outside the host, the resulting biflagellate zygote encysting and
infecting the host in the manner described above. The resultant cell
becomes thick-walled and is often more or less angular. It is usually
smaller than the asexually produced sporangia. These thick-walled resting
spores may live for a period of several months until favorable conditions
occur, when they swell, cracking open the outer thick wall and permitting
the emergence of an exit tube from which the zoospores escape. The
union of the two nuclei does not occur until the following spring shortly
before the germination of the resting spores. It is probable that meiosis
occurs early in the series of nuclear divisions within the resting spore.

Fig. 9. Chytridiales, Family Olpidiaceae. Olpidium viciae Kus. (A) Zoospores. (B)
Zoospores uniting. (C) Biflagellate zygotes. (D) Infection of host cell by zygote. (E)
A thin-walled zoosporangium and a resting zoosporangium in the same host cell. (F)
Thin-walled zoosporangia in host cells. (G) Host cell with many resting zoosporangia.
(H) Resting zoosporangium discharging zoospores. (After Kusano: J. Coll. Agr., Imp.
Univ. Tokyo, 4(3):141-199.)

ORDER CHTTRIDIALES

It is likely that all zoospores of these species are potentially either
gametes or zoospores, depending upon the environment and the oppor-
tunity for union of the cells. The earlier emerging zoospores usually do
not fuse but those emerging later, especially if from separate zoosporangia,
do. For most species of the genus, sexuality has not been demonstrated.
Treatment with chloriodide of zinc does not reveal any cellulose reaction
in those species of the genus that have been tested except in 0. radicale
Schwartz and Cook (1928). (Fig. 9.)

The genus Pseudolpidiopsis was founded by Minden (1915) to include
an organism studied by Zopf (1884), parasitic
in algae of the family Zygnemataceae , which
was similar to Olpidium in asexual reproduc-
tion but different in sexual reproduction. He
named it Olpidiopsis schenkiana Zopf. Sexual
reproduction occurs within the host cell by
the union of two adjacent cells, the contents
of one passing into the other which forms a
thick wall to which the empty smooth cell
wall of the male cell adheres. Since this is the
type of sexual reproduction characteristic of
the genus Olpidiopsis in the Order Lageni-
diales doubt has been thrown on the accu-
racy of Zopf's description and figures (1884)
of the zoospore as posteriorly uniflagellate.
Sparrow (1942) and others are inclined to
unite the two genera and place them in the
Olpidiopsidaceae. Pleotrachelus, with many
exit tubes from the sporangium is otherwise
similar to Olpidium. Nothing is known as
to the sexual reproduction although resting
spores are produced in one of the three
definitely known species. These species occur
in molds of the genus Piloholus and in the fresh-water alga Oedogonium.
(Fig. 10.)

Rozella includes eleven species recognized by Sparrow. They are all
parasitic in fungi of the Blastocladiales, Monoblepharidales, Sapro-
legniales, Peronosporales, Chytridiales, and a few other groups, mostly
in fresh water but one species in salt water. After entering the host cell
or filament, which frequently becomes considerably hypertrophied, the
organism remains naked for some time, finally producing a thin wall
which fuses with the host wall. In some species the organism becomes
septate into a series of several sporangia. The posteriorly uniflagellate
zoospores escape through one or more inoperculate exit papillae. Resting

Fig. 10. Chytridiales, Fam-
ily Olpidiaceae. Pleotrachelus
fulgens Zopf. Zoosporangium
in Piloholus sp. (After Zopf.
Courtesy, Sparrow : Aquatic
Phycomycetes, Exclusive of
the Saprolegniaceae and Pyth-
ium, Ann Arbor, University
of Michigan Press.)

50 PHTCOMTCETEAE

spores are formed in several species and are smaller than the host cells,
lying free from the host wall. They may be smooth or spiny. Sexual repro-
duction is unknown. Sphaerita is an incompletely known genus of which
S. dangeardii Chatton and Brodsky is parasitic in species of Euglena.
The description of the single flagellum attached anteriorly but trailing
posteriorly casts doubt on the correctness of its position here since in the
genus Pseudosphaerita of the Olpidiopsidaceae the two flagella, attached
near the anterior end are unequal in length, the shorter one anterior and
the longer one trailing posteriorly. Several other genera are recognized
but will not be discussed here.

Family Achlyogetonaceae. This family containmg three genera,
occurs in fresh-water algae, and reportedly in some Nematodes. In the
genus Septolpidium, occurring in Diatoms, the fungus elongates within
the host cell and by successive septation forms a series of zoosporangia
from which the posteriorly uniflagellate zoospores escape through single
exit tubes.

Family Synchytriaceae. In this family we find fungi whose para-
sitism is confined largely to the higher plants. Synchijlrium is the chief
genus. Its swarm spores possess one posterior flagellum. After settling
upon the host, which only occurs in a drop or film of water, the swarm
spore dissolves an opening into an epidermal cell and enters as a whole,
not leaving an empty cyst attached externally. The presence of the
parasite usually stimulates the host cell to hypertrophic growth. This
cell may elongate and enlarge to a balloon-shaped external structure as in
Erodium cicutarium (L.) L'Her. infected by S. papillatum Farlow, the
other cells of the host remaining normal. The enlargement of the epi-
dermal cell may be chiefly lateral and inward in some plants so that it
does not project from the surface but more or less crushes the adjacent
host cells. The enlargement may be both outward and inward, some of
the adjacent cells developing hyperplastically and even undergoing hyper-
trophy, the result being a wart-hke structure, as in S. vaccinii Thom. on
the cranberry, Oxycoccus macrocarpus (Ait.) Pursh. When great hyper-
plasia and hypertrophy both occur in the host tissues, the fungi may
remain in the inwardly enlarged epidermal cells or by the division of the
latter may come to lie more deeply in the massive hyperplastic tissues.
This is the case in S. endohioticum (Schilb.) Perc, the cause of the wart
disease of the potato, Solanum tuberosum L. M. T. Cook (1945) has made
a study of the different types of response by the host to infection by
different species of Synchytrium. (Fig. 11.)

Within the host cell the ])arasite enlarges and soon develops a cell
wall. This is reported by von Wcttstein (1921) and von Gutenburg (1909)
to contain cliitin, a component of the cell walls of most of the higher
fungi. At first the parasites remain uninucleate. By division of the nucleus

OBDER CHTTEIDIALES

Fig. 11. Chytridiales, Family Synchytriaceae. Synchytrium endobioticuni (Schilb.)
Perc. (A) Zoospores. (B-D) Stages in the infection of the host. (E) Parasite in the
upper part of host cell. (F) Parasite enlarged. (G, H) Sorus pushing out from pro-
sorus. (I) Beginning of nuclear multiplication in sorus. (J) Beginning of segmentation
in sorus. (K) Sorus containing five zoosporangia. (L) Single zoosporangium containing
swarm cells. (M-0) Stages in union of swarm cells. (P-R) Infection of host by zygote.
(S) Resting sporangium in host cell. (T) Resting sporangium producing swarm cells.
(After Curtis: Trans. Roy. Soc. London, B, 210:409-478.)

PHYCOMYCETEAE

and delimitation of walled uninucleate cells the parasite becomes a
"soms." These cells may remain thin-walled and polyhedral by mutual
pressure or may round up and become separate, with thicker walls.
Within each of these cells the nucleus divides until a large number
(sometimes hundreds) are present. As these sporangia enlarge, the whole
cell is ruptured and the sporangia themselves escape separately, as in
S. decipiens (Peck) Farlow, or rupture within the host cell releases the
swarm spores. In many species the original fungus cell does not directly
become the sorus of sporangia but while still in the uninucleate condition
buds out into a thin-walled sorus in the outer part of the host cell, the
sporangia and swarm spores being formed in this sorus instead of in the
original cell, w^hich in this case is called a prosorus.

The swarm spores may infect the host directly or may unite by twos,
forming biflagellate zygotes which settle on the epidermis of the host
and encyst, the nuclei then fusing. The host cell is then entered and after
some growth the parasite becomes a thick-walled resting spore. After a
longer or shorter period this may function directly as a sporangium, or
as a sorus of sporangia, or as a prosorus, releasing swarm spores which
infect the host asexually or after sexual union with other swarm spores.
In some species there are several generations in a season, at least of the
asexually produced individuals, while in others only the thick-walled
overwintering generation is known. Whether this is asexually or sexually
produced has not been determined. Sexuality has been demonstrated for

Fig. 12. Cliytridiales, Family Syncliytriaceac. Synchytrium fulgrns Schroet.
(1-8) Summer cycle. (I-VIII) Winter cycle. (After Kusano: Japanese J. Botany,
5(1):35-132.)

ORDER CHYTRIDIALES 53

only a few species as yet, e.g., S. endohioticum by Miss Curtis (1921) and
S. fulgens Schroet., on Oenothera, by Kusano (1930). In the former the
overwintering sexually produced resting cells become deep-seated in the
tissues of the host by the hyperplastic division of the cells. These resting
cells become sporangia directly, not sori of sporangia. In S. fulgens both
the summer and winter generations become prosori. (Figs. 11, 12.)

Two species of Synchytrium are of economic interest, S. vaccinii,
causing small galls on the leaves and fruits of the cranberry (Oxycoccus
macrocarpus) , and S. endohioticum, the cause of the very destructive wart
disease of the potato.

Three other genera occur in this family as parasites in algae. One of
these is Micromyces of which the species M. zygogonii Dang, and M. longi-

FiG. 13. Chytridiales, Family Synchytriaceae.
Micromyces longispinosus Couch. (A) Sorus, previously
extruded from the spiny prosorus, breaking up into
numerous zoosporangia, some of which are discharging
swarm cells. (B) Spiny resting spore discharging a
sorus, only one nucleus present. (Courtesy, Couch:
M?/co/o6ria, 29(5):592-596.)

spinosus Couch have been observed in America by Couch (1937) in
Mougeotia and Spirogyra, respectively. The infected cells of the host are
usually somewhat enlarged. The plant body of the fungus is spherical
with numerous long spines. Through a small opening the contents emerge
and form a thin-walled sorus which divides into several sporangia. Within
these are produced numerous uniflagellate swarm spores which may infect
the host directly or may first unite by twos. The smaller, thick-walled
resting spores, which are likewise spiny, are probably the product of
infection by the zygotes, but this has not been proved. The resting spore
germinates by the extrusion of a sorus within which sporangia are pro-
duced. The zoospores escape when the host cell, which has usually become
much swollen, bursts at one side. Scherffel (1925) sets apart as the genus
Micromycopsis some species in which there is an exit tube from the fungus
body through the wall of the algal host so that the sorus, containing two

54 PHYCOMYCETEAB

or three sporangia, is external to the host cell instead of internal as in
Micromyces. He points out that though in general the cell walls of the
Synchytriaceae do not show a cellulose reaction, yet the wall of the
sporangial sorus and of the tube on which this is borne becomes red-violet
in color with iodine in potassium iodide solution, in Micromycopsis cristata
Scherffel and also in Synchytrium mercurialis (Lib.) Fuckel. Miss Canter
(1949) has described a third genus in this family parasitic in fresh-water
algae, the genus Endodesmidium, parasitic in Desmids. Like the two fore-
going genera this is endobiotic and holocarpic. The prosorus of the only
recognized species is smooth-walled, not spiny as in Micromyces and most
species of Micromycopsis. The relatively large sorus grows out of one end
of the prosorus within the cavity of the host cell. The contents divide into
numerous, movstly spherical sporangia which escape through papillae into
the cavity of the host wall or into the surrounding medium if the papilla
pierces the cell wall. These sporangia occasionally possess a posterior
flagellum which is feebly active. They produce 2 to 5 minute, posteriorly
uniflagellate zoospores which swim actively. (Fig. 13.)

The eucarpic, monocentric Chytrids fall into two families according
to Miss Whiffen and three according to Sparrow, who sets the operculate
forms apart from the inoperculate as a distinct family. In general the
fungus body can be distinguished into a uninucleate enlargement and a
non-nucleate haustorial or rhizoidal system attached directly or indirectly
to the former. This varies from a short peg to an extensive mass of usually
tapering and more or less branched rhizoids. These may be entirely
intramatrical or, except for the tips of the branches, may be entirely
extramatrical. The body of the encysted zoospore may enlarge and be-
come the sporangium or it may become a prosporangium from whose apex
the sporangium arises. On the other hand an enlargement of the germ
tube may be formed, the contents of which eventually pass into the
sporangium which arises by enlargement of the original zoospore cyst.
In Miss Whiffen's family Entophlyctaceae the zoospore cyst empties
itself completely into the subjacent portion of the germ tube, this enlarge-
ment becoming the sporangium or a prosporangium out of which the
sporangium buds. The old zoospore cyst falls away or remains only as an
empty fragment.

The eucarpic monocentric families may be distinguished as follows
according to Miss Whiffen:

Rhizidiaceae: zoospore cyst enlarging into a sporangium or prosporangium.
Entophlyctaceae: zoospore cyst not further functional; the upper part of the
germ tube enlarging into a zoosporangium or prosporangium.

Both of these families as delimited by Miss Whiffen contain operculate
as well as inoperculate genera. Sparrow segregates the operculate forms
into the family Chytridiaceae. He divides the inoperculate genera on the

ORDER CHTTRIDIALES 55

basis of their position with reference to the substratum into family
Phlyctidiaceae, epibiotic and endobiotic, and family Rhizidiaceae, inter-
biotic, i.e., only the tips of the rhizoids penetrating into the substratum.

Family Rhizidiaceae. As in the Olpidiaceae and Synchytriaceae, the
swarm spores in this family are provided with but one, posterior, flagellum.
Upon reaching the host they send each a haustorial process into the
matrix, the main body of the encysted swarm spore remaining outside.
The haustorium may be a short undivided peg-like structure or slender
and more or less branched, sometimes penetrating other host cells as well.
The external portion may enlarge directly to form the zoosporangium or
the latter may be formed above it, or as a swelling of the subjacent part
of the haustorium. The organism remains uninucleate until the spo-
rangium begins to develop when the nucleus divides many times to form
the nuclei of the swarm spores. The haustorium does not at any time
contain any nuclei and can hardly be considered as homologous to a
mycelium. Resting cells are produced in some species but their mode of
origin is unknown in most cases. Sexual reproduction by the union of two
well-developed cells has been observed in a few forms. Fusion of swarm
spores has been reported but whether it is a true sexual process or merely
a sort of rejuvenescence of weak or exhausted cells has not been deter-
mined. Petersen (1903), Sparrow (1936), and Karling (1945) have demon-
strated sexual reproduction in Siphonaria, and Sparrow (1937) also in
Rhizoclosmatium and in Asferophlyctis. A special rhizoid-like outgrowth
proceeds from one cell, presumably to be considered the male cell, to
another cell. This tube may be long or short or in some cases the two cells
are in direct contact. When this process is completed the female cell
enlarges and forms a thick-w^alled resting spore. After a variable length of
time this zygote serves as a prosporangium, a small pore being produced
and the contents emerging, enclosed by a thin cell wall, and forming an
external zoosporangium. The host organisms of the parasitic forms of this
family are mostly algae, pollen grains, or small aquatic animals, but
Rhizophydium graminis Ledingham is parasitic in the roots of Triticum
and Panicum. Many species are saprophytic.

Phlyctochytrium grows on algae into which its much-branched haus-
torium penetrates. This arises from an apophysis just within the host
wall while the portion external to the wall enlarges to become the zoospo-
rangium. Only the presence of the apophysis distinguishes it from
Rhizophydium. (Fig. 14.)

Rhizophydium occurs mostly in water on various substrata. It con-
sists, when mature, of an enlarged, more or less spherical, thin-walled
external sporangium, with a usually tufted haustorium within the host
cell. Upon the maturity of the numerous swarm spores they escape through
one or more inoperculate pores in the sporangium wall. Sometimes the

PHYCOMYCETEAE

Fig. 14. Chytridiales, Family Rhizidiaceae. Phlyctochytrium hallii Couch. (A)
Plant nearly mature. (B) Zoospores differentiated. (C) Discharge of zoospores. (D, E)
Resting sporangia. (After Couch: J. Elisha Mitchell Sci. Soc, 47(2):245-260.)

Fig. 15. Chytridiales, Family Kliizidiaceae. Rhizophydium coronum Hanson. (A)
Zoospore. (B) Zoosporangium with swollen exit papilla. (C) Mature resting spore
(stained). (D) Germinating resting spore. (Courtesy, Hanson: Am. J. Botany,
32(8):480-483.)

ORDER CHYTRIDIALES

external cell becomes a thick-walled resting spore which produces its
swarm spores only after some time. Couch (1932) has shown for R. couchii
Sparrow that these resting spores arise as follows: A zoospore "comes to
rest on the host, penetrates the wall, and develops apparently just like a
sporangium. Later another spore comes and attaches itself to the larger
body. . . . The smaller cell discharges its entire contents into the larger.
. . . This now secretes around itself a thick wall and goes into the resting
state." In R. ovatum Couch the same author (1935) reports that the male
cell encysts on the algal host (Stigeodonium) and produces its rhizoids.
The female cell comes to rest upon the male cell and both cells enlarge,
the Avail between becoming perforated. The male nucleus passes into the
larger female cell and unites with its nucleus. This cell becomes a thick-
walled resting spore. (Fig. 15.)

Rhizophlydis differs from Rhizophydium mainly in that the rhizoids
are numerous and arise from more than one point on the sporangial wall.
The four or five species are parasitic on fresh-water algae or saprophytic
on insect exuviae and vegetable debris. R. petersenii Sparrow can be
cultivated on cellophane or filter paper in pure water cultures. Karlingia
rosea (de Bary and Wor.) Johanson which differs in the production of an
operculum near the base of each of the several exit papillae was formerly
assigned to this genus under the name R. rosea. This species grows
saprophytically on organic matter in the soil, the large sporangia some-
times exceeding 0.1 mm. in diameter. These become rose-colored and give
a rosy tinge to the soil. This species also may be cultivated on cellophane
and various kinds of vegetable matter such as sterilized onion skin, grass
leaves, etc. The position of the operculum at the base of the exit papilla
instead of at its apex is of interest.

Ohelidium grows upon the exuviae of aquatic insects. Its external body
bears a spine and a cup or funnel-like base, and the branching rhizoids
arise from a small apophysis. The zoospores escape from a lateral, ino-
perculate opening below the spine. No resting stage is known. It occurs
in Europe and Asia and Sparrow (1938) reports 0. mucronatum Now.
also from the United States. Siphonaria forms a round or ellipsoid
zoosporangium and strongly developed rhizoids. One species has spines
laterally and apically but does not possess the cup or funnel-like base of
the preceding genus. Sexual reproduction occurs by the union of two
thalli through a short or long slender tube, the female thallus becoming
a thick-walled resting spore. (Fig. 16.)

The genus Polyphagus is usually placed in this family but its rela-
tionship to the other genera is doubtful. P. euglenae Now. is parasitic on
species of Euglena and other one-celled green organisms. Instead of being
an internal parasite it lives externally. According to Wager (1913), the
germinating zoospore sends out in various directions slender processes

PHTCOMYCETEAB

Fig. 16. Chytridiales, Family Rhizidiaceae. Siphonaria petersenii Karl. (A) Mature
plant with zoospores escaping. (B) Union of male and female plants through a narrow-
tube. (C) Sexually produced thick-walled zygote with empty male plant still attached.
(Courtesy, KarHng: Am. J. Botany, 32(9):580-587.)

which enter the host cells encountered. Sometimes, where the latter are
crowded in considerable numbers, as many as fifty may be attacked by
the haustoria from one parasite. The latter remains uninucleate and is
invested by a firm thin wall. Within the swelling (prosporangium) repre-
senting the original zoospore, the nucleus divides to form the nuclei of
the new zoospores, and the whole contents bud out into a somewhat
elongated thin-walled sporangium within which the division into the
uninucleate zoospores takes place. As many as several hundred zoospores
may be produced. Upon the occurrence of conditions unfavorable for
further asexual reproductions there may occur the conjugation of two
cells. A somewhat smaller cell sends out a slender process (perhaps a
modified haustorium), the tip of which enlarges when it comes into
contact with a larger cell. Into this enlarged tip the nucleus and contents

ORDER CHTTRIDIALES 59

of the original cell pass. Then the nucleus of the other cell with which it
is in contact enters through a small opening and the wall thickens to form
a thick-walled resting spore. After several months a zoosporangial sac is
formed in which the two nuclei fuse and then divide to form the nuclei
of the zoospores. Three species of Polyphagus are recognized by Sparrow.

The genus Harpochytrium is sometimes placed in this family. The
plant consists of a slender or stout tubular, straight or curved, and sessile
or stalked zoosporangium with blunt or pointed apex. It adheres to the
surface of the host (mostly fresh-water filamentous algae) by a foot,
rarely penetrating the algal wall. When young it is uninucleate but be-
comes multinucleate as it grows. The upper half to three-quarters of the
protoplasm becomes divided transversely into four or five up to many
posteriorly uniflagellate zoospores which escape out of an inoperculate
apical opening, attaching themselves to a host cell by the tip of the
flagellum. The protoplasm, containing one or more nuclei, which remains
in the zoosporangium grows and produces a new sporangium by prolifera-
tion. Very much resembling this genus is the alga Chytridiochloris, based
on H. viride Scherf., which has a chloroplast and reproduces in the same
manner. For this reason the correctness of the assignment of Harpo-
chytrium to this family is doubted by some authors, especially by Jane
(1946) who has made a monographic revision of the genus.

Because of its operculate manner of dehiscence Sparrow (1943) places
the genus Chytridium in a separate family, Chytridiaceae, while Miss
Whiffen includes it and other operculate forms of the same general struc-
ture in the Rhizidiaceae. The original encysted zoospore becomes the
sporangium. At its base is the endobiotic haustorium or rhizoidal system
which may produce an apophysis in some species. This haustorial system
may be a simple unbranched peg or a typical branched system of rhizoids
may arise from the tip of the peg, from the base of the sporangium, or
from the apophysis. The operculum is apical. Resting spores are endo-
biotic, apparently asexually produced. They act as prosporangia, giving
rise to extramatrical operculate sporangia. Sparrow recognizes twenty-
seven species besides a number of doubtful ones. They are mostly para-
sitic in fresh-water algae but a few occur in marine algae. They are re-
ported from Europe, Asia, and North America.

Family Entophlyctaceae. In this family the zoospore cyst becomes
emptied and usually soon disappears or remains as an empty cap. The
fungus body enlarges within the host and sends out rhizoids which in
some species are very extensive, over 0.5 mm. long and up to 10 m thick.
Some forms are strictly parasitic, mostly upon algae, but many are
saprophytes on various types of material. In Entophlyctis the germ tube
enlarges to become the sporangium within the cell of the algal host. The
usually single exit papilla is inoperculate. Resting spores are known in

60 PHYCOMYCETEAE

some species. In E. vaucheriae (Fisch) Fischer the latter acts as a prospo-
rangium, giving rise to a thin-walled spherical sporangium. Sexual repro-
duction is not known. In Endochytrium the exit tubes are operculate,
otherwise the fungus is like Entophlyctis. In Diplophlydis the germ tube
enlarges as in the two foregoing genera but this enlargement is a prospo-
rangium from which grows out an external inoperculate sporangium. The
resting spores in D. intestina (Schenk) Schroet. are formed, according to
Sparrow (1936), by the anastomosis of the rhizoidal systems of two plants,
the contents of the smaller passing into the larger which becomes thick-
walled and covered externally by minute short sharp spines. They may
function directly as zoosporangia or as prosporangia. In Phlyctorhiza
(Hanson, 1946) the rhizoids radiate from the germ tube, branching freely
and anastomosing to form a reticulum. The proximal portions of these
rhizoid branches form, at first, a thin, angular sporangium which even-
tually becomes a round, shallow^, somewhat lenticular, thin-walled zoospo-
rangium lying underneath the radiating branches. Through a low,
inoperculate papilla the zoospores escape into a vesicle where they soon
become active and swim away. Thick-walled somewhat tuberculate
resting spores may be formed, apparently asexually, in place of the
zoosporangia. In germination they act as prosporangia. Rarely some of
the rhizoid branches produce secondary zoosporangia so that such indi-
viduals are polycentric. The only known species, P. endogena Hanson,
grows in the basement membrane of insect integuments.

Nephrochytrium, saprophytic mostly on algae or grass leaves in water,
corresponds to Diplophlydis except that the exit tubes are operculate.
Whether its resting spores are produced by a sexual process or not is
unknown.

Macrochytrium may possibly belong in the Entophlyctaceae but needs
further study. The zoospore gives rise to a stout germ tube which produces
a cluster of coarse rhizoids. Laterally from the germ tube, near its point
of origin, arises the subspherical sporangium which may attain a diameter
of 0.5 mm. It opens at the apex by a large operculum allowing the escape
of up to a thousand zoospores. This has been found in Europe and in the
United States, saprophytic on submerged fruits or twigs. Whether this is
a typical member of the Chytridiales or not is uncertain. The coarse
rhizoidal system suggests the possibility of a coenocytic mycelial structure
with many nuclei instead of an enucleate system.

The polycentric, eucarpic Chytridiales are placed by some students
in one, two, or three families. Until their life histories and cytology are
known more completely any classification of these genera will have to be
tentative. The author recognizes two families as well distinguished, with
the probability that more may have to be recognized when the life
histories of the members of the group are better known.

ORDER CHYTRIDIALES 61

The dozen or so genera of Chytridiales that are polycentric differ from
the eucarpic, monocentric genera in the production of many nucleate
expansions on the rhizoidal system, each of which may become a center
for the production of zoosporangia, resting spores or other rhizoidal
branches. As a result the fully developed individual consists of many
centers of varying size and shape, connected by (usually) slender, non-
nucleate rhizoidal strands or tubes. This system of tubes and rhizoids
was given the name "rhizomycelium" by Karling (1932). One of the
crucial points in the recognition of the affinity of these organisms is to
determine whether they possess a true mycelium, with nuclei along its
course, or a rhizomycelium. Furthermore, does this distinction necessarily
separate these fungi into different orders or do we have here a gradual
gradation within the same order from organisms scarcely distinguishable
from the Rhizidiaceae or Entophlyctaceae to those in which the rhizo-
mycelium has become truly mycelial?

The two families tentatively accepted by the author are as follows:

Cladochytriaceae : intra- or extramatrical, eucarpic, with many centers connected
by slender, branching, rhizoidal or tubular threads, with swellings here and
there. Sporangia thin-walled, terminal or intercalary, operculate or in-
operculate. Resting spores thick-walled, apparently not sexually produced.

Physodermataceae : epibiotic, monocentric, and eucarpic at first, then by inde-
pendent infection endobiotic and strongly polycentric, producing resting
spores only. The latter eventually produce internally one to several thin-
walled, inoperculate zoosporangia. Probably the epibiotic stage produces
gametes while the resting spore sporangia produce zoospores but this has
not been proved.

Family Cladochytriaceae. In the Cladochytriaceae the chief genus
with inoperculate sporangia is Cladochytrium, which is saprophytic and
forms a system of fine branching rhizoids within the tissues of the sub-
stratum. Here and there arise swellings which may become the round
or pyriform sporangia or which may become spindle-shaped, once septate
"turbinate organs," one of these cells becoming a sporangium and per-
haps later the other. Septa are formed to separate the sporangia and
resting cells from the remainder of the rhizomycelium. The zoosporangia
empty through an inoperculate exit tube which may be quite long. Some
of the swellings become colorless resting spores with a thickened wall.
These, according to Karling (1934), germinate by becoming sporangia in
which numerous zoospores are produced by cleavage and escape through
a short sporangial neck, in C. replicatum Karl. The cytology of this fungus
has been studied by Karling (1937). (Fig. 17A-C.)

Physocladia (Sparrow, 1932) differs in habit from the foregoing in
that it is external to the matrix (pine pollen in the case of P. ohscura
Sparrow) which is penetrated only by the fine tips of the rhizomycelium.
It consists of a slender, branched thread with rhizoidal branches and

Fig. 17. (See legend on lacing page.)
62

ORBEK CHYTRIDIALES 63

intercalary swellings here and there and, rarely, with septate turbinate
cells, and with large terminal spherical zoosporangia, each with an
apophysis. These produce large numbers of zoospores which escape
through a nonoperculate opening and swim actively for some time in a
large vesicle before its membrane is ruptured. Large spherical, thick-
walled resting spores are formed terminally. Their further fate has not
been determined.

Polychytrium (Ajello, 1942) differs from both the foregoing genera in
its coarse branched rhizomycelium which has occasional rhizoids but no
turbinate cells or swellings. At the ends of the branches and sometimes in
intercalary positions the zoosporangia arise, usually in pairs, sometimes
several close together. They are pyriform. In them the zoospores are
developed and push out through the apical exit papilla or tube surrounded
by a slimy matrix in which at first they lie motionless. Soon they become
active and break away and swim off. They have a long posterior flagellum
and a conspicuous lunate nuclear cap. After the discharge of the zoospores
new zoosporangia are formed by proliferation. In addition to these typical
zoosporangia, yellowish-brown very tuberculate zoosporangia may occur,
in pairs or in clusters. From these the zoospores escape in the same
manner. No resting spores have been observed.

The principal operculate genera of this family are Nowakowskiella,
Septochytrium, Megachytrium, and Catenomyces. In Nowakowskiella the
slender rhizomycelium is nonseptate, except where the enlargements be-
come zoosporangia, with usually a large apophysis. Zoospores escape in a
mass before separating or those few remaining behind creep out one by
one. The zoosporangia sometimes proliferate. Resting spores are formed
in some species of the genus. Roberts (1948) shows that in some species
of Nowakowskiella there is a marked morphological differentiation be-
tween the vegetative (or trophic) and the reproductive portions of the
rhizomycelium. The former is mostly confined to the interior of the
substratum and produces no reproductive organs from the enlargements.
The nuclei are found only in the swellings but not in the isthmuses or
rhizoids. In the reproductive portion, which is mostly external to the
substratum, the nuclei occur not only in the enlargements, which may
eventually become the zoosporangia, but also in the intervening fila-
mentous structures. In Septochytrium the primary swelling becomes a

Fig. 17. Chytridiales, Family Cladochytriaceae. (A-C) Cladochytrium tenue Now.
(A) General view of part of the thalhis, showing spindle organs, zoosporangia in vari-
ous stages of development, and rhizomycelium. (B) Resting spore which developed in
a spindle organ. (C) Resting spore germinating to produce external zoosporangium.
(D-F) Nowakowskiella macrospora Karl. (D) General view of part of the thallus, show-
ing rhizomycelium and swellings and operculate zoosporangia. (E) Resting spore. (F)
Resting spore germinating to produce external zoosporangium. (Courtesy, Karling:
Am. J. Botany, 32(l):29-35.)

PHYCOMYCETEAE

Fig. 18. Chytridiales, Family Cladochytriaceae. Catenomyces persicinus Hanson.
Portion of plant showing coarse rhizomycelium and zoosporangia with endo-operculate
exit tubes. (Courtesy, Hanson: A7n. J. Botany, 32(7):431-438.)

large sporangium from whose sides arise many moderately stout rhizoidal
tubes with occasional constrictions and septa, and fusiform swellings,
some of which may become large or small zoosporangia or resting spores.
The latter act as prosporangia when they germinate. In S. variabile
Berdan, Miss Berdan (1939) demonstrated that the sporangial walls and
intercalary swellings give a pronounced violet color on treatment with
chloriodide of zinc, indicating the presence of cellulose. This reaction is
absent in most of the members of the family. M egachytrium differs from
the two foregoing genera in the possession of a very coarse, tubular,
branched rhizomycelium instead of fine tapering threads. The swellings
of this rhizomycelium become operculate sporangia or resting spores. The
latter act as prosporangia when they germinate. The genus Catenomyces
(Hanson, 1945) is quite similar to the three foregoing genera but has an
endo-operculum deep down in the exit tube, not at its apex. Resting
spores have been doubtfully observed. The zoosporangia may be terminal
or intercalary, with one to several exit tubes. The rhizomycelium is stout
but tapers to slender rhizoid-like tufts. (Fig. 17D-F, Fig. 18.)

The inoperculate genus Catenaria, assigned by Sparrow (1943) to the
subfamily Catenarioideae, has been shown by Couch (1945) to be proba-
bly more closely related to Order Blastocladiales and will accordingly be
discussed in the next chapter.

Family Physodermataceae. Two genera, Physoderma and Uro-
phlyctis are usually included in this family, but since the only constant
difference is the effect upon the host the validity of this distinction is
doubtful. Both are parasitic on and in higher plants. Physoderma may
discolor and eventually kill the infected tissues without causing marked
hypertrophy, while Urophlyctis induces strong gall formation. Tentatively

ORDER CHYTRIDIALES

Fig. 19. Chytridiales, Family Physodermataceae. Physoderma zeae-maydis Shaw.
(A) Resting sporangium, side view. (B) Resting sporangium germinating to form
zoosporangium. (C) Young ephemeral zoosporangia. (D) Emptied ephemeral zoo-
sporangia, one with beginning of proliferation. (E) Swarm spore from ephemeral
zoosporangium. (F) Swarm spore from resting spore sporangium. (Courtesy, Sparrow:
Am. J. Botany, 34(2):94-97.)

the genera are kept distinct, especially since phytopathological literature
maintains this separation. They differ from the foregoing polycentric
forms in that the primary infection produces an external sporangium with
rhizoids penetrating the epidermal cells of the host. This rather flattened,
or even slipper-shaped, zoosporangium opens without an operculum and
sets free numerous swarm cells. Successive sporangia may arise by
proliferation within the empty walls. The polycentric stage is originated
in a manner characteristic of the Cladochytriaceae, probably by infection
from these swarm cells

In Physoderma macular e Wallr., Clinton (1902) has shown that the
germinating zoospore sends rhizoids into the host cell (submerged leaf of
Alisma) and enlarges externally to become a zoosporangium much in the
manner of Rhizophydium. After the zoospores escape through an exit
papilla a second zoosporangium may be formed within the empty wall of
the first, and so on two to four times. Other zoospores settling on older
leaves, or perhaps upon leaves not permanently emersed, send in a fine
filament which enlarges to become a storage cell (or "Sammelzelle") and
produces haustorial processes or fine filaments which may penetrate to
other cells and there in their turn produce similar cells, and so on. From
each such cell there may arise a bud which eventually becomes larger
than the original cell. This is a resting sporangium and is thick-walled
and somewhat flattened on one side. After a period of rest it enlarges and

66 PHTCOMYCETEAE

bursts off the upper side of the thick wall with a circular split, forming a
very large lid. On the thin inner wall an inoperculate exit papilla is
formed, setting free the posteriorly uniflagellate zoospores. The suggestion
may be made that possibly the externally formed zoospores are capable
of functioning as gametes, the external zoosporangia arising from zoospore
infection, the internal infection by zygotes. Physoderma zeae-maydis Shaw
is sometimes destructive to corn (maize, Zea mays L.) in the southern
parts of the United States and in Asia. Usually only the internal rhizo-
mycelium and the "Sammelzellen" and resting sporangia are observed
(Tisdale 1919) but Sparrow (1934, 1947) has shown that it is possible to
obtain the production of the external slipper-shaped sporangia by placing
pieces of young maize leaves in a hanging-drop culture with zoospores
from the resting sporangia. As in P. maculare new zoosporangia are
formed within the emptied older ones. These planospores are markedly
smaller than those arising from the resting sporangia and Sparrow sug-
gests that they may possibly be gametes. (Fig. 19.)

Jones and Drechsler (1920) and also Bartlett (1926) have shown that
Urophlyctis has much the same life history as Physoderma but causes
extensive gall production by the host, this being practically the only
distinction between the two genera. The external zoosporangia have been
reported in several species of Urophlyctis. Within the infected epidermal
cell there develops a " Sammelzelle " or "turbinate cell," at first uni-
nucleate but soon multinucleate. At its distal end is formed a terminal
tuft of haustoria. At several places on the cell, buds are formed into each
of which a nucleus passes, following which the bud grows out as a some-
what enlarged end of a very slender non-nucleate filament. This enlarge-
ment in turn becomes a turbinate cell and may give rise to other similar
cells, usually three to five cells from each. In the center of the distal tuft
of haustoria there soon buds out a thin-walled cell which grows rapidly
and becomes much larger than the cell from which it originates. This
becomes thick-walled and bears a crown of haustoria. During its growth
the cytoplasm and nuclei from the turbinate cell pass into it. This is not
an act of fertilization, as was believed by earlier mycologists. The con-
necting rhizomycelium soon disappears and finally only the resting
sporangia are to be found in the gall tissue. After some time these spo-
rangia are capable of germination. Scott (1920) has studied this process
in U. alfalfae Magn. It produces one to fifteen or more zoosporangia
varying in diameter from 10 to 40 microns, which push out through
irregular fissures in the brown wall. Zoospores escape through short exit
papillae, there being several such papillae on the larger sporangia. The
zoospores are 4 to 8 microns long with a posterior flagellum 30 to 50
microns in length. Scott observed no conjugation of these zoospores. On
the other hand, 0. T. Wilson (1920) reports that the zoospores are

ORDER CHYTRIDIALES

Fig. 20. Chytridiales, Family Physodermataceae. Urophlyctis alfalfae Magn. (A)
Rhizomycelium, "Sammelzellen," and resting sporangia. (B) Top view of resting
sporangium, (After Jones and Drechsler: J. Agr. Research, 20(4):295-324.)

bifiagellate, one flagellum being very short, and that they are of two
sizes. These conjugate before infection takes place. This needs confirma-
tion as there is a possibiHty that these supposed zoospores or gametes
may have been organisms parasitic within the resting sporangia of the
Urophlyctis. Several species of Urophlyctis are known. (F'ig. 20.)

As a tentative addition to the operculate Chytridiaceae, Tetrachytrium
and Zygochytrium, reported by Sorokin (1874) from Russia may be con-
sidered here for want of further information. They are soil or water fungi,

PHYCOMYCETEAE

FiG.[21. (See legend on facing page.)

ORDER HYPHOCHYTRIALES G9

less than 0.1 mm. tall, and consist of an upright, stout and few-branched,
nonseptate hypha with a basal holdfast. Whether this is a true hypha or
a rhizomycelium is not known, since neither species has been found again
since their original description.

In Zygochytrium the main stalk is forked, each branch terminating in
a spherical, operculate sporangium. The contents escape as a naked mass
which soon forms a thin membrane. In the interior the protoplasm divides
into many posteriorly uniflagellate zoospores which escape by the irregu-
lar rupture of the enclosing membrane, and soon germinate. Between the
main forks of the fungus short branches grow which meet and unite,
forming a zygospore in a manner very suggestive of zygospore formation
by the Mucorales. This thick-walled spore germinates soon by a slender
hypha and produces a new plant resembling the parent plant. This fungus
is yellow and grows on dead, submerged insects. (Fig. 21A-F.)

In Tetrachytrium, which is blue-green in color, the nonseptate main
stalk bears three or four somewhat recurved branches on some of which
develop terminally spherical, operculate gametangia. Their contents
escape in the same manner as from the zoosporangia of Zygochytrium
and in a similar manner become invested in a thin membrane. Within
this are formed four posteriorly uniflagellate gametes which unite by
twos to form thin-walled, nonflagellate zygospores. After a little while
these germinate to form new plants. The gametes which do not unite do
not germinate. Occurring on dead plant material and even on dead
beetles in water. (Fig. 21G-M.)

Until these fungi can be found again it will be impossible to decide
whether they really belong to the Chytridiales or to a group of fungi that
have a true mycelium. Their relationship may be with the operculate
Cladochytriaceae.

Order Hyphochytriales. The organisms making up this order show a
close parallelism in body structure to the Chytridiales with which they
have been usually associated. They differ, however, in the type of zoospore
which is anteriorly uniflagellate. In Rhizidiomyces apophysatus Zopf,
Couch (1941) demonstrated that this single anterior flagellum is of the
tinsel type in contrast to the posterior flagellum of the Chytridiales which
is of the whiplash type. The cell walls may show positive cellulose reaction
in a few cases but in some species chloriodide of zinc calls forth no charac-
teristic cellulose coloring. Karling (1943) has grouped these forms with a

Fig. 21. Chytridiales. (A-F) Zygochytrium aurantiacum Sor. (A) Mature plant.
(B) Emptied zoosporangium, showing operculum. (C) Extruded protoplasm has
formed zoospores. (D) Zoospores escaping. (E) Plant with beginning of conjugation.
(F) Mature zygospore. (G-M) Tetrachytrium triceps Sor. (G) Mature plant. (H) Proto-
plasm escaping from gametangium. (I) Four planogametes formed in the extruded
protoplasm. (J) Gametes escaping. (K) Gametes uniting. (L) Zygote beginning to
germinate. (M) Young plant. (After Sorokin: Botan. Ztg., 32(20) :305-315.)

PHYCOMYCETEAE

Fig. 22. Hyphochytriales. (A-C) Family Anisolpidiaceae. Anisolpidium edocarpii
Karliiig. (A) Zoosporangium with unopened exit tube. (B) Zoosporangium discharg-
ing the anteriorly uniflagellate zoospores. (C) Portion of filament of Ectocarpus
containing one empty zoosporangium and one resting spore. (D) Family Rhizi-
diomycetaceae. Rhizidiomyces apophysatus Zopf. (E) Family Hyj^hochytriaceae.
Hyphochytriuni catenoides Karling. (A-C, courtesy, Karling: Am. J. Botany, 30(8) :637-
648. D-E, courtesy, Sparrow : Aquatic Phycomycetes, Exclusive of the Saprolegniaceae
and Pythium, Ann Arbor, Univ. Michigan Press.)

single anterior flagellum into three families, emplo3ang the ordinal name
Anisochytridiales. They may be distinguished as follows:

Anisolpidiaceae: holocarpic, intramatrical, monocentric, the thallus becoming
a sporangium or a resting spore.
Three genera: Anisolpidium., Reesia, and Cystochytrium.
Rhizidiomycetaceae : eucarpic, monocentric, sporangia and resting spores
extramatrical, rhizoidal system intramatrical.
Two genera : Rhizidiomyces and Latrostium.

KEY TO THE MORE IMPORTANT FAMILIES AND GENERA 71

Hyphochytriaceae' : polycentric, intramatrical, forming a more or less branched
mycelium-like structure which here and there, terminally as well as in
intercalary positions, enlarges to form the sporangia or resting spores.
Two genera: Hyphochytriwn and Catenariopsis.

Family Anisolpidiaceae. Anisolpidiwn resembles Olpidium very
closely in its life history except that sexuality has not been observed.
The pyriform zoospore possesses a long, anteriorly attached flagellum.
Two or three species parasitic in marine Phaeophyceae. In A. ectocarpii
Karling, the chromosome number is five or six in the dividing nuclei in
the sporangium. Reesia differs from the foregoing genus in the persistence
of the naked, amoeboid stage until the organism has attained almost full
size when a cell wall is produced. In R. amoeboides Fisch, the resting
spores arise from infection by bifiagellate zygotes formed by the union
of two swarm spores. Two species, parasitic in Lemna. (Fig. 22A-C.)

Family Rhizidiomycetaceae. Rhizidiomyces resembles Rhizophydium
(or Phlyctochytrium, in one species, because of the presence of an apophy-
sis), but the zoospore is anteriorly uniflagellate, with the tinsel type of
flagellum. Four species are known, two saprophytic in soil, one parasitic
in algae, and one parasitic in the oogonia of Saprolegniaceae. Latrostium,
parasitic in the oogonia of Vaucheria, possesses only one recognized
species. The zoospores escape not through an exit tube as in Rhizidiomyces
but by the deliquescence of a large exit papilla. (Fig. 22D.)

Family Hyphochytriaceae. Hyphochytrium infestans Zopf was first
described by its author in 1884 and has not been recognized since. It was
parasitic in the apothecium of a species of Helotium. It consists of an
extensive, coarse, tubular, branched, occasionally septate rhizomycelium
(or true mycelium?). The zoosporangia arise as terminal and intercalary
swellings on the mycelium. They open by a subapical orifice releasing the
anteriorly uniflagellate zoospores. No resting spores have been observed.
Two other species are known: H. hydrodidii Valkanov on Hydrodictyon
and H. catenoides Karling, mostly saprophytic. The former produces
resting spores. Catenariopsis possibly belongs in the foregoing genus. Its
difference appears to be a greater contrast between the enlarged zoospo-
rangia and the short connecting isthmuses. (Fig. 22E.)

Key to the More Important Families and Genera of Order Chytridiales

Holocarpic, i.e., endobiotic, without rhizoids, discharge tubes inoperculate.
Vegetative cell enlarging to form a single zoosporangium.

Family Olpidiaceae
Zoosporangium not completely filling the host cell.

Flagellum anteriorly attached but traihng posteriorly. (Doubtfully be-
longing in this family.) Sphaerita

1 This family as here delimited differs greatly from the treatment accorded in the
first edition of this textbook, both as to its suggested relationship and contents.

72 PHYCOMYCETEAE

Flagellum posterior, discharge tubes rarely more than one.

Olpidium
Flagellum posterior, discharge tubes usually numerous. Pleotrachelus
Zoosporangium completely filling the host cell, the cell walls of the fungus
and host in close contact. RozeUa

Vegetative cell elongating and by septation forming a series of zoosporangia.

Family Achlyogetonaceae
More than two zoosporangia formed in a series.

Zoospores encysting at mouth of exit tube. Achlyogeton

Zoospores swimming away from exit tube without encysting.

Septolpidium
Only two zoosporangia formed, separated by an isthmus.

Bicricmm
Vegetative cell dividing internally into numerous zoosporangia or serving as
a prosorus, the zoosporangia arising in a cell developing
from the prosorus. Family Synchytriaceae

Parasitic in higher plants; vegetative cell rather large, becoming a sorus or

prosorus or resting spore. Synchytrium

Parasitic in algae; vegetative cell small, becoming a prosorus or a resting
spore.
Sorus of zoosporangia formed within the host cell.

Sorus dividing into a few sporangia which show no signs of flagella, each

producing a number of zoospores. Micromyces

Sorus dividing into many zoosporangia which are set free, part within the
host cell, part through a papilla into the surrounding water.
Occasionally a zoosporangium bears a feebly active pos-
terior flagellum. Zoospores minute, 2-5. Endodesmidium
Sorus of zoosporangia formed outside the host cell at apex of a discharge
tube which pierces the host cell wall and produces only a
few nonmotile zoosporangia. Micromycopsis

Eucarpic, i.e., endobiotic, epibiotic, or interbiotic, with rhizoids or haustoria.

Monocentric. Including both operculate and inoperculate
genera.
Zoospore cyst enlarging into a zoosporangium or prosporangium.

Family Rhizidiaceae
Zoospore cyst enlarging into a zoosporangium.
Exit papilla or tube inoperculate.

Subsporangial swelling (apophysis) lacking.

Interbiotic (i.e., not closely attached to host cell, which is entered
only by the tips of the rhizoidal branches).
Rhizoidal system arising from the main axis. Rhizidium

Rhizoidal system with several branches from the body of the
sporangium. Rhizophhjdis

Epibiotic (i.e., closely associated with the host cell which is pene-
trated by the rhizoidal system). Rhizophydium
Subsporangial swelling present (scarcely differentiated in Ohelidium).
Ej^ibiotic. i Phlyctochytrium
Interbiotic.

Apex of zoosporangium with mucro; a cup- or funnel-like base.

Obelidium
No pronounced mucro; no cup- or funnel-like base.

Zoosporangium somewhat stellate. Asterophlyctis

KEY TO THE MORE IMPORTANT FAMILIES AND GENERA 73

Zoosporangium spherical or ellipsoidal.

Rhizoids delicate. Rhizodosmatium

Rhizoids coarse." Siphonaria

Exit tube operculate near base, interbiotic. Karlingia

Exit tube operculate at apex, epibiotic. Chytridium

Zoospore cyst enlarging into a prosporangium.

Epibiotic, operculate, small, rhizoids not massive. Chytridium

Epibiotic, operculate, rhizoidal system massive. Macrochytrium

Interbiotic, inoperculate. Polyphagus

Zoospore cyst not enlarging into zoosporangium or prosporangium, but upper
part of germ tube so enlarging.

Family Entophlyctaceae
Exit tube or papilla inoperculate. Endobiotic.

Upper part of germ tube becoming a zoosporangium. Entophlydis
Zoosporangia arising by enlargement of the proximal portions of the

rhizoidal branches. Phlyctorhiza

Upper part of germ tube becoming a prosporangium. Diplophlyctis
Exit tube operculate, germ tube enlarging into a zoosporangium.
Epibiotic. Rhizoids constricted into catenulate segments.

Catenochytrium
Endobiotic. Rhizoids not constricted. Endochytrium

Exit tube operculate. Endobiotic. Germ tube enlarging into a prosporangium.
Eucarpic and polycentric.

Nephrochytrium

Mostly saprophytic, intra- and extramatrical, with many centers connected

by slender (rarely stout) tubes. Sporangia thin-walled,

terminal and often intercalary. Resting spores thick-walled,

apparently not sexually produced.

Family Cladochytriaceae
Sporangial openings inoperculate.

Rhizomycelium slender, branching, intramatrical, with turbinate cells.
Zoosporangia intercalary and terminal. Resting spores thick-
walled, acting as prosporangia when germinating.

Cladodiytrium
Rhizomycelium slender, branching, mostly extramatrical, rarely with
turbinate cells. Zoosporangia and resting spores single at the
apex of hyphae or branches. On pine pollen in water.

Physodadia
Rhizomycelium coarse, branched, without turbinate cells or intercalary
swellings. Zoosporangia usually in clusters of two or more
at tips of hyphae or intercalary. No resting spores although
thicker-walled, tuberculate zoosporangia are formed which
set free their zoospores promptly. Polychytrium

Sporangia operculate at the mouth of the opening.

Intra- and extramatrical rhizomycelium slender and branched, with
various more or less fusoid swellings. Zoosporangia formed
mostly on the extramatrical rhizomycelium, terminal or
intercalary, apophysate. Thicker-walled resting spores some-
times formed from a mass of parenchyma-like cells. Zoospo-
rangia sometimes proliferating. Nowakowskiella
Mainly intramatrical, rhizomycelium moderately stout, constricted occa-
sionally and septate or partially so at the constrictions.

74 PHYCOMYCETEAE

branching and tapering to fine points. Numerous fusiform
swellings which may become large or small zoosporangia or
resting spores, the latter acting as prosporangia when they
germinate. Zoosporangia not proliferating.

Septochytrium
Extra- and intramatrical, rhizomycelium rather thick and undulate,
branched, not tapering into fine points. Zoosporangia ter-
minal and intercalary, rarely proliferating. Resting spores
intercalary, on germination acting as prosporangia.

Megachytrium
Exit tubes of zoosporangia operculate near base (endo-operculate). Rhizo-
mycelium moderately stout. Catenomyces
Parasitic epibiotically and endobiotically on higher plants, the epibiotic stage
monocentric, consisting of a proliferating, slipper-like
zoosporangium with a tuft of rhizoids in the epidermal cell,
the endobiotic stage being a slender branching rhizomy-
celium with numerous centers which give rise to further
rhizomycelium and to thick-walled resting sporangia, mostly
flattened on one side. Family Physodermataceae
Endobiotic stage not causing warts or galls on the host.

Physoderma
Endobiotic stage causing warts or galls. Urophlydis

Appendix to the Operculate Cladochytriaceae, but with relationship doubtful.
Stalk forked and bearing terminally the operculate zoosporangia. Sexual re-
production by formation of thick-walled zygospore by a
mucoroid process of conjugation. Zygochytrium

Stalk branched and bearing at its tips operculate gametangia producing 4
gametes in each. No thick-walled resting spores known.

Tetrachytrium

Key to the Families and Genera of Order Hyphochjrtriales

{Based upon Karling, 1943)

Holocarpic, monocentric, endobiotic. Family Anisolpidiaceae

Vegetative stage provided early with a cell wall and becoming a round or
ellipsoidal zoosporangium with 1-3 exit tubes. Sexual reproduction unknown.
Parasitic in Phaeophyceae. Anisolpidiuni

Vegetative stage naked and amoeboid almost until maturity of the rounded
zoosporangium. Exit tube single. Sexual reproduction by the union of uni-
flagellate isogametes. Parasitic in Lemnaceae. Reesia
Vegetative structure becoming elongated, sometimes septate, to form one
elongated or several rounded zoosporangia with thick walls. Zoospores
escaping through a median aperture. Sexual reproduction unknown. Para-
sitic in roots. Cystochytrium
Eucarpic, monocentric, epibiotic. Family Rhizidiomycetaceae
Zoosporangia with or without apophysis, zoospores undergoing cleavage within
a vesicle at the apex of the exit tube. Parasites or saprophytes.

Rhizidiomyces
Zoosporangia without apophysis, zoospores fully developed within the
zoosporangium. Parasites in oospores of Vaucheria. Latrostium

Eucarpic, polycentric, hypha-likc, without rhizoids, with terminal and inter-
calary zoosporangia. Family Hyphochytriaceae

LITERATURE CITED 75

Only one well established genus. Parasitic in various hosts.

Hyphochytrium
The genus Catenariopsis may be distinct from Hyphochytrium. In it the zoo-
sporangia are separated from one another by short isthmuses.

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Scott, C. Emlen: A preliminary note on the germination of Urophlyctis alfalfae,

Science, N.S., 52(1340) :225-226. 1920.
SoROKiN, N.: Einige neue Wasserpilze, Botan. Ztg., 32:305-315. PI. 6. 1874.
Sparrow, Jr., Frederick K. : Observations on the aquatic fungi of Cold Spring

Harbor, Mycologia, 24(3) :268-303. Pis. 7-8. Figs. 1-4. 1932.
: The occurrence of true sporangia in the Physoderma disease of corn.

Science, N.S., 79(2060) :o63-564. 1934.
: Evidences for the possible occurrence of sexuality in Diplophlyctis,

Mycologia, 28(4) :321-323. Figs. 1-2. 1936.
: Some chytiidiaceous inhabitants of submerged insect exuviae, Proc.

Am. Phil. Soc, 78(l):23-53. Pis. 1-4. Figs. 1-5. 1937.
: The morphology and development of Obelidium mucronatum, Mycologia,

30(1) A-U. Figs. 1-44. 1938.
: A classification of aquatic Phycomycetes, ibid., 34(1):113-116. 1942.

LITERATURE CITED 77

Sparrow: Aquatic Phycomycetes, exclusive of the Saprolegniaceae and Pythium,
xix + 785 pp. Q34: figs. Ann Arbor, Univ. Michigan Press, 1943.

: Observations on chytridiaceous parasites of Phanerogams: II. A pre-
liminary study of the occurrence of ephemeral sporangia in the Physoderma
disease of maize, Am. J. Botany, 34(2):94-97. Figs. 1-17. 1947.

TiscHLER, G.: Pflanzliche Chromosomen-Zahlen, Tabulae Biologicae, 4:1-83.
1937.

TiSDALE, W. H.: Physoderma disease of corn, /. Agr. Research, 16(5):137-154.
Pis. A and B (colored) and 10-17. Fig. 1. 1919.

Wager, H. : The Ufe-history and cytology of Polyphagus Eugienae, Ann. Botany,
27(106) :173-202. PZs. 16-19. 1913.

VON Wettstein, Fritz: Das Vorkommen von Chitin und seine Verwertung als
systematisch-phylogenetisches Merkmal im Pflanzenreich, Sitz. her. Akad.
Wiss. Wien, Math, natunv. Klasse, Abt. I, 130(1) :3-20. 1921.

Whiffen, Alma J. : A discussion of taxonomic criteria in the Chytridiales, Far-
lowia, l(4):583-597. 1944.

Wilson, 0. T.: Crown-gall of alfalfa, Botan. Gaz., 70(l):51-68. Pis. 7-10. 1920.

ZoPF, Wilhelm: Zur Kenntnis der Phycomyceten : I. Zur Morphologie und
Biologic der Ancylisten und Chytridium, zugleich ein Beitrag zur Pliyto-
pathologie. Nova Acta Leopoldina, 47:141-236. Pis. 12-21. 1884.

PHYCOMYCETEAE: BLASTOCLADIALES
AND MONOBLEPHARIDALES

THE organisms treated in this chapter show in very many points a
close relationship to the Rhizidiaceae in the Chytridiales, but exhibit
a greater development of the vegetative structure and a greater com-
plexity of sexual reproduction in most of the cases where this is known.
Instead of being a single uninucleate cell with a non-nucleate haustorial
system which immediately becomes a sporangium by enlargement and
internal division into zoospores, the vegetative body is a multinucleate
clavate, cylindrical or spherical, or hypha-like structure which may be
simple or branched. True septa are wanting except to delimit injured
regions or sporangia or gametangia, but coarsely perforated pseudosepta
occur in Allomyces and "cellulin" plugs may occur at the constrictions
in Gonapodya. From this vegetative body arise one to many sporangia or
gametangia. These organisms are aquatic or perhaps more often inhabit
the soil. Most are saprophytic on vegetable material but the genus
Catenaria contains species which may grow parasitically, in nematodes,
in fluke eggs, and in the case of one species (Couch, 1945a) in the hyphae
of the aquatic fungi Allomyces and Blastocladiella. Coelomomyces is para-
sitic in the larvae of mosquitoes.

The zoospores and gametes, where motile, are as in the Chytridiales
posteriorly uniflagellate, with the whiplash type of flagellum. In a few
species in the genera Blastocladia and Monoblepharis biflagellate zoo-
spores have been observed occasionally. Cotner (1930a,b) demonstrated
that these are abnormal structures, being binucleate as well as biflagellate,
and that their occurrence is due to unfavorable conditions of develop-
ment, usually unfavorable temperature, so that the typical zoospore of
this whole group should be looked upon as being uniflagellate. The cell
walls do not give the cellulose reaction with chloriodide of zinc except
after treatment for some time with warm KOH or NaOH solutions which
apparently saponify fatty deposits in the cell walls. After such treatment

ORDER BLASTOCLADIALES 79

and thorough washing the wall in Monohlepharis, according to von Wett-
stein (1921) and to Harder (1937), shows a violet coloration with chlor-
iodide of zinc. The author obtained the same reaction with an unidentified
species of Blastocladia. Harder claims that in AUomyces and certain other
members of the family the cell wall consists fundamentally of chitin.
Nabel (1939) makes the same claim for AUomyces, Blastocladia, and
Blastocladiella. In Coelomomyces a cell wall has not been demonstrated
on the vegetative mycelium.

Order Blastocladiales. In the first edition of this textbook Blasto-
cladiales and the following order, Monoblepharidales, were included under
the latter name. Because of the slight differences in the structure of the
zoospores and of the type of sexual reproduction, where it is known, and
the production of resting sporangia the author now follows Sparrow (1943)
in recognizing two orders. In the Blastocladiales the posteriorly uni-
flagellate swarm cells (zoospores or gametes) usually possess a well-
marked "nuclear cap" attached mostly to the anterior surface of the
nucleus. In the Monoblepharidales the nuclear cap is not noticeable but
in the swimming cells numerous small granules are gathered at the apex
sometimes forming a sort of papilla.

In the Blastocladiales sexual reproduction, where known, is by the
union of two equal or unequal motile gametes to form a biflagellate zygote
which germinates without becoming a resting spore. In the Mono-
blepharidales the egg is nonflagellate and after fertilization forms a thick
wall to become a resting spore. In the Blastocladiales, in addition to the
thin-walled zoosporangia, there are found also thick-walled resting spo-
rangia which, in Blastocladiella and AUomyces, have a close connection
with the alternation of sporophytic and gametophytic generations — an
alternation that is unknown in the Monoblepharidales. These resting
sporangia are spherical, ovoid, or pyriform, with a thick outer wall often
perforated by numerous pits, and a thin inner wall. On germination the
outer wall cracks open and the expanding inner wall produces one or more
exit papillae from which the swarm spores emerge. These latter are called
"planonts" by Sparrow (1943).

Three families are tentatively recognized, Catenariaceae (Couch
1945a), Coelomomycetaceae (Couch, 1945b) and Blastocladiaceae. They
are distinguished as follows:

Family Catenariaceae. This family is parasitic in worms or fungi,
or saprophytic in various other plant or animal substrata. The plant body
at first is tubular, mostly unbranched, coenocytic, swelling at more or less
regular intervals to form reproductive organs which are connected by
short, narrow isthmuses, septate at each end and sometimes in the
middle. These reproductive bodies are either thin-walled zoosporangia
discharging by exit tubes or, on more exhausted media, thick-walled

^ i ^'-1

"'•■■■■• IV^

s '-5

■?a'/

.*-_; 4>

■ n -'■- i'

V^i*:»-^

1" ff^^n

S ■*' .'•'«■

L, , ' «■

n'iH

n>g.6? '^

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^'•.•'^■■■f .

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Fig. 23. (See legend on facing page.)
80

ORDER BLASTOCLADIALES 81

resting spores usually free from the hyphal membrane. These resting
sporangia are smooth or minutely granular. In germination the outer
wall cracks open and a tube emerges through which the zoospores are
discharged. Rhizoids are produced at almost any point on the thallus,
from the isthmuses as well as the reproductive swellings. Sexual repro-
duction occurs by the union of equal, motile gametes. One genus only is
known, Catenaria, which was formerly placed in the family Clado-
chytriaceae of the order Chytridiales until Couch (1945a) demon-
strated that the mode of reproduction is typical of some forms of the
Blastocladiales.

Of the two well-studied species of the genus, sexual reproduction is
unknown in C. anguillulae Sorokin. It produces both thin-walled and
resting zoosporangia which appear to produce exactly similar zoospores.
In C. allomycis Couch, the zoospores from the thin-walled zoosporangia
produce plants with either or both types of sporangia. Those from the
resting sporangia produce less active zoospores which encyst at or near
the mouth of the exit tube. After about two hours these cysts produce
four gametes, each, which escape through a short papilla and sooner or
later unite by pairs to form posteriorly biflagellate zygotes. These swim
for a time, the two flagella lying close together and being synchronous in
their motions. The zygotes eventually encyst and penetrate the host
tissue by means of a slender germ tube. Apparently, pairing of the gametes
is rare between those from any one cyst. This type of reproduction is
practically identical with that of Blastocladiella cystogena Couch and
Whiff en (1942) and comparable to that of Allomyces cystogenus Emerson
as interpreted by McCranie (1942). (Fig. 23A-G.)

Family Coelomomycetaceae. Coelomomycetaceae are parasitic in
the larvae of insects, chiefly mosquitoes. Mycelium coenocytic, non-
septate, lacking rhizoids and without demonstrable cell walls, branched
and somewhat anastomosing. Terminal portions of the short branches
enlarge and break free and develop resting spores surrounded by the
plasma membrane of the mycelium. These spores possess a thick outer,
colored wall, usually pitted or striate, and a thinner inner wall. Dehiscence
by a longitudinal slit. Zoospores posteriorly uniflagellate with an, at most,
imperfectly formed nuclear cap. No thin-walled sporangia nor sexual stage

Fig. 23. Blastocladiales. (A-G) Family Catenariaceae. (A-F) Catenaria allomycis
Couch. (A) Chain of zoosporangia within filament of Allomyces. (B) Zoosporangium
with zoospores about to escape. (C) Resting sporangia within host filament. (D)
Zoospores emerging from germinating resting sporangium and encysting. (E) Cysts at
mouth of exit tube of resting sporangium. (F) Biflagellate zygotes formed by union of
gametes emerging from the cysts. (G) Catenaria anguillulae Sorokin, fluke egg with four
resting sporangia and several emptied thin-walled zoosporangia. (H, I) Family Coelo-
momycetaceae. Coelomomyces lativittatus Couch and Dodge. (H) Germinating resting
sporangium. (I) Zoospores. (A-G, courtesy, Couch: Mycologia, 37(2):163-193. H-I,
courtesy, Couch and Dodge: J. Elisha Mitchell Sci. Soc, 63(l):69-79.)

82 PHTCOMTCETEAE

observed. Twelve or more species. Only genus known Coelomomyces
(Keilin, 1921; Couch, 1945b; Couch and Dodge, 1947). (Fig. 23H, I.)

Family Blastocladiaceae. The vegetative portion of the plants of
this family consists of a more or less extensive system of tapering and
branching rhizoids and of a globular or clavate external portion from
which may arise directly the sporangia and gametangia or a system of
branches on which these organs are borne. In Allomyces the hyphae are
more nearly uniform, without a broad basal portion, and are frequently
constricted here and there. At these points usually occur coarsely perfo-
rate "pseudosepta." Three or more genera are recognized in this family,
which may be distinguished as follows:

Blastocladia: plant body with a simple, lobed or branched basal portion, often
with secondary axes. Zoosporangia with a single apical exit papilla. Alter-
nation of sporophytes and gametophytes unknown.

Blastocladiella (including according to Couch and Whiffen (1942) Clavochytrium
and Sphaerocladia) : plant body a spherical vegetative structure bearing
rhizoids at the base or on all sides, or more or less clavate, becoming directly
a reproductive organ or bearing at its apex the single reproductive organ.
Alternation of generations demonstrated in some species. Discharge papillae
one or several.

Allomyces: plant body a cylindrical basal segment giving rise, usually di-
chotomously, to cylindrical, often constricted, branches with pseudosepta.
Discharge papillae usually more than one. Alternation of sporophytes and
gametophytes known in some species.

Blastocladiella. In Blastocladiella stuhenii Couch and Whiffen (1942)
the plant body of the sporophyte is spherical, with tapering, much-
branched rhizoids emerging at all sides. It bears a thin-walled zoospo-
rangium with one to several discharge tubes or a dark, thick-walled,
resting sporangium which produces swarm cells that give rise to indis-
tinguishable male or female gametophytes similar to, but smaller than,
the sporophytes. The emerging gametes are equal in size and cannot be
distinguished by color. They fuse in pairs and at once produce the
sporophytes. In B. variabilis Harder and Sorgel (1938) the sporophyte is
cyhndrical, bearing at its base much-branched rhizoids and at its apex
either a thin-walled zoosporangium or a thick-walled resting sporangium.
The zoospores from the former give rise to sporophytes again but those
from the latter (planonts) give rise to similar gametophytes, each with a
single clavate gametangium, orange-colored in the male and colorless in
the female gametophyte. The biflagellate zygote gives rise at once to a
sporophyte. Five other species are distinguished. In B. stomophila (Couch
and Cox) Couch and Whiffen only thin-walled zoosporangia have been
obser\'ed. In B. simplex Matthews and two other species both zoospo-
rangia and resting sporangia are produced, but the zoospores from the
latter reproduce sporophytes, no gametophytes having been recognized.

ORDER BLASTOCLADIALES

Fig. 24. Blastocladiales, Family Blastocladiaceae. (A-D) Blastocladiella cystogena
Couch and Whiffen. (A) Plant with mature resting sporangium. (B) Uniflagellate
zoospores, two greatly magnified, just after discharge from resting sporangium. (C)
Cysts, some still containing and some discharging gametes. (D) Biflagellate zygote.
(E) Blastocladiella asperosperma Couch and Whiffen, cluster of plants, some with rest-
ing sporangia and some with thin-walled sporangia. (F, G) Blastocladia pringsheimn
Reinsch. (F) Plant with thin-walled sporangia. (G) Resting sporangia. (A-E, cour-
tesy. Couch and Whiffen: Am. J. Botany, 29(7):163-193. F-G, courtesy, Thaxter:
Botan. Gaz., 21(2):45-52, Univ. Chicago Press.)

84 PHYCOMYCETEAE

In B. cystogena Couch and Whiffen the thin-walled zoosporangia are
wanting. The zoospores from the resting sporangia encyst almost immedi-
ately and soon produce four smaller gametes each, also posteriorly
uniflagellate, which fuse in pairs and germinate at once to produce the
original stage. When cultured on an agar medium, several other species
besides B. siuhenii are spherical, with rhizoids on all sides, but on other
media clavate or cylindrical with a septum setting off the sporangium or
gametangium. (Fig. 24A-E.)

Blastodadia. This genus is more complicated in its structure. The
basal segment, bearing at its base tapering branched rhizoids, may be
spherical (in B. glohosa Kanouse) and bear on its surface the cylindrical
zoosporangia and subspherical or ovoid resting sporangia, sometimes
interspersed with slender threads. On the other hand, it may be cylindrical
and branched into more or less dichotomously dividing cylindrical hyphae
which bear the sporangia of both types. Often the hyphae bear a spo-
rangium apically and then branch sympodially so that the successive
sporangia appear racemose in arrangement. Slender setae may also be
present. The commonest species is apparently B. pringsheimii Reinsch,
which varies from a large clavate structure bearing the zoosporangia and
resting spores and setae at its apex or which may branch as described
above. In what was apparently this species the author (1939) observed
the union of equal swarm cells whose subsequent fate could not be fol-
lowed and whose origin, whether from thin-walled zoosporangia or thick-
walled resting sporangia was not ascertained. Miss Blackwell (1940)
studying apparently the same species very intensively was unable to
confirm this observation. (Fig. 24F, G.)

Allomyces. About six species are recognized in this genus. They are
distinguished mainly by their reproductive structures and life cycles.
Vegetatively they consist of a basal cylindrical segment attached by
tapering branched rhizoids to the substratum, and branching dichoto-
mously or sympodially into gradually more slender hyphae, sometimes
forming a tangled mycelial mass. The branches are blunt at the tip and
show here and there the pseudosepta characteristic of the genus, there
often being a constriction at each pseudoseptum. These are sometimes
wheel-Uke in appearance with radiating rods separating a circle of more
or less triangular openings. At the ends of the branches, singly or some-
times in chains, are produced the cylindrical or somewhat oval, thin-
walled zoosporangia. By sympodial branching these may come to have
the appearance of a racemose arrangement. The zoospores escape by one
or several inoperculate openings. The same plants may also bear the
thick-walled resting sporangia which vary in shape from spherical to
ovoid or even lemon-shaped and with the brown outer wall deeply
punctate. The zoospores from the thin-walled zoosporangia produce the

ORDER BLASTOCLADIALES

Fig. 25. Blastocladiales, Family Blastocladiaceae. Allomyces arbusculus Butler. Life
cycle. (After Emerson. Courtesy, Sparrow: Aquatic Phycomycetes, Exclusive of the
Saprolegniaceae and Pythium, Ann Arbor, Univ. Michigan Press.)

same stage of development. Those from the thick-walled resting sporangia
behave differently according to the species. In the main the reproduction
is of two distinct types. In the section Euallomyces, represented typically
by A . javanicus Kniep and A . arbusculus Butler, the zoospores from the
thick-walled resting spores produce gametophytes of approximately the
same structure as the sporophytes described above. In place of the
sporangia, chains of gametangia are produced terminally on the branches.
There are usually two, rarely more, in each chain. These consist of a
smaller and a larger gametangium, the former being terminal in A. java-
nicus, the latter terminal in A. arbusculus. The smaller, male gametangium
is salmon-pink to orange in color, the larger, female gametangium is
colorless. The gametes emerge from one to three exit papillae. The male
gametes are smaller and faintly colored and about twice as numerous as
the colorless female gametes. They fuse promptly and the biflagellate
zygote germinates almost at once to produce the sporophyte. This type
of sexual reproduction and alternation of generations in this genus was
first reported by Kniep (1929, 1930) in A. javanicus. Hatch (1933) demon-
strated it in A. arbusculus. Since then Hatch (1935, 1938), Emerson
(1941), McCranie (1942), and Wolf (1941) have given the genus intensive

86 PHYCOMYCETEAE

study. Emerson (1938) demonstrated another type of reproduction in
several species which he placed in the section Cystogenes. This has been
studied by McCranie upon whose observations the following life cycle is
based. In A. cystogenus Emerson the sporophyte is as in section Euallo-
myces, except that the resting sporangia remain attached to the sporo-
phyte in the latter and are deciduous in the former. Emerson reported
that on germination of these resting sporangia, there emerged large
posteriorly biflagellate swarm cells which quickly encysted and then gave
rise to four posteriorly uniflagellate swarmers from each cyst. These pro-
duced the sporophytic generation. McCranie's description differs from
that of Emerson in that he reports the emergence of nonflagellate, some-
what amoeboid, spores from the resting sporangium. These encyst and
within them are formed four uniflagellate gametes equal in size, which
unite to form posteriorly biflagellate zygotes from which develop the
sporophytes. The fact that in Blastocladiella cyslogena Couch and Whiffen
the swarmers emerging from the cysts act as isogamous gametes would
seem to make McCranie's conclusions as to the sexual processes in
A. cystogenus nearer the truth than Emerson's. The resting sporangia of
Allomyces are able to survive in the dry state many months or even years.
The species of the genus are more often found in the warmer parts of the
world, such as the southern United States, Mexico, Central and South
America, West Indies, Southern Asia, East Indies, Africa, and southern
Europe. (Fig. 25.)

Order Monoblepharidales. The chief differences between this order
and the Blastocladiales are the fertilization of large, nonmotile eggs by
posteriorly uniflagellate sperms; the hypha-like mycelium, like that of
Allomyces but lacking the perforated pseudosepta of that genus; the
absence of thick-walled resting sporangia and of alternation of genera-
tions, with minor differences in the structure of the swarm cells. Of the
three genera here included in this order sexual reproduction is unknown
in Gonapodya and the correctness of the inclusion of that genus in the
order is doubtful. Possibly it should be placed in the foregoing order.
The two other genera, Monoblepharis and Monohlepharella, are closely
related. With the doubtful inclusion of Gonapodya but one family is
recognized. The genus Myrioblepharis has been placed in this order by
some investigators, but it is probable that this is not an independent
organism but some other aquatic fungus parasitized by a ciliate Pro-
tozoan. The only species, M. paradoxa, was described by Thaxter in
1895, and found once by von Minden in 1915. The papers of these authors
should be referred to.

Family Monoblepharidaceae. Gonapodya is a genus of two species,
growing saprophytically on vegetable matter in fresh water. When rose
fruits are placed in water for several weeks, this fungus frequently de-

OKDER MONOBLEPHARIDALES

velops in little tufts, usually covered by bacterial slime, on the surface
of the fruits. The mycelium is variously branched, arising from a tuft of
branching, tapering rhizoids. The external mycelium is constricted at

Fig. 26. Monoblepharidales, Family
Monoblepharidaceae. Gonapodya siliquae-
formis (Reinsch) Thaxt. (A) Portion of
fungus showing typical habit. (B) Unopened
zoosporangium with zoospores outlined. (C)
Zoosporangium with escaping zoospores.
(D) Zoospore. (After Thaxter: Botan. Gaz.,
20(ll):477-485, Univ. Chicago Press.)

frequent intervals producing cylindrical or moniliform coenocytic seg-
ments. At the constrictions there are usually plugs of a carbohydrate
substance that has been called cellulin. At the ends of the branches the
terminal segment becomes the zoosporangium, ovate or lanceolate in

PHYCOMYCETEAE

outline, and producing numerous zoospores of the same type as those of
Monoblepharis. After these have escaped through the apical pore suc-
cessive zoosporangia may be formed in the empty sporangial wall by
proliferation. Sexual reproduction similar to that of Monoblepharis was
reported by Cornu (1871) but has never been found again in either of the

two species now recognized. (Fig. 26.)

Monoblepharis, with two species, was
first described by the French mycologist
Cornu in 1871; since then four or five
other species have been recognized.
Lagerheim (1900) gave the name Di-
blepharis to two species of this genus in
which biflagellate swarm cells were ob-
served, but in view of the investigations
of Cotner referred to earlier, and since
the sperm cells in those species were
described as uniflagellate, it seems desir-
able to consider these as belonging to
Monoblepharis. The species of this genus
are saprophytic, usually on twigs or
other vegetable matter in fresh water.
They are attached to the substratum
by rhizoids and form unbranched or
branched hyphae which are nonseptate
except where reproductive organs are
formed. The zoosporangia are terminal
to the main hypha or its branches, and
are mostly more or less cylindrical. After
the sporangium is emptied another may
be formed within the empty wall by
proliferation, or terminally on a sympo-
dial branch arising at the base of the
old sporangium. The zoospores escape,
fully formed, from an opening dissolved
in the apex of the sporangium and are typically posteriorly uniflagellate.
As mentioned above the frequent biflagellate condition of the zoospores
is probably abnormal.

The oogone in Monoblepharis may be formed terminally or by the
enlargement of a subterminal segment. In the former case a second seg-
ment immediately below the oogone becomes the antherid; in the
latter case the terminal segment becomes the antherid. In M. macrandra
(Lagerheim) Woronin the antherids may be terminal on slender
branches distant from the oogones. The number of sperms produced in an

■'-H

Fig. 27. Monoblepharidales,
Family Monoblepharidaceae. Mon-
oblepharis insignis Thaxter. (A)
Hyphae showing antherids, young
oogones and oogones with endo-
genous oospore. (B) Oogone ready
for fertilization and antherids with
sperms. (After Thaxter: Botan.
(?a2., 20(10):433-440,Univ.Chicago
Press.)

OKDER MONOBLEPHARIDALES

Fig. 28. Monoblepharidales, Family Monoblepharidaceae, M onoblepharis poly-
morpha Cornu. (A) Zoospore. (B-E) Zoosporangia. (F) Fertilization of oogone. (G)
Exogenous oospore. (After Sparrow: Ann. Botany, 47(187) :517-542.)

antherid is usually 4-7, but in M. insignis Thaxter there may be up to
24-32. They are posteriorly uniflagellate and swim to the oogone or creep
along its surface in a jerky manner as the flagellum waves. The oogones
become uninucleate according to Laibach (1927) before the basal septum
is formed. They are pyriform or ovoid and when ready for fertilization
become open at or near the apex, sometimes at the top of a short broad
neck. The protoplasmic contents round up into a more or less spherical
egg near the base of the oogone or close to its opening. Upon entry of the
sperm into the egg the flagellum of the former disappears and the fertil-
ized egg remains in the oogone and forms a thick wall (endogenous
oospore) or creeps out through the oogonial opening and encysts exter-
nally (exogenous oospore), in some cases then falling ofT. Rarely both
modes of oospore formation may occur in the same species, e.g., M.
macrandra. The two sex nuclei do not unite in the oospore until shortly
before its germination by a slender germ tube. (Figs. 27, 28.)

Monohlepharella was distinguished as a separate genus by Sparrow

PHYCOMTCETEAE

(1940) with one species M. taylori Sparrow, but since that date a few
other species have been described. It is distinguished from Monohlepharis
by the behavior of the exogenous oospore which swims free by means of
the persistent fiagellum of the fertilizing sperm, becoming encysted after

Fig. 29. Monoblepharidales, Family Monoble-
pharidaceae. Monoblepharella mexicana Shanor. (A)
Hypha with zoosporangia. (B) Hypha with oogones
and antherids. (C) Oogone just after entry of sperm
cell. (D) Zygote swimming away by means of the
flagelkmi of the sperm cell. (E) Encysted zygote.
(Courtesy, Shanor: Mycologia, 34(3) -.241-247.)

having progressed some distance. Occasionally in M. taylori the oogone
may contain up to four eggs, though one is the more usual number. In
M. elongata Springer, in about half of the oogones more than one egg is
formed, sometimes as many as eight. (Fig. 29.)

Looking back from Monohlepharis we can see a fairly unbroken series

KEY TO THE FAMILIES AND GENERA OF ORDER BLASTOCLADIALES 91

from the eucarpic, monocentric, inoperculate Rhizidiaceae, through
Blastocladiella, Allomyces, and Monoblepharella to Monohlepharis, with
the uninucleate vegetative body in the more primitive forms becoming
transformed directly into a zoosporangium or gametangium, as in
Rhizophydium, while in Blastocladiella the vegetative structure is ovoid or
elongated with many nuclei, bearing the reproductive organs terminally.
Here the motile gametes are isogamous. In Allomyces the plant body is
more hypha-like and branched, and the reproductive organs are numer-
ous. In section Euallomyces the motile gametes are heterogamous. In
Monoblepharella the female gametes are one to eight in the oogone and
are motile, not by their own flagella, but by means of the flagellum of
the sperm. In Monoblepharis the egg is single in the oogone and is not
at all motile except in the species with exogenous oospores, where it
creeps out of the oogone after fertilization but has no power of further
locomotion.

Key to the Families and Genera of Order Blastocladiales

Plant body at first tubular, coenocytic, then forming alternately zoosporangia
and narrow isthmuses. Rhizoids arising at isthmuses or at all points. Resting
spores formed sometimes in place of zoosporangia.

Family Catenariaceae

Only genus. ^ Catenaria

Plant body coenocytic, branching and anastomosing, apparently without a cell
wall in its vegetative stage, multinucleate. Tips of the branches enlarging
to form thick-walled resting spores with the outer wall marked punctately
and with longitudinal lines. These produce posteriorly uniflagellate zoospores
soon, or after a resting period in the dry state. Parasitic in the larvae of
mosquitoes. Family Coelomomycetaceae

Only genus. Coelomomyces

Plant body with basally attached tuft of branching rhizoids and a globular or
clavate external basal segment upon which directly, or upon branches of
which, arise the reproductive organs (zoosporangia, resting spores, game-
tangia) . Sometimes the distinction between the basal piece and the branches
arising from it is only slight. Family Blastocladiaceae

Plant body with a simple or lobed or branched basal portion, often with sec-
ondary axes. Zoosporangia with single apical exit papilla. Alternation of
sporophytic and gametophytic generations unknown. Sexual reproduction
mostly unknown. Blastocladia

Plant body a more or less spherical structure which is directly transformed into
a reproductive organ, or more or less clavate, bearing at its top a single
reproductive organ. Discharge papilla one or several. Alternation of gen-
erations demonstrated in some species. Blastocladiella

Plant body consisting of a cylindrical basal segment giving rise — usually
dichotomously — to cylindrical, often constricted, branches with pseudo-
septa at the constrictions. Usually more than one discharge papilla. Sexual
reproduction by anisoplanogametes or isoplanogametes. Alternation of
sporophytic and gametophytic generations known in some species.

Allomyces

92 PHYCOMYCETEAE

Key to the Family and Genera of Order Monoblepharidales

Single family. Family Monoblepharidaceae

Branching mycelium constricted into cylindrical or rounded coenocytic seg-
ments, terminal ones becoming zoosporangia. Sexual reproduction not
certainly demonstrated. Gonapodya

Mycelium often not extensively branched, not constricted. Zoosporangia ter-
minal. Antherids and oogones terminal or subterminal. Egg cells not
flagellate.
Egg cells not motile by means of the flagellum of the sperm.

Monoblepharis
Egg cells motile by means of the flagellum of the fertilizing sperm.

Monoblepharella

Literature Cited

Bessey, Ernst A.: Isoplanogametes in Blastocladia, Mycologia, 3(3) :308-309.

1939.
Blackwell, Elizabeth: A life cycle of Blastocladia Pringsheimii Reinsch,

Brit. Mycol. Soc. Trans., 24(l):68-86. Figs. 1-9. 1940.
CoRNU, Maxime: Note sur deux genres nouveaux de la famille des Saprolegniees,

Bull. soc. botan. France, 18:58-59. 1871.
CoTNER, Frank B.: The development of the zoospores in the Oomycetes at

optimum temperatures and the cytology of their active stages. Am. J.

Botany, 17(6) :51 1-546. Ph. 30-32. Fig. 1. 1930a.
: Cytological study of the zoospores of Blastocladia, Botan. Gaz., 89(3):

295-309. Figs. 1-10. 1930b.
Couch, John N.: Observations on the genus Catenaria, Mycologia, 37(2) :163-193.

Figs. 1-78. 1945a.
: Revision of the genus Coelomomyces, parasitic in insect larvae, /.

Elisha Mitchell Sci. Soc, 61(1-2) :124-136. Pis. 1-2. 1945b.
, and H. R. Dodge: Further observations on Coelomomyces, parasitic on

mosquito larvae, ibid., 63(l):69-79. Pis. 15-20. 1947.
-, AND Alma J. Whiffen: Observations on the genus Blastocladiella, Am.

J. Botany, 29(7) :582-591. Figs. 1-66. 1942.
Emerson, Ralph: A new life cycle involving cyst-formation in AUomyces,

Mycologia, 30(2) :120-132. Fi>s. 1-11. 1938.
: An experimental study of the life cycles and taxonomy of AUomyces,

Lloydia, 4(2) :77-_144. Figs. 1-16. 1941.
Harder, Richard: tjber das Vorkommen vom Chitin und Zellulose und seine

Bedeutung flir die phylogenetische und systematische Beurteilung der Pilze,

Nach. Ges. Wiss. Gottingen. Math, physik. Klasse, Fachgruppe VI, N.F.,

3(l):l-7. 1937.
, und Georg Sorgel: Uber einen neuen planoisogamen Phycomyceten

mit Generationswechsel und seine phylogenetische Bedeutung, ibid., 3(5):

119-127. Figs. 1-4. 1938.
Hatch, Winslow R.: Sexualitv in AUomyces arbuscuhi Butler, /. Elisha Mitchell

Sci. Soc, 49(1):163-170. PI. 12. 1933.
: Gametogenesis in AUomyces, Ann. Botany, 49(196) :623-650. Figs. 1-33.

1935.
: Conjugation and zygote germination in AUomyces arbuscula, ibid., N.S.,

2(7):583-G14. P/s. lS-22. Figs. 1-13. 1938.

LITERATURE CITED 93

Keilin, D. : On a new type of fungus: Coelomomyces stegomyiae n.g., n.sp., para-
sitic in the body cavity of the larva of Stegomyia Scutellaria Walker (Diptera,
Nematocera, Culicidae), Parasitology, 13(3) :225-23i. Figs. 1-7. 1921.

Kniep, Hans: AUomyces javanicus n. sp. ein anisogamer Phycomycet niit Plano-
gameten, Ber. deut. hotan. Ges., 47:199-212. Figs. 1-7. 1929.

: tJber den Generationswechsel von AUomyces, Z. Botan., 22(9):433-441.

Figs. 1-2. 1930.

Lagerheim, G.: Mykologische Studien: II. Untersuchungen liber die Mono-
blepharideen, Kgl. Svenska Vetenskapsakad. Handl., 25, Afd. 3, No. 8: 1-42.
Pis. 1-2. 1900.

Laibach, Friedrich: Zytologische Untersuchungen iiber die Monoblepharideen,
Jahrb. wiss. Botan., 66: 596-630. Pis. 12-13. Figs. 1-12. 1927.

McCranie, James: Sexuality in AUomyces cystogenus, Mycologia, 34(2) :209-213.
Fig. 1. 1942.

VON MiNDEN, M.: Chytridiineae, Ancylistineae, Monoblepharidineae, Sapro-
legniineae, in Kryptogamenflora der Mark Brandenburg, vol. 5, pt. 3, pp.
353-496. Leipzig, Gebriider Borntraeger, 1911.

Nabel, Kurt: Uber die Membran niederer Pilze, besonders von Rhizidiomyces
bivellatus nov. spez., Arch. Mikrobiol., 10:515-541. Figs. 1-7. 1939.

Sparrow Jr., Frederick K. : Phy corny cetes recovered from soil samples col-
lected by W. R. Taylor on the Allan Hancock 1939 expedition, Allan Hancock
Pacific Expeditions, 3(6):101-112. Pis. 16-17. 1940.

: The Monoblepharidales, Ann. Botany, 47(187) :517-542. PL 20. Figs.

1-2. 1933.

Aquatic Phy comy cetes, exclusive of the Saprolegniaceae and Pythium,

xix + 785 pp. QMfigs. Ann Arbor, Univ. Michigan Press, 1943.
Thaxter, Roland: New or peculiar aquatic fungi: 1. Monoblepharis, Botan.

Gaz., 20 :433-440.P/. 29. 1895a; 2. Gonapodya Fischer and Myrioblepharis nov.

gen., ibid., 20:477-485. PL 31. 1895b; 3. Blastocladia, ibid., 21:4:5-52. PL 3.

1896.
von Wettstein, Fritz: Das Vorkommen von Chitin und seine Verwertung als

systematisch-phylogenetisches Merkmal im Pflanzenreich, Sitz. ber. Akad.

Wiss. Wien, Math, naturw. Klasse, AbL I, 130(1) :3-20. 1921.
Wolf, Fred T. : A contribution to the life history and geographic distribution of

the genus AUomyces, Mycologia, 33(2):158-173. Figs. 1-2. 1941.

PHYCOMYCETEAE: LAGENIDIALES
AND SAPROLEGNIALES

IN THE two foregoing chapters have been discussed organisms, chiefly
aquatic or soil inhabiting, with posteriorly uniflagellate swarm cells
and whose cell walls are deficient in cellulose or do not show the cellulose
reaction without preliminary treatment to remove some masking sub-
stances. Also organisms with a single anterior flagellum of the tinsel type
were discussed. In this and the next following chapter are treated forms,
many of them aquatic or soil inhabiting, in which the swarm cells are
anteriorly or laterally biflagellate and in which the cellulose reaction is
normally shown immediately upon application of chloriodide of zinc
without other preliminary treatment. The anterior flagellum is of the
tinsel type, the posterior one of the whiplash type. Here, too, a rather
well connected series can be followed from holocarpic, enddbiotic, mono-
centric forms up to fungi with extensive mycelium and complicated
modes of reproduction, both asexual and sexual. This series, of course,
may be read in the reverse direction, leading from the complex to simpli-
fied forms. The author does not follow Sparrow (1943) in considering that
the Plasmodiophoraceae are related to the other groups included in the
chapter since Plasmodiophora is apparently more closely related to the
Mycetozoa (see Chapter 2).

Order Lagenidiales. The Hmits of this order are far from definite and
it may be that, as in the first edition of this textbook, the genera and
famihes here included should be placed in the order Saprolegniales.
Sparrow recognized the close affinity of some of these to that order by
placing the Ectrogellaceae and Thraustochytriaceae there. At present the
author is not convinced that enough is known of the structures and fife
histories of all of these rather simple or reduced forms to warrant their
distribution in separate orders. It therefore seems preferable to use the
order Lagenidiales as a temporary catchall for a number of genera which
perhaps are not too closely related, but which have in common the
characters given below. The members of this order are (except Thrausto-

OEDEE LAGENIDIALES 95

chytriaceae) endobiptic and holocarpic in that they Hve within their host
cells, mostly as parasites, and lack rhizoids, the whole fungus body
becoming one or several reproductive units. Asexual reproduction is
accomphshed by the transformation of the plant body into a zoospo-
rangium which empties its anteriorly or laterally bifiagellate zoospores
into the water surrounding the host cell through an inoperculate exit
tube or tubes in the manner of Olpidium or Achlyogeton in the Chytridiales.
The zoospores may escape singly and swim away independently or may
encyst at the mouth of the exit tube and later escape and swim away.
In some cases the zoospores are not distinguishable in the sporangium
within the host cell but emerge as a mass of protoplasm to form an
external vesicle (as in Pythium) and there become organized and break
through the plasma membrane of the vesicle and swim away. In almost
all cases the zoospores are of the secondary type, i.e., more or less kidney-
shaped with one of the two lateral flagella directed anteriorly and the
other posteriorly.

Sexual reproduction where known is by the contact of whole plants
or of special segments and passage of the contents of one into the other,
which immediately becomes a zoosporangium or which may become a
thick- walled resting spore (oospore). In the latter case there is usually
no periplasm. The uniting organs (gametangia) may be approximately
equal and similar in appearance or the antherid may be much smaller
than the oogone and fertilization may be accomplished by a conjugation
tube. The thick-walled oospore has been observed in some cases to
germinate as a zoosporangium.

In the order as here defined it is exceedingly uncertain whether we
have a series progressing from a simple monocarpic fungus to a short
hypha of several reproductive units on the way toward the evolution of a
well-developed hypha such as we find in the Saprolegniales or Peronospo-
rales or, what may be equally likely, the fungi of this order represent
different degrees of reduction from various genera of those orders. In
conformity with the procedure in the Chytridiales-Monoblepharidales
series and in the Hyphochytriales the organisms in the series with
bifiagellate zoospores (the Biflagellatae of Sparrow) are arranged with
those of simple structure first.

Karhng (1942) recognizes five famihes in the group that he calls
"Simple Holocarpic Bifiagellate Phycomycetes." He does not wish to
convey the idea that these are necessarily a single phylogenetic series,
especially in view of the insufficient knowledge of the life histories of the
majority of the described species and of the uncertainty of their rela-
tionship, either as primitive or as reduced forms. If the latter, some of
them may represent reductions from Saprolegniaceae, Pythiaceae, or
Leptomitaceae that have a greater or less similarity because in reduction

96 PHYCOMYCETEAE

to simpler vegetative structure they perforce become very similar.
Sparrow (1943) includes these simple or simplified forms in five famihes
in two orders (not coinciding entirely with the famihes recognized by
Karling).

The author follows Sparrow in part in his distinction of three famihes
to which he adds doubtfuhy the Woroninaceae and Thraustochytriaceae.

Woroninaceae : fungus remaining naked and amoeboid for a considerable time,
forming a " Plasmodium "(?)■ Eventually separating into several segments
which produce cell walls and become zoosporangia or the whole "Plas-
modium" enlarging to form a single zoosporangium with its wall in close
contact with the host wall. Zoospores preformed in the zoosporangia. Clusters
of angular or single round resting spores may be formed. Parasitic in algae
and fungi.

Olpidiopsidaceae: one-celled, free in the host cell and early producing a cellu-
lose wall. Zoospores preformed in the zoosporangium. Resting spores formed
sexually or by parthenogenesis. Parasitic in algae or fungi.

Sirolpidiaceae: forming a linear series of zoosporangia, occasionally a single,
elongated zoosporangium. Zoospores preformed in the zoosporangium.
Sexual stage unknown. Parasites in marine algae.

Lagenidiaceae: one-celled or more often a short, constricted or unconstricted,
unbranched or branched row of cells, each of which becomes a zoosporangium
or gametangium. Zoospores completing their development in a vesicle at
the opening of the exit tube. Oospores formed within the female gametan-
gium. Parasitic in algae (mainly fresh-water forms) or microscopic animals
or roots of grasses.

Thraustochytriaceae: resembling Rhizophydium. Parasitic upon marine algae.
Zoosporangium epibiotic, obpyriform, attached to a branched endobiotic
rhizoidal system. Zoospores formed in the zoosporangium but not motile,
and set free by the dissolution of the apical portion of the wall, as angular
cells, from which after some time the biflagellate motile stage emerges.
Zoosporangia proliferating after the discharge of the zoospores. A family of
very doubtful relationship.

Family Woroninaceae. In this family of possibly three genera the
zoospores encyst on the external surface of the host and empty their
contents into the host cell through a slender tube, the empty spore walls
remaining attached for some time. Within the host cell the uninucleate
fungus remains naked and amoeboid and grows, accompanied by multi-
phcation of the nuclei, until it largely fills the cell. Eventually this
plasmodium-like structure separates into a group of several segments
around each of which a cellulose wall is produced. The zoospores produced
in these sporangia escape through exit tubes, those further inside the
sorus emptying through those nearer the surface. The zoospores are
biflagellate anteriorly. Instead of producing zoosporangia the naked
fungus mass may divide into very numerous small angular resting spores,
also with cellulose walls, which are clustered together in more or less
definite compact cystosori of a few to many spores. Each resting spore

ORDEB LAGENIDIALKS

upon germination gives rise to one zoospore or to several zoospores. The
foregoing description applies to Woronina, the type genus of the family,
which is parasitic in the hyphae of Saprolegniaceae and in Vaucheria.
Two other genera, Rozellopsis, parasitic in Pythiaceae, and Pyrrhosorus,
parasitic in Red Seaweeds, are tentatively also placed in this family but
differ in not forming clusters of zoosporangia and in their failure to pro-

FiG. 30. Lagenidiales, Family Woroninaceae. Woronina
polycystis Cornu. (A) "Plasmodium" nearly filling host
cell. (B) Early stage in the formation of zoosporangia.
(C) Empty zoosporangia. (D) Zoospore. (E) Infection
of host by zoospore. (F) Cystosori within host. (After
Cook and Nicholson: Ann. Botany, 47(188) :851-859.)

duce cystosori. The vegetative stage within the host is, as in Woronina,
more or less plasmodial in nature. (Fig. 30.)

Family Olpidiopsidaceae. With the removal of Woronina to a dis-
tinct family the remainder corresponds in the main to the Woroninaceae
of the previous edition. Vegetatively and in their asexual reproduction
the members of this family show great similarity to those of the Olpidiaceae
in the Chytridiales. They differ strongly, however, in producing zoospores
with two anteriorly or laterally attached flagella, one of the tinsel type,
the other of the whiplash type, and in possessing walls which give the

PHYCOMYCETEAE

Fig. 31. Lagenidiales, Family Olpidiopsidaceae. (A, B) Olpidiopsis vexans Barrett.
(A) Zoospores. (B) Zoosporangia of smooth and rough types and one mature oogone
with attached empty antherid. (C, D) Olpidiopsis varians Shanor. (C) Swollen tip of
hypha of Saprolegnia ferax (Gruith.) Thuret with three zoosporangia and one mature
oogone with attached, empty antherid. (D) Germination of oospore. (E) Olpidiopsis
luxurians Barrett, oogone and partially emptied antherid. (A and E, after Barrett:
Ann. Botany, 26(101) :209-238. B-D, after Shanor: /. Elisha Mitchell Sci. Soc,
55(1):167-195.)

cellulose reaction upon the application of chloriodide of zinc without any
preliminary treatment. It is not at all certain that all the genera assigned
to this family really belong here. The principal genus, of about 20 species,
is Olpidiopsis. The zoospores are anteriorly biflagellate or the flagella may
be somewhat laterally attached in a groove, one directed forward (tinsel
type) and one posteriorly (whiplash type), as demonstrated by Couch
(1941). The species of this genus are strictly parasitic in the hyphae of
Saprolegniaceae, Pythiaceae, and various algae. One species is reported
in Riccia and a doubtful species in an insect. The zoospore settles on the
outside of the host cell and encysts with a cellulose wall. It produces a
slender infection tube through which the uninucleate naked protoplasm
enters the host. It remains naked for a while but eventually forms a
cellulose wall and when fully grown becomes a spherical or ellipsoidal
zoosporangium. An exit tube pierces the host cell wall and the zoospores
escape through the softened inoperculate tip. Before the exit tube opens

ORDEK LAGENIDIALES 99

the zoospores become visible in the zoosporangium and after a few
moments the contents again appear homogeneous, the zoospores again
becoming visible and actively motile just before the tube opens. Depend-
ing upon the number of zoospores infecting the host cell there may be
from one to many sporangia formed. The host cell may not show much
enlargement or may be quite strongly hypertrophied. In the species para-
sitic in fungi the zoosporangium may be ornamented by few or many,
short or elongated, slender or stout spine-like processes which are pro-
duced by the host protoplasm, not by the parasite. Sexual reproduction
where known is by the union of two adjacent equal or unequal cells of
the parasite which at the point of contact produce a perforation of the
walls through which the multinucleate contents of one cell pass into the
other, also multinucleate, cell. Barrett (1912) reported the apparent union
of these numerous nuclei in pairs. The zygote cell enlarges somewhat and
then forms a thick, angular, smooth, or coarsely spiny or knobby, wall.
After a little while this resting spore becomes a zoosporangium and the
biflagellate zoospores emerge through an inoperculate exit tube. The
empty male cell, sometimes more than one, remains attached to the
oospore and is sometimes partly covered by the thickened wall of the
latter. Over 20 species have been described but careful culture work with
pure cultures is necessary before the validity of all these species can be
established. The genus was first set up by Cornu in 1872 for several
species occurring in the hyphae of Saprolegniaceae. He described the
zoospores as posteriorly uniflagellate but Fischer (1882) studying what he
believed to be the same species found that the zoospores were anteriorly
biflagellate. In 1884 the great German mycologist Wilhelm Zopf described
0. schenkiana in the filaments of Spirogyra, but described and figured the
zoospores as posteriorly uniflagellate. On this account von Minden (1915)
transferred the species to a new genus which he named Pseudolpidiopsis.
Scherffel and others studying what they believed to be the same species

» found that the zoospores were biflagellate. If the usually accurate ob-
: server Zopf was incorrect the genus Pseudolpidiopsis (tentatively included
in this book in the Olpidiaceae) must be rejected but if he was correct
the genus must be maintained. It must be noted that several cases are
known where apparently almost identical species have been found on
careful study to differ in their flagellation (e.g., Sphaerita dangeardii
Chat, and Brod., in the Olpidiaceae and Pseudo sphaerita euglenae
Dangeard, in the Olpidiopsidaceae, both parasitic in Euglena). (Fig. 36.)
In some species of Olpidiopsis the resting spores may arise partheno-
genetically under certain conditions and sexually under other conditions,
while in other species the sexual stage is unknown as yet. These were set
apart by Fischer (1892) as a separate genus Pseudolpidium. Probably this
does not deserve this generic distinction but Karling (1942) uses this as a

100 PHYCOMYCETEAE

temporary resting place for the species in which resting spores, either
sexually or parthogenetically produced, are unknown while Sparrow
(1943) uses this name for the forms with parthogenetically produced
resting spores. They occur in algae or fungi.

Peter senia was described by Sparrow (1934) for two species parasitic
in marine Florideae and one species in Saprolegnia, which differ from
Olpidiopsis in the production of lobed or tubular zoosporangia, mostly
with more than one discharge tube, and for which, so far, no resting
spores are known. In Pythiella (Couch, 1935) the primary zoospores are
without flagella and creep out through the mouth of the exit tube where
they encyst and after about an hour emerge as zoospores of the secondary
type, with the two flagella laterally attached. The female cell produces a
distinct egg with periplasm and the male cell fertilizes it through a short
conjugation tube. It is possible that this genus represents a very much
reduced form of Pyihium. The only known species is parasitic in the
hyphae of species of Pythium. Pseudosphaerita resembles Sphaerita of the
Olpidiaceae. It is imperfectly known and needs further study.

Family Sirolpidiaceae. In this as in the foregoing family the fungus
lies free within the host cell and produces a cellulose wall. Usually, how-
ever, this cell elongates and becomes divided by cross walls into several
zoosporangia which disarticulate, in Sirolpidium, or remain tubular,
sometimes shortly branched, with cross walls at maturity but not dis-
articulating, in Pontisma. Resting spores are not known in either genus.
Both are parasitic in marine algae, the former in Bryopsis and Cladophora,
the latter in the red alga Ceramium. In both genera the laterally biflagel-
late zoospores are produced in the zoosporangia and escape fully formed
through an inoperculate exit tube. Karling (1942) places Petersenia in
this family because of its elongated, tubular, or lobed cell body. Its type
species was originally described as a species of Pleotrachelus, of the
Olpidiaceae, but differs from that in the biflagellate zoospores.

Family Lagenidiaceae. This family was originally named Ancylisti-
daceae, but with the discovery by Miss Berdan (1938) that the genus
Ancylistes really belongs in the Entomophthorales, the present designa-
tion was given by Karling (1939). In this family the zoospores are mostly
of the secondary type, the primary type being rare. The contents of the
zoosporangium are set free through an exit tube into a vesicle in which
the zoospores attain their final form and then escape by bursting the
vesicle membrane. They encyst and germinate on the outside of the host
cell and form a slender infection tube at whose apex a short or long, simple
or branched, coenocytic plant body is formed. This may remain non-
septate in Lagcna or septate with cylindrical segments in Lagenidium or
constricted at the septa into bead-like segments as in Myzocytium. Each
segment may become a zoosporangium or a gametangium. Adjacent seg-

ORDER LAGENIDIALES

101

Fig. 32. Lagenidiales, Family Lagenidiaceae. (A-D) Lagenidium rabenhorstii Zopf.
(A) The coenocyte has divided into several zoosporangia. (B) Forty-five minutes
later, the cytoplasm is escaping from one of these zoosporangia into a vesicle. (C) Four
successive stages in the development of the same vesicle. (D) Plant with four empty
zoosporangia and an empty antherid and an oogone containing a mature oospore.
(E, F) Myzocytium proUferum Schenk. (E) Plant with empty zoosporangia, one with
a vesicle containing zoospores. (F) Plant with three antherids and three oogones con-
taining mature oospores. (After Zopf: Nova Acta Leopoldina, 47(4):141-236.)

ments in the same hypha or in separate hyphae that are in contact may
function as antherid and oogone, respectively. They may be ahke in size
and other appearances or shghtly different. By means of a conjugation
tube the contents of the antherid pass into the oogone and then a thick-
walled, rounded oospore is formed. There is no distinct periplasm.
Germination of the oospore was described by Dangeard (1903) in Myzo-
cytium vermicola (Zopf) Fischer, which is parasitic in a free-living nema-

102 PHYCOMYCETEAE

tode. The nucleus divides into several nuclei and then an exit tube is
formed, but the actual production of the zoospores was not observed.

The members of this family are parasitic, rarely saprophytic, in cells
of algae (mostly fresh-water species), microscopic animals, or their eggs,
and in the case of Lagena, parasitic in the roots of grasses. Sometimes a
small individual is nonseptate and is scarcely to be distinguished from
one of the Olpidiopsidaceae while in Lagenidimn giganteum Couch the
fungus body is elongated, multiseptate, and quite mycelium-like and up
to 40 fj. thick in extreme cases. It is in cases like this that the supposition
is strengthened that this family represents a reduction in size and com-
plexity from higher, better developed fungi, perhaps close to Pythium.
The three genera recognized by Karling and by Sparrow are Lagenidium,
with 15-20 species, Myzocytium, with 4 or 5 species, and Lagena with one
species.

In the genus Lagenidium the coenocytic segments may be few in
number and all included within one cell of the algal host or they may
pierce the cell walls of the latter to continue through several cells. In
L. giganteum the hosts are small aquatic Crustacea and larvae of mos-
quitoes. Not only does the extensive coarse mycelium fill the host but it
extends out into the surrounding water up to a distance of 0.1 mm. This
species has been brought into pure culture by Couch. Its sexual stage is
unknown. In those species where sexual reproduction has been observed
the female gametangium usually enlarges considerably as the oospore
develops. The antherid is an unmodified or only slightly modified vegeta-
tive cell, either adjacent to the oogone in the same filament or in an
adjacent filament. In the latter case this may indicate heterothallism.
Lagena (Vanterpool and Ledingham, 1930) is a parasite in the roots of
wheat {Triticum aestivum L.) and other grasses, which are weakened by
the presence of this fungus. The organism may be much elongated or
even more or less coiled in the host cell but not septate. It produces a
single exit tube through which the contents of the zoosporangium emerge
into a vesicle in which they undergo their final transformation and swim
away after rupturing its membrane. Ne,ar-by thalli may unite by a
conjugation tube through which the contents of one cell pass into the
other, rounding up there to form a thick-walled oospore. The mode of
germination of the latter is unknown. (Figs. 32, 33.)

Possibly belonging to this family is Aphanomycopsis (Scherffel, 1925)
forming a more or less elongated and coiled, often somewhat branched,
coenocytic thallus in the cell of a Diatom. This thallus is transformed into
a zoosporangium with one or more exit tubes at whose mouths the primary
(nonflagellatc) zoospores encyst, later emerging and swimming away as
zoospores of the secondary type. Sexual reproduction has not been
demonstrated although thick-walled resting spores are formed in enlarged

OKDER LAGENIDIALES

103

Fig. 33. Lagenidiales, Family Lagenidiaceae. Lagena radicicola Vanterpool &
Ledingham. (A) Zoospore and semidiagrammatic representation of infection of root
hair of wheat. (B) Zoosporangia in top and side view and an empty zoosporangium.
(C) Formation of vesicle at apex of discharge tube of zoosporangium. (D) Semidia-
grammatic representation of the conjugation of two thalli. (Courtesy, Vanterpool
and Ledingham: Can. J. Research, 2(3):171-194.)

104 PHYCOMYCETEAE

portions of the tubular thallus. Because of its diplanetism Sparrow (1943)
places this genus in the family Ectrogellaceae in the Saprolegniales.
Ectrogella, also parasitic in Diatoms, consists of short or somewhat
elongated, unbranched coenocytic tubes which become zoosporangia
whose zoospores are diplanetic, usually with the primary zoospores
biflagellate and encysting at the mouth of the exit tube or near by.
Sexual reproduction by the union of two adjacent thalli in the same host
cell has been reported. This results in the formation of a rounded, thick-
walled oospore lying more or less loosely in the oogone to which the
empty male cell remains attached by the persistent conjugation tube.
Eurychasma and Eurychasmidium, occurring in marine algae, have been
assigned by Sparrow to the same family.

Family Thraustochytriaceae. Sparrow (1943) places the genus
Thraustochytrium in a family of its own. This fungus is Rhizophydium-\\ke,
growing saprophytically upon marine algae. There is a system of branched
rhizoids within the host cell and an external obpyriform zoosporangium
which renews itself by proliferation upon the discharge of the zoospores.
The latter are angular and not motile at first and later become trans-
formed into pyriform, biflagellate zoospores. Because of this similarity to
diplanetism Sparrow places this family in the Saprolegniales.

Order Saprolegniales. This order consists of fungi with well-marked
hyphal development. In fact the largest fungus hyphae known are to be
found here. Thus Monsma (1937) found, growing on hemp seed in water,
hyphae of Achlya oblongata de Bary var. globosa Humphrey that attained
a diameter of 270 m near the base and were so stiff that, on removing the
seed from the water, the hyphae stood out straight to a length of 15 mm.
The fungus body in this order consists mostly of a well-developed system
of branching, usually tapering, filaments within the substratum and an
external portion mostly of coarse or slender, unbranched or less strongly
branched hyphae which bear the reproductive organs. These hyphae may
be nearly uniform in diameter or tapering gradually from base toward
the apex or they may be more or less regularly constricted here and there,
sometimes the orifices being closed by plugs of cellulin. In one family the
basal portion of the external mycelium consists of more or less rounded
or clavate bodies from whose upper portion arise slender, usually con-
stricted, hyphae.

The mycelium of this order is coenocytic with a multinucleate layer
of cytoplasm surrounding a large central vacuole in the larger hyphae.
The cell walls show the cellulose reaction immediately upon treatment
with chloriodide of zinc. Usually septa are lacking in the mycelium
except: (1) to delimit zoosporangia, (2) to set apart the gametangia from
the remainder of the hyphae, and (3) to delimit injured portions. In
Achlya polyandra de Bary, Horn (1904) observed and showed to the

ORDER SAPROLEGNIALES 105

author the extensive and rapid formation of transverse and obHque walls
until the hypha was- cut up into innumerable angular segments, under
the influence of exceedingly minute amounts of copper in the water.
Growth of the external hyphae is terminal and by the limited formation
of branches. Unlike the Lagenidiales the mycelium does not become all
converted into reproductive organs but these are mostly terminal or
subterminal on the external hyphae and their branches.

The Saprolegniales are saprophytic on dead plant or animal matter
in the soil or in water (fresh water, rarely brackish) or parasitic in algae
or small animals or even fish, and in some cases in the roots of plants in
the soil. Although spoken of usually as water molds perhaps the majority
are inhabitants of moist soil.

The zoosporangia are of the same diameter as the hyphae or somewhat
enlarged, cylindrical or ovoid. At their inception the protoplasm from
the lower portions of the hyphae crowds into the terminal portions
destined to become the zoosporangia until these are filled with dense,
multinucleate protoplasm with a much reduced central vacuole. At the
base of each a septum is formed setting it apart from the supporting
hypha. Within this zoosporangium cleavage of the protoplasm begins
next to the wall and progresses inwardly to the central vacuole, cutting
out uninucleate naked portions of protoplasm which round up somewhat
and eventually become the zoospores. These are usually pyriform, with
two anterior flagella, one of the tinsel type and the other of the whiplash
type. Such zoospores are called primary zoospores. Escape of the zoospores
occurs through the softened tip and sometimes through a lateral papilla.
In all cases the opening is inoperculate. After the zoosporangium is
emptied the basal septum may arch up into the empty space, filling it
and becoming a second zoosporangium. This formation of zoosporangia
by proliferation may occur repeatedly. Instead of proliferating, the hypha
just below the emptied zoosporangium may branch out laterally, the
branch quickly turning upward to form a new zoosporangium parallel to
the empty one. This sympodial formation of zoosporangia is characteristic
of some genera.

The primary zoospores may escape and swim away some distance
before encysting. In some species of Pythiopsis this encysted primary
zoospore eventually germinates by a slender tube and forms a new plant,
or the encysted zoospore may escape from its cell wall as a primary
zoospore again. More often in other genera the encysted primary zoospore
escapes from its cyst after a few minutes or hours in a different form, the
secondary zoospore. It is kidney-shaped or resembles a grape seed with
the two flagella arising in the groove, usually nearer the more pointed
end than the base. One flagellum, of the tinsel type, is directed forward
while the whiplash type flagellum is directed posteriorly. This zoospore

106 PHYCOMYCETEAE

encysts after a while and may germinate by a slender hypha or re-escape
and re-encyst several times. In some cases the primary zoospore appar-
ently produces no flagella but creeps to the opening of the zoosporangium
and there encysts, escaping as a secondary zoospore some time later. In
still other genera the primary, nonflagellate, zoospores encyst within the
zoosporangium as separate round cells or become compacted into poly-
hedral cells by mutual pressure. In the latter case they form short exit
tubes that pierce the zoosporangial wall so that their contents escape
as secondary zoospores. In a few species the zoospores that encyst within
the zoosporangium do not escape at all as swimming cells but germinate
directly by germ tubes.

It is customary to refer to the characteristic of forming primary
zoospores only, as monoplanetic and of the formation of two successive
types of zoospores as diplanetic. More correctly these two terms should
be monomorphic and dimorphic. Properly speaking monoplanetic means
wandering once, or with only one swimming stage, while diplanetic means
with two swimming stages. Since in Pythiopsis the primary type of
zoospore may swim and encyst several times and in Achlya, Dictyuchus,
and other genera, the secondary type of zoospore may also do the same
thing the customary terms are not used in their correct etymological
sense.

Sexual reproduction occurs by the formation of male and female
gametangia, antherids and oogones, respectively. They may be terminal
on the main hyphae or on short lateral branches, rarely intercalary. The
oogones round up and become filled with multinuclear protoplasm and
then a cross wall is formed. Most of the nuclei degenerate leaving a much
reduced number. A single egg or several eggs may be formed, containing
a few nuclei each, of which only one remains sexually functional, the
others moving toward the outer wall and usually disintegrating. The
whole of the original protoplasm may be used up in the formation of
the egg or eggs or a portion of it may be left surrounding a single egg.
This is called the periplasm and may contain numerous nuclei which
eventually disintegrate. In the genus Araiospora the periplasm becomes
divided by radial sheets of protoplasm in which these nuclei lie and then
radial cell walls are formed so that the egg is surrounded by a layer of
cells. Probably in most cases the periplasm produces much of the exterior
ornamentation of the oospore wall.

The antherids, depending upon the species of the fungus, may be
formed on separate plants from those forming the oogone (heterothallic
species), or both kinds of sexual organs may be produced on the same
mycelium (homothullic species). This distinction is not in all cases sharp
as there may be varying degrees of maleness or femaleness, as shown by
Bishop (1940). The antherids are terminal on long slender branches or on

OEDEE SAPEOLEGNIALES 107

short branches or may arise as enlargements of the oogonial stalk, just
below the oogone. When arising at a distance these antheridial hyphae
are attracted to the oogones apparently by a secretion from the latter.
J. R. Raper (1939, 1940) shows evidence that these are in the nature of
hormones. In heterothallic species of Achlya he demonstrated that a
secretion from the male plant stimulates the formation of oogones on the
female plant and that in turn the substances given off by the developing
oogones lead to the production of antheridial branches in the male plants
and, probably, guide chemotropically the direction of their growth.

Upon reaching the oogone the usually somewhat enlarged tip of the
antheridial branch flattens against it and, if it has not already occurred,
a septum is produced, separating the antherid from the supporting
hypha. The antherid is usually plurinucleate and the nuclei may divide
again. Eventually most of them degenerate. At or near the center of the
contact surface a tube grows from the antherid through the oogone wall
to the egg, or if there are several eggs this conjugation tube may become
branched so that one antherid may fertilize several of them. Couch (1924)
showed that in Leptolegnia caudata de Bary an opening is dissolved
between the antherid and oogone permitting fertilization without the
formation of a conjugation tube. According to Kevorkian (1925) this is
true also of Apodachlya hrachynema (Hild.) Pringsh. while Cooper (1929)
also demonstrated this for Brevilegnia diclina Harvey. After the entry
of a single sperm nucleus into the egg the latter secretes a definite wall
which may become thick, with a smooth or rough exterior. The union of
nuclei does not occur until much time has elapsed. Although it has not
been demonstrated it is assumed that meiosis occurs at the germination
of the oospore. This usually occurs by the formation of a hypha which
may develop to form a new plant or which may produce a zoosporangium.
In many species of this order the oospores develop parthenogenetically.
The formation or nonformation of antherids depends upon the conditions
of nutrition, temperature, etc. It has been asserted that in some cases
antherids, though present, may not function.

It is difficult to make a decision as to whether the pluriovulate con-
dition should be considered the more primitive or a derived condition
in this order. If the ancestral forms were fungi whose female gametangia
contained several large motile eggs (as in Allomyces), we would expect
the more primitive Saprolegniales to be pluriovulate, but if we look to
the Lagenidiales for the stock whose evolution led to the Saprolegniales,
we find that they have but a single egg. If we look to the Siphonales, we
find that in these green algae in the genus Vaucheria the oogone contains
but a single egg while there are other members of that group in which the
oogone contains several eggs. The author is inclined to favor the hypothe-
sis that the pluriovulate condition is derived from the uniovulate con-

108 PHYCOMYCETEAE

dition. Of the more than 20 genera included in this order the egg is single
in all but 6 or 8 genera and also in some of the species of each of these
genera that are normally pluriovulate. The species of this order have
been monographed by Coker (1923) and the North American species by
Coker and Matthews (1937).

The author recognizes three families in this order: Saprolegniaceae,
Leptomitaceae and Rhipidiaceae. The two latter are considered by Miss
Kanouse (1927) and by Sparrow (1943) to deserve segregation into a
separate order, the Leptomitales, with which the author does not agree.
Until the various forms included in this book in the Lagenidiales are
better known, it is uncertain whether some of them should be more
closely associated with the Saprolegniales. There is undeniably a close
relationship between these two orders and also with the next order, the
Peronosporales.

The three families of Saprolegniales may be distinguished as follows:

Saprolegniaceae: mycelium not definitely constricted at intervals nor with
cellulin plugs, mostly cylindrical or gradually narrowing toward the extrem-
ities but not consisting of an enlarged basal portion and slender branches.
Zoospores mostly dimorphic or with modifications of dimorphism, only pri-
mary zoospores produced in one genus. Oogone with one or more eggs and
lacking periplasm.

Leptomitaceae: mycelium definitely constricted at more or less regular inter-
vals, frequently with cellulin plugs. Zoospores dimorphic. Oogones usually
with one egg, in one species with more, with no periplasm.

Rhipidiaceae: mycelium more or less well differentiated into an enlarged basal
portion with slender, mostly constricted, branches which bear the zoospo-
rangia and sexual organs. (In Mindeniella these arise directly on short stalks
from the basal segment.) Oogones with single eggs and with periplasm.

Family Saprolegniaceae. The members of this family are in some
cases strictly aquatic, but a considerable number of species are soil in-
habitants. Contrary to the belief prevailing earlier they are mostly
saprophytic on vegetable matter, less often on animal matter. Only a few
species of Achlya and Saprolegnia are sometimes destructive to young fish
and fish eggs in fish hatcheries. Several species of Aphanomyces and one or
two other genera are parasitic in algae, in the roots of higher plants, or in
aquatic animals.

In about half of the genera the oogone contains but one egg, but the
pluriovulate species are far in the majority. The number of eggs per
oogone in these may vary from 2 or 3 up to 50. Fertilization of the eggs is
accomplished by the passage of sperm nuclei, one to each egg, usually
through conjugation tubes that penetrate the oogone wall from the
adhering antherids. In some cases although antherids are present there
seems to be no opening for the passage of the sperm nucleus so that the
egg develops parthenogenetically. The oospore may germinate by a germ

ORDER SAPROLEGNIALES

109

Fig. 34. Saprolegniales, Family Saprolegniaceae. Thraustotheca prirnoachlya Coker
& Couch. (A) r/ira«sto</iemtypeof zoosporangium. (B) ^c/i/ya type of zoosporangium.
(C) Oogone and antherid. (D) Oospore germinating and producing several small
zoosporangia. (Courtesy, Coker and Couch: J. Elisha Mitchell Sci. Soc, 40(3-4):
197-202.)

tube which produces a new mycelium. In Thraustotheca prirnoachlya
Coker & Couch the germinating oospore may divide into several internal
spores or these may be formed in short germ tubes growing out through
the pits of the oogone wall (Coker and Couch, 1924). Ziegler (1948)
studied the germination of 26 species of this family representing 6 genera
and found the following 4 types. (1) "A long or short germ tube is formed,
with an apical sporangium"; (2) "the germ tube produces a sparsely
branched mycelium with a sporangium at the apex of the main hypha or
a branch"; (3) "the primary germ tube forms a branched mycelium";
(4) "the primary germ tube forms a long unbranched hypha." (Fig. 34 D.)

Apparently the primitive form of zoospore in this family is pear-
shaped with two equal anterior flagella. Only such primary zoospores are
formed in the two known species of Pythiopsis. Far more often the species
of this order are dimorphic. Other forms show various modifications of
the dimorphic plan.

The zoosporangia are typically terminal segments of hyphae, but
sometimes several are formed one behind the other. When the zoospores
are discharged a new zoosporangium may arise by proliferation, some-
times five or six times. In other cases the new zoosporangia are formed
on short or long sympodially produced branches. Usually they are slender,
like the supporting hypha, or clavate or ovoid. Under certain cultural
conditions the hypha may form ovoid or clavate zoosporangia in chains,
each opening by an exit pore near its apical end. Under some conditions

110 PHYCOMTCETEAE

such potential zoosporangia may round up into thick-walled resting
spores or chlamydospores.

Emergence of the zoospores is usually through the softened apex of
the zoosporangium. In Saprolegnia, Leptolegnia, and Isoachlya the primary
zoospores swim away as soon as released, encysting separately at some
distance from the zoosporangium. In Achlya, Aphanomyces, and several
other genera the escaping primary zoospores encyst immediately on
emerging and form a ball of cells which release the secondary zoospores.
In Thraustotheca and other genera the primary zoospores encyst within
the zoosporangium and upon rupture of the latter the encysted spores
are set free and give rise to the secondary zoospores. In Dictyuchus the
encysted spores are polyhedral by mutual pressure and germinate within
the zoosporangium by short exit tubes which pierce the zoosporangium
walls, thus setting free the secondary zoospores individually. In Aplanes
and Geolegnia and some other genera the encysted primary spores germi-
nate by germ tubes within the zoosporangium or after the latter has
disintegrated. Under varying conditions of culture the same species of
Saprolegnia or Achlya may be induced to produce its zoospores in the
manner typical of Saprolegnia, Achlya, Thraustotheca, or Aplanes, showing
that these modifications are not of very deep fundamental importance.
This is corroborated by the fact that Salvin (1942) was able to succeed
in attempts at mating Thraustotheca clavata (de Bary) Humphrey with
Achlya flagellata Coker, the former producing the antherids and the latter
the oogones. The oospores so produced could not be brought to germina-
tion by the methods attempted. (Figs. 34-38.)

Pythiopsis. In this rarely studied genus the sympodially produced
sporangia may be ovoid (P. cymosa de Bary) or slender. The mycelium
is rather stout, as is typical for the family. The zoospores which escape
are of the primary type. After encysting they may germinate by a germ
tube or may produce zoospores again, but these are still of the primary
type in the two species. The oogone usually has but one egg. The antherids
may be up to three in number and may arise from immediately below the
oogone. The species of this genus occur in soil or in fresh water. A third
species assigned to this genus by Harvey (1925) should be transferred,
according to Coker and Matthews (1937) to the genus Isoachlya.

Saprolegnia. This is the most commonly studied genus of the family.
It contains about 20 species, mostly saprophytic, rarely parasitic, on
animal or vegetable matter in water or soil. Dead insects in water or even
larger animals become surrounded by a fringe of the long external hyphae,
the much-branched trophic hyphae being within the animal tissues. These
external hyphae are fairly stout (up to 50 to 100 microns in diameter in
extreme cases) and more or less straight and but little branched. They
terminate in club-shaped zoosporangia within which numerous zoospores

ORDER SAPROLEGNIALES

111

Fig. 35. Saprolegniales. (A-E) Family Saprolegniaceae. (A, B) Saprolegnia monoica
Pringsh. var. glomerata Tiesenh. (A) Proliferated zoosporangia. (B) Oogone and
antherid. (C-E) Achlya racemosa F . Hildeb. (C) Cluster of zoosporangia. (D) Gemmae.
(E) Oogone and antherids. (F) Family Leptomitaceae, Leptomitus ladeus (Roth)
Agardh. Portion of filament showing one empty sporangium, one containing zoo-
spores, and the other not yet mature. (Courtesy, Coker: The Saprolegniaceae with
Notes on Other Water Molds, Chapel Hill, Univ. North Carolina Press.)

112 PHYCOMYCETEAE

are present in no definite arrangement. The primary zoospores squeeze
out one by one from the terminal opening and swim away, encysting at a
distance. The new zoosporangia are formed by proHferation within the
empty ones. Usually a little later, under conditions which can often be
controlled in culture, short or long lateral branches arise which swell at
the apex into a globular oogone separated by a septum from the main
hypha. The oogone may be single or there may be a chain of several
oogones. Within the multinucleate oogone the protoplasm cleaves into
several or many masses which round up to form the naked eggs
(oospheres). These are at first multinucleate but soon all the nuclei but
one disintegrate. On longer, usually more slender, branches from the same
plant (often arising just below the oogone) or from a different plant the
antherids are produced. These are terminal, multinucleate structures,
somewhat larger in diameter than the supporting hyphae, from which
they are separated by septa. Upon reaching an oogone they become
flattened against the outer surface. From the center of the surface of
contact a papilla pushes into the oogone, forming the conjugation tube
which seeks out an egg or branches so as to reach several eggs. A single
sperm nucleus passes into each egg if fertilization actually takes place.
In many cases there is no fertilization and the egg becomes a thick-walled
oospore, parthenogenetically. After the disintegration of the oogone the
oospore may lie dormant in the mud for several months, eventually
germinating by a tube which may or may not be terminated by a zoospo-
rangium. Reduction division probably occurs as the oospore germinates.
(Fig. 35A, B.)

Achlya, with about 25 species, resembles Saprolegnia in habit and
manner of growth. The chief morphological difference is that the hyphae
bearing the zoosporangia grow sympodially so that there is no prolifera-
tion of zoosporangia. The behavior of the zoospores is also characteristic.
The primary zoospores encyst immediately as they emerge from the
mouth of the zoosporangium, forming a very typical cluster of encysted
spores. Only after 15-45 minutes or longer do these spores give rise to
secondary zoospores. The oogones are usually pluriovulate and in most
respects resemble those of Saprolegnia. In both Saprolegnia and Achlya
and in a few other genera the presence or absence of thin places or "pits"
in the oogone walls is of diagnostic importance. Many species of Achlya
are parthenogenetic. Both of these genera are usually exceedingly sensi-
tive to minute traces of copper salts as discovered by Horn (1904), j'-et
Gaumann (1919) found ;S'. monoica Pringsh. growing in the shaft of an
abandoned copper mine in Lapland where the dissolved salts gave the
water a green color. (Fig. 35C-E.)

Aphanomyces has slender zoosporangia with but a single row of
zoospores which behave on emerging as do those of Achlya. The oogone

ORDER SAPROLEGNIALES

113

Fig. 36. Saprolegniales, Family Saprolegniaceae. (A-C) Aphanomyces exoparasiti-
cus Coker & Couch. (A) Encysted zoospores at mouth of zoosporangium. (B) Young
oogone and antherid. (C) Mature oogone containing oospore. (D, E) Aphanomyces
phycophilus de Bary. (D) Discharged zoospores. (E) Oogone and antherid. (A-C, after
Couch: J. EHsha Mitchell Sci. Soc, 41(3-4) :213-227. D-E, after Sparrow: Mvcoloaia
22(3):118-121.)

has but a single egg. The species of this genus are parasitic in algae and
in the roots of higher plants, where they may cause root rots, as well as
upon some aquatic animals, mostly Crustaceans. .4. acinetophagus Bartsch
and Wolf (1938) has been described from a fresh-water protozoan.
Scarcely distinguishable from Aphanomyces is Hydatinophagus parasitic
upon Rotifers. Another closely related genus, also parasitic upon Rotifers,
is Sommersiorffia, with special spike-like branches which catch the host

114

PHYCOMYCETEAE

Fig. 37. Saprolegniales, Family Saprolegniaceae. Leptolegnia caudata de Bary. (A)
Young oogone and two antherids. (B) Same structures five hours later, one antherid
discharging contents into oogone. (C) Same structure two and a half days later,
oospore wall considerably thickened. (D) Stained section showing entry of sperm
nucleus into the egg, with remains of peripheral supernumerary nuclei in both oogone
and antherid. (Courtesy, Couch: Am. J. Botany, 19(7) -.584-599.)

victims. Plectospira (Drechsler, 1927, 1929) is parasitic in plant roots and
resembles Aphanomyces except for the production of lobulate masses of
hyphae which apparently serve as auxiliary storage parts of the zoospo-
rangia, much as occurs in some species of the genus Pythium (see p. 127).
The oogones have a single egg but no periplasm and become surrounded
by many antherids, up to over 50, of which only a few reach full develop-
ment. Leptolegnia resembles Aphanomyces in its slender hyphae and its
slender zoosporangium with a single row of zoospores, as well as in the
production of only a single egg in the oogone. It differs in that the primary ,
zoospores swim away immediately and encyst at a distance as inj
Saprolegnia. (Figs. 36, 37.)

Dictyuchus resembles Achlya and like it produces clavate or cylindricj
zoosporangia which may be single or in chains or more or less sympodially
clustered. They may separate from the supporting hyphae and float

ORDKR SAPROLEGNIALES

115

v^-

around carrying within them the encysted primary zoospores. Normally
these zoospores encyst within the zoosporangium and by mutual pressure
become more or less polyhedral in shape, depending upon their number.
The zoosporangial wall may be persistent, in which case the secondary
type zoospores escape through short exit papillae which pierce the sur-
rounding wall, so that eventually a net-like structure remains. In other
species the zoosporangial wall is evanescent early so that the encysted
primary zoospores are rounded on
their outer face and somewhat flat-
tened where they are in contact.
They may become separated under
pressure. The emerging zoospores
in both types are of the secondary
type and may encyst and emerge
again several times before germi-
nating by germ tubes. In the species
with persistent zoosporangium walls
this is the only type of zoosporan-
gium known. In those with evanes-
cent walls under certain conditions
of culture medium and temperature
the first zoosporangia (according
to Couch, 1931) empty their zoo-
spores through an apical pore where
they encyst as in Achlya. The later
zoosporangia are as in Dictyuchus.
The oogone contains a single egg.
The one species with several eggs,
D. polysporus Linds., assigned here
has been determined by Apinis
(1930) to belong to the genus Pro-
toachlya. Most of the known species
have been isolated from the soil
but some occur also in fresh water. Both homothallic and heterothallic
species are known (Couch, 1926b; Coker and Braxton, 1926). In Thrausto-
theca a similar preliminary production of Achlya-like zoosporangia has
been reported (Coker and Couch, 1924). (Fig. 38.)

Miss Huneycutt (1948) has described a new genus, Aphanodictyon,
with the single species A. papillatum. This has the vegetative structure of
Aphanomyces, with very slender, branching mycelium. The zoosporangia
are globose or subglobose and the primary zoospores encyst within them
as in Dictyuchus. The secondary type zoospores escape through short
exit tubes. The oogones contain 1-8 eggs (mostly 3-6), which are fertilized

Fig. 38. Saprolegniales, Family Sap-
rolegniaceae. Dictyuchus missouriensis
Couch. (A) Early zoosporangia of Achlya
type. (B) More enlarged zoosporangium
of Dictyuchus type. (Courtesy, Couch: /.
Elisha Mitchell Sci. Soc, 46(2) -.225-230.)

116 PHTCOMYCETE AE

by antherids of androgynous or diclinous origin. The oogones have few to
many papillae about 2 n thick and of variable length up to 20 n. The
fungus occurs on keratinized material in the soil and may be cultured on
thin slivers of horse hoof.

Apinis (1935) described a genus which he named Archilegnia in which
fertilization is claimed to be by motile uniflagellate sperm cells. In other
respects such as the mode of asexual reproduction and the structure of
the pluriovulate oogones this fungus is like Saprolegnia. Not only do the
sperms arise in short antherids growing out at right angles from the main
hypha, but the author claims that encysted zoospores may give rise to
four sperms. He reports that the sperms enter the oogone through small
openings. It is likely that this genus was described from a Saprolegnia
parasitized by small Protozoa or Chytridiales. If the correctness of the
reports of Apinis can be confirmed it will throw great light upon the
possible ancestry of the Saprolegniaceae.

Family Leptomitaceae. In this family the mycelium is slender as in
the foregoing family, but is constricted at more or less regular intervals.
The constrictions are sometimes plugged by granules of carbohydrate
nature to which the name cellulin has been given. Similar granules may
be found scattered in the cytoplasm. The cell walls give the cellulose reac-
tion immediately when treated with chloriodide of zinc. The zoospores are
dimorphic. The oogones contain no periplasm. In Apodachlya there is
only one egg in the oogone but in Apodachlyella there are several. All the
known species of the family are saprophytic, chiefly on matter of vege-
table origin. Some species are aquatic, growing on sticks, fruits, etc.;
others grow unattached in water rich in organic matter; and some grow
in soil.

Leptomitus, of which L. lacteus (Roth) Agardh is perhaps the only
species, consists of branching, cylindrical hyphae rather uniform in size,
found in water containing large amounts of organic matter, such as
drainage water from sugar factories, near the mouths of sewers, etc. It is
easily distinguished by its constrictions which may remain open or become
plugged by granules of cellulin. The terminal segments first and then
successively the segments behind them become converted into zoospo-
rangia in which are produced numerous pear-shaped primary zoospores
all of which may escape successively through the terminal zoosporangium
or from separate openings from each zoosporangium. The zoospores
scatter as do those of Saprolegnia, eventually giving rise to zoospores of
the secondary type. Sexual reproduction is unknown. (Fig. 35 F.)

Apodachlya is also much branched and has no specialized holdfast
hyphae. The zoosporangia are distinct from the main hyphae, and are
mostly ovoid or pyriform, tapering below to a short pedicel. The oogone
contains a single egg without periplasm and may be terminal or lateral.

ORDER SAPROLEGNIALES 117

The antherid may subtend the oogone, directly, or arise on a short stalk
from immediately below it. No conjugation tube seems to be produced.
Germination of the oospore is by means of one or two germ tubes.

In ApodachlyeUa zoosporangia have not been observed. The spherical
or pyriform oogones contain 2 to 12, more often 4 to 7 eggs, which become
thick-walled oospores without surrounding periplasm. The slender con-
stricted antheridial branches, 2 or 3 in number, arise from the segment
below the oogone. A conjugation tube is produced by each functioning
antherid.

Family Rhipidiaceae. In this family the mycelium is provided with
well-developed holdfast hyphae bearing a more or less thickened, some-
times even spherical, basal segment from whose upper portion arise
slender, often constricted, hyphae terminated by zoosporangia or by the
sexual organs, or both may arise directly from the basal segment
(MindenieHa). Only zoospores of the secondary type are produced. The
oogone contains a single egg surrounded by periplasm which in one genus
{Araiospora) forms a cellular layer closely investing the oospore.

Sapromyces, with two or more species, grows on sticks, etc., in water,
being attached by its rather few rhizoids. The upright main axis is not
much thickened in comparison to the several slender, constricted branches
that arise at its apex. These bear at their apices one to several obovoid,
clavate, or cylindrical zoosporangia which give rise to numerous biflagel-
late, kidney-shaped zoospores which escape directly or into an evanescent
vesicle. Possibly due to sympodial growth some of the zoosporangia may
appear to be lateral. On the same branches with the zoosporangia or on
separate branches the oogones and antherids arise. These also may be in
terminal clusters of two or more, or single, or may appear to be lateral.
The obovoid oogones contain each a single egg with abundant periplasm.
The clavate antherid is borne on a slender, sometimes coiled hypha arising
from just below the point of attachment of the oogone. It becomes
attached to the oogone at the apex.

Rhipidium, with four species, consists of a thick, more or less cylindri-
cal body with numerous rhizoids, growing on fruits, twigs, etc., in water.
At its top it gives off slender branches which are constricted here and
there. Terminally on these slender branches arise the ovoid zoosporangia
which may later appear lateral on account of the sympodial mode of
growth of the hypha. The protoplasmic contents of the zoosporangium
divide into numerous zoospores which push out into a cylindrical vesicle.
Upon the rupture of the latter the biflagellate, kidney-shaped zoospores
escape. The oogones are also terminal on slender branches and contain
each a single egg surrounded by a layer of periplasm. The antherid may
arise on a slender branch just below the oogone or on a longer branch
from another plant. It attaches itself to the basal portion of the oogone.

118

PHYCOMYCETEAE

Fig. 39. Saprolesniales, Family Rhipidiaceae. (A-D) Rhipidium americanum
Thaxt. (A) Habit sketch. (B) Detail of branch bearing zoosporangia. (C) Discharge
of zoospores. (D) Two oogones, each with a mature oospore. (E-G) Araiospora
pulchra Thaxt. (E) Portion of plant bearing an oogone and zoosporangia of both
types (F) Oogones and antherids. (G) Ordinary and spiny zoosporangia. (After
Thaxter: Botan. Gaz., 21(6):317-331. Univ. Chicago Press.)

KEYS TO THE FAMILIES AND GENERA OF LAGENIDIALES 119

The oospore is thick-walled and roughened areolately. Its manner of
germination has not been reported. (Fig. 39 A-D.)

Araiospora, with four species, grows on vegetable matter in water. It
has a thick supporting or storage body from which arise the numerous
slender constricted branches bearing the reproductive organs. The zoospo-
rangia are of two kinds, ovoid or club-shaped ^^•ith smooth walls, and
subspherical and covered with stout spines. The zoospores are similar in
each type, being biflagellate and kidney-shaped. The spherical oogones
are borne on similar branches, sometimes on distinct plants. They are
characterized by a cellular periplasm layer around the oospore. The
oogone is fertilized by a basally applied antherid which may arise near by.
(Fig. 39 E-G.)

Mindeniella also has zoosporangia of two kinds. The thin- walled
clavate or ovoid sporangia are pedicellate on or near the apex of the stout
cylindrical or clavate basal segment. They may be spiny at the upper end
or without spines. The resting zoosporangia are ovoid or almost spherical
and pedicellate. They are spiny and thick-walled. Their germination has
not been observed. No sexual organs are known.

It is apparent that the Rhipidiaceae form a group of more highly
specialized genera which have probably arisen from the Leptomitaceae.
Whether the latter arose from the Saprolegniaceae or vice versa is uncer-
tain. In view of the somewhat more specialized sexual organs in the
genus Apodachlya it is possible that this family arose from the Sapro-
legniaceae or that both arose from the Lagenidiales. However it is also
possible that the latter represent ends of series of reductions from various
ancestral forms in the Saprolegniales and possibly in the Peronosporales.
Because of the difference in zoospore structure and in the composition of
the cell wall it seems unlikely that these orders have any close relationship
with the Chytridiales-Blastocladiales-Monoblepharidales series.

Keys to the Families and Genera of Lagenidiales

Key to the Genera of Family Woroninaceae

I" Plasmodium " forming a cluster ("sporangiosorus") of numerous cellulose-
walled zoosporangia or a cluster ("cystosorus") of thick-walled, angular resting
spores. Parasitic in Saprolegniaceae and green algae. Woronina

Plasmodium" becoming surrounded by cellulose walls forming an elongated
and sometimes branched thin-walled tube. The contents separate into naked
"spore mother cells" each of which produces 8 zoospores. Resting spores
unknown. Saprophytic in marine Florideae. Pyrrhosorus

'Plasmodium" at maturity filling the infected portion of the host and forming
its wall pressed closely against that of the host. Forming a single more or less
rounded zoosporangium or a row of cylindrical zoosjiorangia. Resting spores
where known round and spiny and free from the cell wall of the host. Parasitic
in Pythiaceae and Saprolegniaceae. Rozellopsis

120 PHYCOMYCETEAE

Key to the Genera of Family Olpidiopsidaceae

Zoospores escaping through discharge tubes.
Zoospores monomorphic (monoplanetic).
Zoosporangia spherical to eUipsoid, mostly with one discharge tube.
Resting spores produced, usually by a sexual process. Olpidiopsis
Resting spores unknown. Pseudolpidium

Zoosporangia irregularly lobed or tubular, usually with more than one dis-
charge tube. Resting spores unknown. Petersenia
Zoospores dimorphic (diplanetic), encysting at the mouth of the discharge tube
and then escaping singly. Zoosporangia spherical or nearly so. Fertiliza-
tion of oogone by a conjugation tube. Oogone with one egg and peri-
plasm. Pythiella
Zoospores escaping by a large irregular break in the wall of the spherical or
elUpsoidal zoosporangium. Resting spores not certainly known.

Pseudosphaerita

Key to the Genera of Family Sirolpidiaceae

Thallus elongate, narrowly tubular, becoming septate and then disarticulating
into separate zoosporangia. Sirolpidium

Thallus elongate, broadly tubular, becoming septate into distinct zoosporangia
which remain attached. Pontisma

Key to the More Important Genera of Family Lagenidiaceae

Thallus elongated, not becoming septate, sometimes coiled, parasitic in roots of
grasses and other plants. Zoospores formed in a vesicle at the tip of the single
exit tube. Sexual reproduction by union of contents of adjacent thalli through
a conjugation tube to form a thick-walled resting spore.

Lagena
Thallus short or elongated, not septate, sometimes branched, in the cells of
diatoms. Primary zoospores encysting at mouth of exit tube and emerging
as zoospores of the secondary type.
Thallus short, not branched, primary zoospores flagellate. Sexual reproduction
by union of adjacent thalli through a permanent conjugation tube, the empty
male thallus remaining attached.

Edrogella
Thallus elongated and coiled, sometimes branched. Primary zoospores not
flagellate. Resting spores formed here and there within the elongated thallus
but no sexual process observed.

Aphanomycopsis
Thallus more or less rounded, parasitic in the exterior cells of Phaeophyceae or
Rhodophyceae. Resting spores not observed.
Sporangia becoming extramatrical at maturity. Primary zoospores encysted
at the mouth of the exit tube or more often against the inner surface of the
zoosporangium. Upon emergence they are of the secondary type.

Eurycha&ma
Sporangia remaining intramatrical with many discharge tubes. Primary
zoospores encysting near the mouths of these tubes and emerging as sec-
ondary zoospores. Eurychasmidium
Thallus elongated and coenocytic, mostly later divided into multinucleate seg-
ments by cross septa. If the latter are wanting perhaps the species should be
transferred to the genus Lagena. Each segment becomes a zoosporangium

KEYS TO THE FAMILIES AND GENERA OF SAPROLEGNIALES 121

or a gametangium. Zoospores forming in a vesicle at the tip of the exit tube.
In fresh-water algae, pollen grains, and microscopic aquatic animals, etc.

Thallus strongly constricted at each septum. Antheridial cell poorly differ-
entiated. Myzoajtium

Thallus not, or only slightly, constricted at the septa. Antherids sharply dis-
tinguished from the oogones, with well-developed conjugation tube. Some-
times septa fail to be formed.

Lagenidium

Thallus and sexual reproduction much as in Lagenidium, but no zoospores
formed. Aerial conidiophores discharge the single conidia violently from the
apex. See Ancylistes

in Order Entomophthorales (Chap. 7)

Key to Family Thraustochijtriaceae
Single genus. Epibiotic zoosporangium with rhizoids. Thraustochytrium

Keys to the Families and Genera of Saprolegniales

Key to the More Important Genera of Family Saprolegniaceae

(Based in part upon Coker and Matthews, 1937)

Fertilization of oogones reportedly by uniflagellate sperm cells, otherwise as in
Saprolegnia. Probably a species of the latter with Chytridiaceous
or Protozoan parasites. Archilegnia

Oogone fertilized by male nuclei introduced from adhering antherids; develop-
ment sometimes parthenogenetic.
Sporangia rare or wanting, the spores encysting in the sporangium without a
swimming stage and germinating by germ tubes, very rarely by
swimming cells. Aplanes

Sporangia abundant, the spores encysting within them.
Oogones usually with more than one egg.

Encysted spores liberated by the irregular rupture of the sporangial wall,
then germinating by germ tubes or by emergence of zoospores of
the secondary type. Thraustotheca

Encysted spores liberated in successive groups by the breaking off of the
apical portion of the sporangium, later germinating by emergence
of zoospores of the secondary type. Calyptralegnia

Encysted spores escaping as zoospores of the secondary type through
exit papillae which pierce the zoosporangium wall. Mycelium very
slender, growing on keratinized media. Aphanodictyon

Oogones with only one egg.

Mycelium of vigorous and extensive growth; encysted spores in several
rows producing a net-like appearance, germinating by germ tubes
or more often by emergence of zoospores of the secondary type.

Didtjuchus
Mycelium of very limited growth, dense and opaque.

Encysted spores in one to several rows, thin-walled, escaping by disso-
lution or rupture of sporangial wall, usually germinating by germ
tubes, more rarely by emergence of zoospores of the secondary type.

Brevilegnia
Encysted spores in one row, thick-walled, multinucleate, never forming
a swimming stage. Geolegnia

122 PHYCOMYCETEAE

Sporangia abundant, the spores emerging from an apical mouth before encyst-
ing.
Zoospores of primary type only, after escaping from sporangium germinating
by germ tube or by repetition forming zoospores again of the pri-
mary type. Pythiopsis
Zoospores of primary type swimming some distance before encysting, then
emerging as secondary type zoospores.
Zoospores in a single row in the zoosporangium. Leptolegnia
Zoospores in more than one row.

New sporangia formed by proliferation. Saprolegnia

New sporangia formed by cymose branching. Isoachlya

Zoospores of primary type with or without flagella and encysting at the
mouth of the zoosporangium, later emerging as secondary type
zoospores.
Zoospores in more than one row in the sporangium.

Zoospores all encysting at the mouth of the zoosporangium. New spo-
rangia formed by cymose branching. Achlya
Zoospores encysting in part at the mouth of the sporangium and some
swimming away before encysting. New sporangia formed by cymose
branching and sometimes also by proliferation. Protoachlya
Zoospores in one row in the slender zoosporangium.

Masses of lobulate inflated segments auxiliary to the zoosporangia are

produced. Plectospira

No lobulate auxihary masses.

Branches of mycelium of usual type. Aphanomyces

Spike-like'^^branches' with sticky tips which catch and parasitize
rotifers. Sommerstorfia

Key to the Genera of Family Leptomitaceae

(Based upon Sparrow, 1943)

Thallus coarse, branched, constricted into numerous cjdindrical segments. Be-
ginning at the apex the segments become zoosporangia in basipetal succession
without change of size or shape. No sexual reproduction known. Growing
saprophytically, in heavily polluted water. Leptomitus

Thallus more slender, constricted, the zoosporangia with definite pedicels and
mostly ovoid or pyriform. Usually producing sex organs.
Oogones with one oospore. Apodachlya

Oogones with more than one oospore; zoosporangia not reported.

Apodachlyella
Key to the Genera of Family Rhipidiaceae

Basal cells giving rise to branches which bear the reproductive organs.

Basal cells slender, sporangia smooth walled, oospores with undulate outer wall.

Sapromyces
Basal cells usually stout, sporangia smooth walled or spiny or both in the
same species. Oospores with reticulate or cellular outer wall.
Oospore wall cellular, both smooth and spiny sporangia present.

Araiospora
Oospore wall reticulate, sporangia with smooth walls. Rhipidium

Reproductive organs arising directly from the basal cell and of varying degrees
of spininess. Mindeniella

LITERATUEE CITED 123

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Bartsch, Alfred F., and Fred T. Wolf: Two new Saprolegniaceous fungi,
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Bishop, Harlow: A study of sexuality in Sapromyces Reinschii (Schrot.) Fritsch,
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CoKER, William Chambers: The Saprolegniaceae with Notes on Other Water
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Pis. 4-5. 1924.
: Notes on the genus Aphanomyces, with a description of a new semi-
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40(160) :849-881. Pis. 35-38. Figs. 1-3. 1926b.
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124 PHYCOMYCETEAE

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PHYCOMYCETEAE: PERONOSPORALES
AND PROTOMYCETALES

Order Peronosporales. The fungi discussed in the previous chapters
were mainly inhabitants of soil and water, and largely saprophytic or
parasitic in habit. This order contains many genera and species that are
very strict parasites on Flowering Plants (Anthophyta or Angiosperms)
and which have hitherto resisted all attempts to grow them on artificial
culture media. However, most if not all of the members of the family
Pythiaceae respond more or less readily to efforts to bring them into cul-
ture upon nonliving media. Some genera of this family were placed in
the past among the Saprolegniales because of characters intermediate
between the two orders.

In the main the asexual reproduction of the Peronosporales exhibits
a step in evolution above that in the Saprolegniales. In the latter the
zoospores are mostly produced from zoosporangia that remain attached
to the main mycelium when the zoospores are set free. However, it must
be noted that the hyphae may break up into separate rounded segments
called "gemmae" which eventually are capable of functioning as zoo-
sporangia. In the majority of Peronosporales the tips of aerial hyphae
enlarge and are separated from the main hypha by a cross wall and are
then set free and distributed, usually by air currents. These so-called
"conidia" show their true nature as potential zoosporangia when they
fall into water, as then tlioir contents may divide internally into zoo-
spores. The zoospores produced in this order are always of the secondary
type. These zoospores may in some cases encyst and become free-swim-
ming several times.

Sexual reproduction occurs by the fertihzation, through a conjugation
tube from an antherid, of the normally single egg in an oogone to form
a thick-walled oospore. With a few possible exceptions the egg is sur-
rounded by periplasm as in the Rhipidiaceae of the order Saprolegniales
and Pythiella in the order Lagenidiales.

There is no sharp distinction of root-like holdfast hyphae and external

126

OEDER PERONOSPORALES 127

hyphae of quite different nature as occurs in many of the Saprolegniaceae
and in the Rhipidiaceae. The mycehum usually shows a cellulose reaction
upon application of a solution of chloriodide of zinc. The hyphae are
mostly much more slender than in the above-mentioned families but, like
them, are normally coenocytic and nonseptate except to set off repro-
ductive organs or injured portions or sometimes empty portions of the
mycelium.

Following Fitzpatrick (1930) the 400-500 species are divided into
three families as follows:

Pythiaceae: saprophytic or parasitic, mycelium intracellular, less often inter-
cellular with haustoria. Zoosporangia filamentous to ovoid or spherical, in
some cases the ovoid forms becoming separable "conidia," if aerial. These
conidia are borne singly, rarely in chains, at the tips of unbranched coni-
diophores or at the tips of the branches of a sympodially branched coni-
diophore. They germinate by the formation of zoospores or by a germ tube.
The oospore germinates by becoming a zoosporangium or by forming a short
hypha terminated by a conidium or by producing mycelium.

Albuginaceae: strictly parasitic in herbaceous Anthophyta (Angiospermae).
Mycelium intercellular with globular haustoria. Conidiophores clavate,
clustered in extensive sori under the epidermis of the host which is ruptured
by the pressure of the conidia which are produced successively in chains at
the apex of each conidiophore. Conidia germinating by the formation
of zoospores or of germ tubes. Oospore germinating by the formation of
zoospores.

Peronosporaceae: strictly parasitic in herbaceous, and, more rarely, woody
Anthophyta (Angiospermae). Mycelium intercellular with globular or fila-
mentous haustoria. Conidiophores emerging through the stomata singly or
two or three together, unbranched or monopodially much branched, bearing
the conidia singly at the tips of the branches or on short sterigmata on the
top of the unbranched conidiophore. Conidia germinating by the formation
of zoospores or by a stout germ tube. Oospore germinating by a germ tube
or by a short unbranched conidiophore.

Family Pythiaceae. The Pythiaceae have been variously divided
into from 4 to 14 genera. Of these, Pythium (in its wider delimitation) and
Phytophthora contain the greater number of species and are best known.

The species of Pythium are mostly soil or water inhabitants, probably
living most of the time as saprophytes. Some of these soil species, how-
ever, are capable of becoming destructive parasites upon plants, causing
rotting of the tissues or damping-off. A few are parasitic on other species
of the genus. Some species (e.g., P. proliferum de Bary, which can be
caught by hanging fruits in bodies of water) are saprophytic upon various
vegetable objects submerged in water. Of the 66 species recognized by
Middleton (1943) three are saprophytic and one parasitic on animal
matter, nine parasitic in fresh-water algae, and one saprophytic in a
marine red seaweed, about nine in soil and vegetable debris and not
known to be parasites, and forty or so capable of parasitism in higher

128

PHYCOMYCETEAE

Fig. 40. Peronosporales, Family Pythiaceae. (A-D) Pythium dictyosporum Racib.
(A) Formation of zoospores within vesicle at the open tip of the discharge tube of the
filamentous sporangium. (B) Zoospores. (C) Young oogone and antherid. (D) Mature
oogone and oospore. (E) Pythium torulosum Coker and Patterson, showing the toruloid
zoosporangium and vesicle. (F, G) Pythium proliferum de Bary. (F) Spherical zoo-
sporangium with vesicle. (G) Proliferated zoosporangium. (A-D, courtesy. Sparrow:
Mycologia, 23(3):191-203. E-G, courtesy, Matthews: Studies on the Genus Pythium,
Chapel Hill, Univ. North Carolina Press.)

plants, mainly roots and stems and in one species in the leaves. In the
forms parasitic in land plants the mycelium is mainly intracellular, usu-
ally killing the cells rapidly, in some cases appearing to kill them in
advance of the arrival of the hyphae.

The mycelium is slender and, as in the Saprolegniales, is a branching
coenocyte. Septa are formed to set off the antherids and oogones and,
usually, the zoosporangia, and to separate the empty older portions of the
mycelium from those containing protoplasm. Sexual reproduction has
been studied by Trow (1901), Patterson (1927), Edson (1915), and others,
and has been found to resemble most closely that in the Rhipidiaceae.
The oogone arises terminally, less often intercalarly on longer or shorter
unmodified hyphae. It is ovoid or spherical and smooth or papillate or
even echinulate. Sometimes it is tubular-elongate at one or both ends.
Of the several nuclei in the oogone, all but one migrate to the periphery

ORDER PERONOSPORALES 129

into the periplasm which surrounds the centrally placed egg except in the
rare cases in which two to six eggs are produced. In P. polysporum Poitras
(1949) nearly one fourth of the oogones produce from two to six oospores.
The antherid is also plurinucleate at first but only one nucleus functions in
fertilization of each egg. The filament bearing the terminal antherid may
arise at a great distance from the oogone ("diclinous") or from near by
on the hypha bearing it ("monoclinous"). In some cases the antherid is a
small segment of the hypha immediately under the oogone, the conjugation
tube entering the latter through the base ("hypogynous"). In a few
species the antheridial filament wraps in a spiral around the hypha bear-
ing the oogone. Often the oospore develops parthenogenetically. Some-
times many antherids may become attached to the same oogone although
only one antherid functions in its fertilization. Through a conjugation
tube one male nucleus is introduced into the egg which then forms a
thick wall and becomes an oospore. The oospore may completely fill the
oogone ("plerotic") or may leave a space between it and the oogone wall
("aplerotic"). It is usually thick-walled and smooth but may be reticu-
lately thickened externally. Germination may be delayed for a long
while. It is effected by the production of a long germ ^be or of a short
germ tube terminating in a zoosporangium, or in some cases the oospore
may produce the zoospores internally, emptying them through a short
beak or exit tube into a vesicle as in the germination of a conidium
(Drechsler, 1947). (Fig. 40.)

Asexual reproduction is mostly by means of zoospores. Typically the
zoosporangium forms a beak (usually short but sometimes several times
as long as the diameter of the zoosporangium). The tip of the beak softens
and out of it flows the protoplasm of the zoosporangium, to form a spher-
ical mass, the scncalled vesicle. Within this the differentiation into zoo-
spores is completed and the kidney-shaped biflagellate zoospores rupture
the plasma membrane of the vesicle and escape singly, or several in a
clump w^hich separate subsequently. After swimming for a while the zoo-
spore encysts and germinates by a germ tube or, as apparently first
reported by Cornu (1872), sends out a short papilla from whose apical
opening emerges a single zoospore, with or without a vesicle, or sometimes
a vesicle in which are produced several zoospores. The process may be
repeated three or four times, the zoospores of the successive crops being
smaller each time.

Three more or less distinctive types of zoosporangium may be dis-
tinguished as follows:

1. Slender filaments, simple or branched, of the same size and appear-
ance as the vegetative mycelium, and opening at the tip or tips of the
hyphae to form a short or long emission tube at whose apex a vesicle is
formed. This filamentous zoosporangium may be separated from the re-

130 PHYCOMYCETEAE

mainder of the mycelium by a septum (subgenus Nematosporangium of
Fischer, 1892) or the septum may be lacking (subgenus Aphragmium of
Fischer). (Fig. 40 A, B.)

2. The more or less filamentous zoosporangium is somewhat thickened
and lobed, often only near the base, to form the so-called toruloid struc-
ture, the plasmatoonkosis of Sideris (1931). This is a storage organ ena-
bling the zoosporangium to produce larger numbers of zoospores in the
vesicles at the tips of the emission tubes. (Fig. 40 E.)

3. The zoosporangium is spherical, ovoid or limoniform, terminal or
intercalary, single or in short chains {Sphaerosporangium of Fischer).
(Fig. 40 F.)

Schroeter (1897) and Sideris (1931) included types 1 and 2 in a genus
which they called Nematosporangium, applying the name Pythium to
Fischer's subgenus Sphaerosporangium. Inasmuch as the type species of
the genus, P. monospermum Pringsheim (1858), forms zoosporangia of
type 1, the name Pythium must be retained for Schroeter's Nematospo-
rangium, if the genus is divided. The later monographers of the genus
(Butler, 1907; Miss Matthews, 1931; and Middleton, 1943) do not make
this division but include all three types of sporangia in one genus.

In some submerged species with spherical or ovoid sporangia new
zoosporangia arise within the emptied ones by proliferation as in Sapro-
legnia, or a branch arises just below the point of attachment of the zoo-
sporangium and is terminated by a new one, sympodial development
proceeding as in Achlya. In some species the zoosporangia may be pro-
duced on aerial hyphae in which case they may become detached and
distributed by currents of air, germinating when they fall into water
either in the usual manner or by a germ tube. These wind-carried de-
tached zoosporangia are called conidia. Usually the submerged zoo-
sporangia and the aerial conidia are alike in the same species. Pythium
deharyanum Hesse, with its zoosporangia representing potential conidia,
is common in soil as a damping-off parasite of seedlings when the soil is
too moist or the seedlings too much crowded. It can often be obtained by
placing a little soil in a dish with some cooled boiled water and placing
in the latter a few boiled hemp seeds (not too many or the bacteria will
become numerous). On these seeds Pythium and various Saprolegniales
will appear in a few days. Pythium aphanidermatum (Edson) Fitzpatrick,
the cause of black-root disease of radishes (especially noticeable in the
White Icicle variety) and rotting of sugar-beet seedlings and roots of
many other plants, is a species with the second type of sporangia. It was
originally described as the type of a new genus, Rheosporangium, by
Edson (1915).

The genus Phytophthora contains 15 or 20 species some of which live
as saprophytes in the soil, developing as parasites in the presence of suit-

ORDER PERONOSPORALES

131

-m^'-i

Fig. 41. Peronosporales, Family Pythiaceae. Phytophthora cactorum (L. & C.)
Schrot. (A) Aerial filament in moist air, producing sympodially a succession of spor-
angia ("conidia"). (B) Formation of zoospores within the sporangium and emission
mto a vesicle. (C) Fertilization of oogone. (D) Mature oospore. (E) Germinating
oospore. (Courtesy, Blackwell: Brit. Mycol. Soc. Trans., 26(1-2) :71-89.)

132 PHYCOMYCETEAE

able host plants. Other species are usually found as parasites on higher
plants, although even these are capable of cultivation on nonliving cul-
ture media. The zoosporangia of the soil-inhabiting types may be sub-
merged and then remain attached to the mycelium, or may emerge into
the air in which case they may become detached and wind-distributed.
In the more strictly parasitic forms the conidiophores emerge through
the epidermis of the host, piercing it or passing through the stomata.
They are simple or may branch sympodially. In all species the zoospo-
rangium or the conidium when it falls into water may produce zoospores
which emerge singly or pass into a vesicle which soon ruptures so as to
allow them to escape. They are of the secondary type and resemble those
of Pythium. They swim for a while and then encyst. They germinate by
a germ tube. Instead of producing zoospores the conidia may produce
short conidiophores upon which one or more small conidia arise or may
germinate by a long germ tube. Sometimes the conidia germinate in situ
without becoming detached. In P. infestans (Mont.) de Bary the my-
celium overwinters in infected tubers which then give rise to diseased,
spore-bearing shoots which serve as centers of infection. (Fig. 41.)

As in Pythium the mycelium of many species is intracellular, directly
killing the invaded cells, but in some species it also grows intercellularly,
sending haustoria into the adjacent cells (Cooper and Porter, 1928;
Szymanek, 1927; Klebahn, 1909; Butler, 1910). In general the species
act as destructive parasites, killing the tissues very rapidly.

Sexual reproduction is essentially like that in Pythium, the single egg
surrounded by periplasm being fertilized by a male nucleus which passes
from the adhering antherid into the egg through a conjugation tube.
The antherid may arise as a separate branch and become attached to the
oogone at any point. In some species the antherid appears to surround
the base of the oogone. This has been interpreted as an antherid coiled
around the hypha bearing the oogone (Shanor, 1938), but Pethybridge
(1913) claims that development is as follows: On the end of a hypha an
antherid is formed and through this another hypha grows, piercing the
antherid completely and swelling then to form the oogone above it. B. D.
Mundkur (1949) confirmed the occurrence of this amphigynous type
of antherid in another species (Phytophthora himalayensis Dastur). In
some species oogones may be found with these basal (amphigynous)
antherids and others with antherids in the lateral position (paragynous).
After fertilization the thick-walled oospore rests for some time and then
germinates to form mycelium or a short conidiophore. Infection of the
host may occur by the zoospores produced in the conidia or by the germ
tubes from the oospores. With the soil-inhabiting species the mycelium
may directly penetrate the subterranean portions of the host. Narasinhan

(1930) reports that P. arecae (Golem.) Pethyb. is heterothallic but Tucker

(1931) believes that this needs further study. (Fig. 42.)

ORDER PERONOSPORALES

133

Fig. 42. Peronosporales, Family Pythiaceae. Phijtophthora stellata Shanor. (A)
Habit sketch of hypiia with numerous sporangia and a sexual reproductive branch.
(B-E) Successive stages in the development of the same antherid and oogone, showing
possible explanation of the so-called amphigynous development. (F) Fertilization of
oogone. (Courtesy, Shanor: /. Elisha Mitchell Sci. Soc, 54(1):154-162.)

Phytophthora infestans (Mont.) de Bary, the cause of the late bhght
of potato {Solarium tuberosum L.), tomato {Ly coper sicon esculentum Mill.),
and rot of potato tubers, was first observed as a serious enemy of the
potato about 1845. After being studied by various investigators it was
first fully described by de Bary in 1876. For many decades oogone forma-
tion was unknown in this species until Clinton in 1911 reported their
production in culture on oat agar. In 1927 Murphy reported finding them
on the surface of tubers and in the surrounding soil. As in some other
species these oospores were mostly parthenogenetic in origin although
a basal antherid was observed in one case. In P. phaseoli Thaxter, the
oogones were shown by Chnton (1906) to be produced in the seeds while
the conidiophores covered the surface of the pods of the lima bean (Phase-
olus Umensis Macf.). P. cactorum (L. & C.) Schrot. and some other species
are troublesome rot-producing and damping-off fungi of many kinds of
cultivated plants.

The border line between some of the root-inhabiting species of Phy-
tophthora and some of the conidium-producing species of Pythium is so

134

PHYCOMYCETEAE

vague that a few mycologists, e.g., Fitzpatrick (1923), have suggested
uniting the two genera, which would then have to take the older name
Pythium. However the two genera may still be kept separate on the basis
of the method of zoospore formation: within the zoosporangium (con-
idium) in Phytophthora and in an external vesicle in Pythium. In the rare
cases where a vesicle is formed in the former the zoospores are produced
within the zoosporangium before passing into the vesicle. Thomas (1942,
1943) made a chemical investigation of the composition of the cell walls
of these two genera. In both of these the major part of the wall consists
of cellulose with an outer deposit of fatty substance. Outside of this there
is a layer of some pectic compound in the case of Pythium and of some
other carbohydrate in Phytophthora. After the carbohydrates and fatty
substances are all dissolved the hyphae still retain their form and this
residual matter proved to be chitin. This is particularly interesting in
view of the conclusions of von Wettstein (1921) that cellulose and chitin

Fig. 43. Peronosporales, Family Pythiaceae. Pythiogeton transversum Minden. (A)
Portion of plant with a zoosporangium not quite mature, a zoosporangium discharging
its contents, and an empty zoosporangium. (B) Portion of the discharged contents
separating into zoospores. (C) Young oogone and antherid. (D) Oogone and antherid
after fertilization. (E) Mature oospore. (After von Minden: Mykologische Unter-
suchungen and Berichtc, 1(2). •146-255.)

ORDER PERONOSPORALES 135

are mutually exclusive. Schroter (1897), recognizing the similarity to the
Saprolegniales, placed Pythium in that group, but retained Phytophthora
in the Peronosporales. This cannot be upheld in view of the closeness
of the two genera, but merely shows the difficulty of drawing sharp
delimiting lines in some cases. A number of other genera have been de-
scribed in this family, some being soil-inhabiting or aquatic saprophytes
and others being serious parasites of various economic plants. By some
mycologists they are merged with the genus Phytophthora and by others
are maintained as distinct genera.

Pyihiogeion, described by von Minden (1916), is an aquatic saprophyte
whose elongated or sac-like zoosporangium has its axis more or less
transverse to the hypha on which it is borne terminally or in an inter-
calary position. The zoospores are expelled in a mass which seems to lack
the tough plasma membrane characteristic of the vesicle of Pythium and
soon breaks up into the individual zoospores. Where known the oospores
have very thick walls. (Fig. 43.)

Other genera assigned to this family are Diasporangium (Hohnk,
1936), a soil-inhabiting parasite, Trachysphaera (Tabor and Bunting,
1923), etc. Pythiomorpha was described by Petersen (1910) and con-
sidered by him to be worthy of the establishment of a separate family.
Several species have been ascribed to this genus but recent studies by
Blackwell, Waterhouse, and Thompson (1941) seem to indicate that
these all represent various species of Phytophthora growing in water.

Family Albuginaceae. This family consists of the single genus
Albugo or, as it is often called, Cystopus. In view of the fact that the
former name was given in 1821 and the latter in 1847 the latter must be
abandoned. The species number about twenty-five. In contrast with the
members of the foregoing family the species are strictly parasitic, never
occurring as saprophytes. They do not lend themselves to cultivation on
culture media. Within some species — e.g., A. Candida (Pers.) Kuntze —
there are numerous specialized races that are adapted only to certain
host species or groups of species. The mycelium is strictly intercellular
except for the small globular haustoria which are borne on the ends of
short, very slender processes which pierce the host cell wall. Melhus
(1915) has shown that the mycelium of A. Candida may overwinter in the
tissues at or below the crown of such host plants as are winter annuals
or biennials, growing out into the new shoots in the spring. The conidio-
phores are formed on ends of short sympodially branching hyphae which
arise from a mass of mycelium gathered in a limited area underneath the
epidermis of the host (the so-called sorus). They are club-shaped and
stand, closely packed together, perpendicular to the surface of the epi-
dermis, between it and the subepidermal cells. From the apex of each
conidiophore are abstricted successively the spherical or ovoid pluri-

136

PHTCOMTCETEAE

e^M^:^Q

Multinucleate egg about to be i.ri^Z^^^^^Ty p riplasm'contZ '"^- ^^^
numerary nuclei. (C) Antherid and multinucleate conj^,gat7on "be^D S^fr''^^"
mature oospore, showing numerous nuclei. (E) ^/i^^oZSacalrDr Pi . ""^

and conjugation tube each with a single functional mfp^.,!? ^ '^ ^y^^tz^, egg
remaining in antherid and in the periplasm rAD^Z^ Supernumerary nuclei
28(3):149-170, Univ. Chicago Press. l^^^^^^^.L^i.:^^^^^^ S^a.)^^^'

OKDER PERONOSPORALES 137

nucleate conidia which are separated from each other by slender connec-
tions, the disjunctors, whose dissolution permits the conidia to fall apart.
The chains of conidia thus formed raise and eventually rupture the over-
lying epidermis, permitting the conidia to escape and to be distributed
by air currents. The similarity of these sori, except for the color, to those
of Rusts led to the name "White Rust" often applied to fungi of this
genus. Upon falling into water the conidia divide internally into several
uninucleate, biflagellate, kidney-shaped zoospores which escape by disso-
lution of a special spot in the conidial wall. After swimming for a short
time these zoospores encyst and germinate by a germ tube. In some
species Palm (1932) has shown that the conidia germinate usually by
the production of a stout germ tube which infects the host without pro-
ducing zoospores. Sexual reproduction takes place in the tissues of the
host. Often the portions of the host plant in which this occurs are much
hypertrophied. This is especially the case with Albugo Candida in which,
the inflorescence and individual flowers of the host may be much thick-
ened and enlarged. The distorted flowers remain green and are some-
times several times as large as the normal flowers. On the ends of hyphal
branches the almost spherical oogones are separated from the hyphae by
septa. Stevens (1899, 1901) studied the process of fertilization in several
species. The oogone is at first multinucleate, the number of nuclei
being as high as 300. These may all pass to the periplasm leaving but a
single egg nucleus in the egg or after passing to the periplasm they may
divide, half of the daughter nuclei remaining in the periplasm and the
other half in the egg so that eventually the latter may contain 100 or
more nuclei. The multinucleate antherid on the end of a hyphal branch
attaches itself to the oogone and eventually sends into the egg a conju-
gation tube through which one male nucleus passes, in the first case
mentioned above, or 100 or more in the second case. These male and
female nuclei fuse by pairs. The fertilized egg produces a thick wall, con-
sisting of a thin endospore and a thick roughened epispore. In the first
type of fertilization the zygote nucleus divides repeatedly so that the
oospore overwinters as a multinucleate structure. In the spring zoospores
are formed and the epispore is ruptured, the endospore pushing out
through the break as a bladder which in its turn ruptures and permits
the zoospores to escape. Just where meiosis occurs is not yet certain. It
has been suggested that the nuclear divisions occuring in the antherid
and oogone before fertilization represent this process, or it may occur in
the first nuclear division in the fertilized egg. In North America the
common species are A. Candida (Pers.) Kuntze, on various crucifers
(Brassicaceae) ; A. portidacae (DC.) Kuntze, on purslane {Portidaca ole-
racea L.);A. bliti (Biv.-Bern. ) Kuntze, on various species of Amaranthus; A .
tragopogonis (DC.) S. F. Gray, on salsify {Tragopogon porrifoliusL.) and

138

PHYCOMYCETEAE

Fig. 45. Peronosporales. (A) Family Albuginaceae. Albugo portidacae (DC.)
Kuntze. Conidiophores and conidia. (B-F) Family Peronosporaceae. (B) Basidiophora
entospora Roze & Cornu, conidiophores and mature oospore. (C) Rhysotheca australis
(Speg.) Wilson, conidiophore. (D) Peronospora ficariae Tul., conidiophore and oogone
with oospore. (E-F) Bremia lactucae Kegel. (E) Tip of conidiophore branch. (F)
Conidia showing germination by germ tube and by zoospores. (A-E, after Berlese:
Icones Fungorum, Padua. F, after Milbrath: /. Agr. Research, 23(12) :989-994.)

other composites; and A. ipomoeae-panduranae (Schwein.) Swing., on the
sweet potato (Ipomnra batatas (L.) Lam.) and related plants. In most
cases the disease caused is of minor importance. (Figs. 44 and 45 A.)

Family Peronosporaceae. The Peronosporaceae, Hke the Albu-
ginaceae, differ from the Pythiaceae in their strictly parasitic habit, the
mycelium always being intercellular with haustoria penetrating the ad-

ORDER PERONOSPORALES

139

Fig. 46. Peronosporales, Family Peronosporaceae. Rhysotheca viticola (B. & C.)
G. W. Wilson. (A) Germination of oospore to form one large conidium. (B) Zoospores
with flagella, and encysted and germinating. (C) Infection of host tissue through a
stoma. (After Gregory: Phytopathology, 2(6):235-249.)

jacent host cells. Like those of the Albuginaceae the haiistoria of this
^ family may be knob-like but in many species they are filamentous or
B finger-like. The conidiophores are external to the host and produce the
^conidia singly on the ends of the branches. These conidia are plurinucleate
^Band germinate in most cases by the formation of zoospores, as in Albugo.
^■In Peronospora and Bremia the typical mode of germination is by means
^■of a stout germ tube without the formation of zoospores. In Rhysotheca
■ (included in Plasmopara by many authors) the conidia germinate by the
Bformation of zoospores but in Plasmopara (in the narrower sense) the
whole protoplasmic contents of the conidium escape as a naked pluri-
nucleate but nonflagellate mass which quickly rounds up and encysts
and then germinates by a germ tube. This difference in the mode of ger-
mination is the basis for Wilson's (1907) division of Plasmopara into the
two genera. In nearly all genera in which germination by means of zoo-
spores is typical the conidia may, under special conditions, germinate
directly by germ tubes. A. de Bary (1876), Melhus (1915), Jones and
Torrie (1946), have shown that as in Albugo Candida, so also in this
family, particularly in the genus Peronospora, the mycelium can live over
winter in the tissues of a biennial, winter annual, or perennial host and
thus infect the new plants in the spring without the aid of conidia or
oospores. Sexual reproduction is like that in those species of Albugo in
which the mature oogone contains a uninucleate egg. Mostly the oospores

140

PHYCOMYCETEAE

germinate by a stout germ tube or by the formation of a conidiophore
terminated by a single large conidium (as reported by Gregory, 1912, in
Rhysotheca viticola (B. & C.) G. W. Wilson), sometimes by the direct
formation of zoospores. (Fig. 46.)

The six or more genera of Peronosporaceae are mainly distinguished
on the basis of their asexual characters. In the genus Basidiophora, with
two species parasitic on Composites (Family Asteraceae), the conidio-
phore is club-shaped with its slightly swollen
apex covered with numerous short sterigmata,
each bearing a nearly spherical conidium which
produces zoospores when it germinates. (Fig.
45 B.)

In Sclerospora the more or less dichoto-
mously branched conidiophores are much
thickened. The conidia germinate typically by
zoospores in aS. graminicola (Sacc.) Schrot., but
in most of the other species of the genus by
germ tubes. The oogone wall may remain thin
and lie closely against, but not grown fast to,
the thick-walled oospore (in S. graminicola
(Sacc.) Schrot., according to McDonough,
1937) or it may become thickened and some-
what folded, separate from the oospore wall
which has a thin wrinkled outer layer and a
thick inner layer (in S. macrospora (Sacc.)
McDonough, according to McDonough, 1947).
The thirteen or more species are chiefly par-
asites of grasses (Family Poaceae). In the
East Indies they cause serious injury to sugar
cane (Saccharum officinariim L.) and to Indian
corn or maize {Zea mays L.). Sclerospora
graminicola is found frequently throughout
the north temperate regions on foxtail grasses
(Setaria). The conidiophores form a downy layer on the under side of
the infected leaves early in the morning but quickly dry down as the air
becomes warmer and drier. After the oospores develop the leaves die and
shred longitudinally into thread-like strips on which the oospores may be
seen readily by the aid of a hand lens. Weston (1920, 1921, 1923, 1924)
has given various species of this genus very careful study. (Fig. 47.)

Plasmopara and Rhysotheca, usually united under the former name,
produce slender, much branched conidiophores whose branches arise
nearly at right angles. The tips of the branches are truncate. The two
genera differ, as mentioned above, by the mode of germination of the

Fig. 47. Peronosporales,
Family Peronosporaceae.
Sclerospora graminicola
(Sacc.) Schrot., conidio-
phore. (After Weston: J.
Agr. Research, 27(10) :771-
784.)

OEDER PERONOSPORALES 141

conidia: by zoospores in Rhysotheca and by a single naked mass in Plas-
mopara. The most important species from the economic standpoint is
R. viticola (B. & C.) G. W. Wilson, which causes the downy mildew and
brown rot of the foliage and fruit, respectively, of grape (various species
of Vitis).

Pseudopero?iospora (called Peronoplasmopara by some although the
former name has priority) has slender conidiophores branching at acute
angles and with pointed tips. The usually violet-tinged conidia germinate
by zoospores. Ps. cubensis (B. & C.) Rostow., first described from Cuba,
is probably native to Russia where its destructive effects have been known
for many years although the fungus was first recognized there in 1903 by
Rostowzew. It is a very serious enemy of the cucumber (Cucumis sativus
L.) and muskmelon (C. melo L.). Another species, Ps. ceUidis (Waite)
G. W. Wilson, attacks the hackberry {Celtis) while other species are found
on hemp {Cannabis), on hops (Humulus), and on nettle {Urtica). Hoerner
(1940) has shown that the species on hops {Ps. humuli Miyabe) is capable
of infecting Celtis, Cannabis, and Urtica and suggests that there is one
species infecting Urticaceae, perhaps with various physiological strains.

Bremia and Peronospora have slender conidiophores, branching at
acute angles in a more or less dichotomous manner. Their conidia ger-
minate typically by germ tubes, although zoospore production also has
been reported for Bremia by Milbrath (1923) and for P. spinaciae (Grew.)
Laub. by Eriksson (1919). Schultz (1937) could not observe zoospore
production in the strains of B. lactucae Regel growing in Germany. In
Bremia the tips of the branches enlarge into disk-like structures bearing
sterigmata on their edges. B. lactucae is sometimes destructive to lettuce
grown under glass. In Peronospora the tips of the branches taper to a
point. Gaumann (1923) recognizes 268 species of this genus. They are of
economic importance in but few cases. P. spinaciae is sometimes destruc-
tive in plantings of spinach {Spinacia oleracea L.) and P. parasitica
(Pers.) de Bary on various crucifers (Brassicaceae). (Fig. 45 D-E.)

Gaumann (1918a, b), Wartenweiler (1918) and others have shown
that the earlier recognized species of this family are separable by bio-
metric and cultural means into large numbers of closely related species
confined to very limited numbers of host species and differing constantly,
but only slightly, in the size and shape of the conidia and conidiophores.
When such studies have been extended to all parts of the world and to
all the forms occurring on different host species the number of species of
Peronosporaceae will doubtless be very greatly increased.

Several other genera have been described which may be found to be
justified. The physiology of conidial germination must be studied care-
fully in these as well as in the older genera. Only after such studies can
we be certain that some of the generic distinctions now maintained, or

142 PHYCOMYCETEAE

recently proposed, are really valuable. Should zoospore formation in
Peronospora be confirmed it would seriously weaken the distinction be-
tween that genus and Pseudoper'onospora.

The evolutionary tendencies within the Peronosporales are of interest
to students of phylogeny. The species of Pythium with long, narrow,
hypha-like zoosporangia would undoubtedly be included in the Sapro-
legniales were it not for those other species of Pythium with ovoid or
spherical zoosporangia which may even function as separable conidia,
thus forming a transition to Phytophthora. This genus still shows in some
species a close relationship to Pythium, in the production of submerged
zoosporangia and facultative saprophytic habits, while in other species
with well-developed conidiophores and strictly parasitic habits the genus
approaches closely the Peronosporaceae. The fact that the single egg in
the oogone is surrounded by periplasm, as in the Rhipidiaceae in the
Saprolegniales, would suggest that the relationship of the Pythiaceae is
closer to this family than to the Saprolegniaceae in which there is no
periplasm and the majority of species have numerous eggs in the oogone.
It has been suggested by some mycologists that from the Pythiaceae have
been derived on the one hand the Saprolegniales (through the Rhipi-
diaceae) and on the other hand the remainder of the Peronosporales.
As obligate parasitism became prevalent in the Peronosporales evolution
appears to have proceeded in several lines. The catenulate conidia of
Albugo call to mind the proliferating zoosporangia of some species of
Pythium as well as of Saprolegnia, while the sympodial conidiophores of
the more advanced species of Phytophthora remind one of the sympodial
branching in Achlya and some species of Pythium. The monopodia]
conidiophores of the Peronosporaceae do not resemble so closely any
structures in Pythium. As parasitism has progressed we also find the
transition from attached zoosporangia to separable zoosporangia (conidia)
leading finally to the conidium as found in Peronospora, in which zoospore
formation has been lost, although the plurinucleate condition persists.

The fungi that have been considered in the preceding chapters of this
book have been largely aquatic in habit or reveal their aquatic ancestry
by producing naked flagellate cells in some stages of their development,
although in some genera these have been suppressed. This has been true
even for the majority of the strictly parasitic species of the Peronosporales
which have abandoned the aquatic habit to assume that of parasitism in
land plants. Besides the production of zoospores the great majority of
the foregoing organisms, except most of the Chytridiales series, show the
cellulose reaction promptly upon the application of chloriodide of zinc
solution. Many of those which fail to show this reaction promptly do so
when certain masking substances are removed. The presence of true chitin
is demonstrated for only a minority of the species. The forms with well-

ORDER PROTOMYCETALES 143

developed hyphae show pronounced anisogamy in most cases. For the
latter reason the Monoblepharidales, Lagenidiales, Saprolegniales, and
Peronosporales were included in the subclass Oomyceteae of the Class
Phycomyceteae in the first edition of this book, following the practice of
Fitzpatrick (1930) and other mycologists. These were set apart from the
subclass Zygomyceteae, in which approximate isogamy was held to war-
rant such a distinction, accompanied as it was by loss of the power to
produce motile cells and by the greater predominance of chitin in the cell
walls. The fact that the simpler forms included in the Oomyceteae, such
as Chytridiales, and some of the Blastocladiales and Lagenidiales, show
isogamy and that many forms whose closest kinship seems to lie with
the Mucorales and Entomophthorales have pronounced anisogamy makes
it doubtful whether this character should be used to distinguish sub-
classes. Hence these two group names have been abandoned in this
edition.

Order Protomycetales. As a very doubtful appendix to the Phyco-
myceteae so far considered must be added the Order Protomycetales,
with a single family Protomycetaceae. The true position of this group
among the fungi has long been the subject of speculation. The author
follows Fitzpatrick (1930) in placing these fungi among the Phyco-
myceteae but with very little idea as to what groups of that class may
have given rise to them. The most recent and extensive investigations
on the group are those of Sappin-Trouffy (1897) and of von Biiren (1915,
1922). The family seems to have no affinity to the Ascomyceteae or to
the other Higher Fungi, though it has been assigned to various positions
among these. It consists of one well-established genus, Protomyces, with
12 or more species to which 3 other genera have been added, Taphridium,
Volkartia, and Protomycopsis. Until life history studies are more complete,
the validity of their segregation is uncertain. All the well-known species
of these 4 genera are parasitic in the stems, leaves, or fruits of Ammiaceae
(Umbelhf erae) , Asteraceae (Compositae), and Cichoriaceae.

A number of species have been described from various other families
but need further study before their validity is assured.

The mycehum is subepidermal or intercellular in the underlying
tissues. It is septate at occasional intervals, each segment being pluri-
nucleate. The cell walls give a strong cellulose reaction with chloriodide
of zinc. Some of the segments of the mycelium enlarge and become
multinucleate (30-40 nuclei, in Protomyces inundatus Dang.). Within the
original wall the cell enlarges and becomes surrounded by a thick, three-
layered cellulose wall, the nuclei dividing several times to become 100-200
in number. These resting sporangia (or "chlamydospores" as some
authors call them) may remain in the tissues over winter or may ger-
minate the same season. In some species they are subepidermal and in

144

PHYCOMYCETEAE

Fig. 48. Protomycetales, Family Protomycetaceae. (A-F) Protomyces macro-
sporus Unger. (A) Young mycelium with intercalary young resting sporangia. (B)
Germination of resting sporangium. (C) Protoplasm forming parietal layer of spore
mother cells. (D) Spore mother cell divided into spores. (E) Discharge of ball of
spores. (F) Successive stages in fusion of spores, showing union of nuclei in the conju-
gation tube and division of the zygote nucleus. (G) Protomycopsis leucanthemi Magn.,
terminally formed resting sporangium. (A-E, G, after von Biiren: Beitr. Kryptogamen-
flora Schweiz, 5(l):l-95. F, ibid., 5(3):l-94.)

others scattered at various depths. In germination the sporangia of
Taphridhim produce their spores without breaking the exospore, while
those of the other genera burst the exospore on one side and the contents
bulge out like a balloon, still surrounded by the inner wall. The nuclei in
Protomyces all migrate to the periphery and a large central vacuole is
formed. The thin layer of peripheral cytoplasm is divided by cleavage
planes, starting at the outside, into little uninucleate cells which divide
twice (meiosis?) forming four ellipsoidal spores out of each cell. These
spores then mass at the center or apex of the sporangium and by breaking
of the latter are thrown out along with the slimy contents of the vacuole.
In some species, apparently not in others, the spores fuse by twos after
being set free, sometimes before. The nuclei unite in the conjugation
tube. The spores in culture media germinate to form yeast-like cells but
when inoculated on the proper host produce endophytic mycelium. (Fig.
48.)

The affinities of this family are exceedingly uncertain. The final divi-

KEYS TO THE FAMILIES AND MORE IMPORTANT GENERA OF PERONOSPORALES 145

si on of the nuclei in the sporangium may be a reduction division. The
cells that divide into four spores each have been called asci and the whole
sporangium a "synascus" but the absence of nuclear fusion in these
"asci" before the formation of the spores seems to exclude that possi-
bility. The cellulose nature of the cell wall, it has been suggested, speaks
for the phycomycetous relationship. If related to the Lagenidiaceae we
must assume the loss of motility of zoospores or gametes. Relationship
to the Chytridiales is less likely because of the presence of well-developed
mycelium with cellulose walls. Baker, Mrak, and Smith (1943) suggest
that Coccidioides immitis Rix. and Gil., the fungus causing the disease of
man called coccidioidomycosis may possibly belong in the Protomycetales.

Keys to the Families and More Important Genera of Peronosporales

Key to the More Important Genera of Family Pythiaceae

Zoospores not preformed in the zoosporangium but developing in an extruded
mass which may or may not be enclosed in a plasma membrane.
Extruded mass of protoplasm not in a definite vesicle, zoosporangia elongated

transversely to the supporting hypha. Pythiogeton

Zoospores developed in a definite vesicle.

Hyphae bearing short lateral branches adapted -to the capture of the rotifers

upon which the fungus feeds. Zoosporangia hyphal. Zoophagus
No special hyphal branches for capturing prey. Zoosporangia hyphal or tor-
uloid or more or less spherical. Pythium

Zoospores preformed in the mostly more or less spherical zoosporangium (con-
idium) and escaping individually or sometimes in a temporary vesicle.

Phytophthora
No zoospores. External warted spherical conidia. Oogones with amphigynous
type of antherid. Of doubtful relationship. Trachysphaera

Key to the Genus of Family Albuginaceae

Only genus. Conidia catenulate on subepidermal conidiophores.

Albugo

Key to the More Important Genera of Family Peronosporaceae

Conidiophores clavate or somewhat cylindrical, somewhat swollen above with
numerous short sterigma-like branches. Conidia without appendages, ger-
minating by formation of zoospores. Parasitic on Composites.

Basidiophora
Conidiophores not much branched. Conidia with prominent beak and a basal
appendage consisting of the adhering upper end of the conidiophore.
Conidia germinating by production of zoospores. Possibly belonging to
the Pythiaceae. Kawakamia

Conidiophores with prominent branches.

Conidiophores stout, with heavy branches clustered near the apex, quickly
fugacious. Conidia germinating by germ tubes or by the formation of
zoospores. Parasitic on grasses. Sclerospora

Conidiophores more slender, branching monopodially, usually nearly at right
angles. Tips of branches obtuse.

146 PHYCOMTCETEAE

Conidia germinating by the formation of a single large naked nonflagellate
plasmatic mass which encysts and then germinates by a single germ tube.

Plasmopora
Conidia germinating by the production of biflagellate zoospores.

RJujsotheca
Conidiophores slender, more or less dichotomously branched, mostly at acute
angles. Conidia usually germinating by single germ tube, or by zoospores.
Tips of branches enlarged into disks with sterigmata at the margin. Parasitic

upon Cichoriaceae. Zoospores rare. Bremia

Tips of branches swollen, but not disk-like. Parasitic on Viola. Zoospores

rare or none produced. Bremiella

Tips of branches acute, conidia germinating by a germ tube.

Peronospora
TiDS of branches acute, conidia germinating by zoospores.

Pseudoperonospora

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PHYCOMYCETEAE: MUCORALES,

ENTOMOPHTHORALES, ZOOPAGALES,

ECCRINALES

THE first three orders that form the subjects of this chapter agree in
the more or less coenocytic structure of the mycehum, in the union
of equal or unequal gametangia to form "zygospores," and in the total
absence of flagellate sexual or asexual cells. Also, chitin is present in most
of these forms and the cell walls do not show the cellulose reaction upon
the application of a solution of chloriodide of zinc.
These three orders may be distinguished as follows:

Mucorales: mycelium very extensive, nonseptate or septate in older aerial
hyphae. Asexual reproduction typically by aplanospores formed in terminal
sporangia. In a few genera these are reduced to indehiscent sporangioles
which function as conidia. Sexual reproduction usually present.

Entomophthorales : mycelium not very extensive, at first coenocytic but sooner
or later becoming septate and often falling apart into hyphal bodies. Asexual
reproduction typically by conidia which are usually shot off with violence.
Sexual reproduction frequently replaced by the parthenogenetic development
of "azygospores."

Zoopagales: mycelia very slender, nonseptate or septate when older, attached
to their hosts (aquatic or soil-inhabiting amoebae, nematodes, or insect
larvae) by more or less complicated haustoria. Asexual reproduction by
various types of conidia. Sexual reproduction by the union of gametangia
to form zygospores of various shapes.

In addition to the foregoing are the Eccrinales which form a group
of fungi whose relationship is very doubtful. They are placed here as a
sort of appendix to the Phycomyceteae so that they may not escape
attention.

Order Mucorales. The Mucorales are widely distributed fungi with
a stout, well-developed, much branched coenocytic mycelium, very
similar to that of some representatives of the Saprolegniales. In the older
mycelium, especially in the aerial portions, septa may divide it into pluri-
nucleate segments, but the young mycelium and that submerged in the

150

ORDER MUCORALES 151

substratum usually remain nonseptate. Ayers (1935) has shown that
septation in Dispira cornuta van Tiegh, begins in the germ tube and that
numerous septa are formed in both young and old mycelium. The cell
wall is reported by von Wettstein (1921) to contain chitin and pectose
compounds and no cellulose, but Mangin (1899) reports true cellulose
in young sporangia of some species and Hopkins (1929) found both
cellulose and chitin in Mucor rouxianus (Calmette) Wehmer. It must be
noted that Nabel (1929) could find no cellulose in this species nor in any
other members of the order.

Asexual reproduction is typically by the formation of nonmotile, en-
cysted spores (aplanospores) in sporangia terminal to the hyphae. These
sporangia are formed in the same manner as in the Saprolegniales and
Peronosporales, by the passage of a portion of the contents of the hypha
into a terminal enlargement which is then cut off from the hypha by a
septum. The multinuclear contents of the sporangium are divided by
cleavage planes into naked, at first polyhedral, cells containing one or
more nuclei each. These then round up and encyst and escape by the
rupture or dissolution of the sporangium wall. They germinate by a stout
germ tube. Swingle (1903) has shown that all of the protoplasm in the
sporangium is used up in the formation of the spores. There are very
interesting evolutionary modifications of this typical sporangium, leading
to the development of structures comparable to conidia.

I The greater part of the mycelium may be represented by that within
the substratum, the aerial portion consisting of scarcely more than enough
to give rise to the sporangia. On the other hand in many species the aerial
mycelium may be very extensive, forming a large cottony mass from
which arise, here and there, the sporangiophores. It is usually white but
often the sporangiophores are dark colored. As it grows older this aerial
mycelium may become septate but not truly cellular for the segments
formed are plurinucleate coenocytes. A number of species of Mucor when
growing in a medium rich in nutrients and of rather high osmotic pres-
sure (e.g., a rather concentrated sugary medium) form a yeast-like growth
instead of the normal filamentous mycelium.

The Mucorales are mainly saprophytic on vegetable matter, more
rarely on animal matter, and are abundant in the soil and in plant debris.
Many are coprophilous. Some are weak parasites on living plant tissues
which are rich in stored food but not active, such as the roots of the
sweet potato {Ipomoea batatas (L.) Lam.). A number of species are para-
sitic upon other fungi, even upon other Mucorales. A few have been
described as parasites on animal tissues.

The probable course of sporangial evolution within the Mucorales
can best be followed by studying the sporangia of a selected list of genera.
The genera chosen may not represent direct lines of descent, since the

PHYCOMYCETEAE

Fig. 49. Mucorales, Family Mortierellaceae. Mortierella rostafinskii Bref. (A)
Sporangium. (B) Hyphal cluster at base of sporangiophore. (C-E) Stages in production
of zygospore. (A-B, after Brefeld und von Tavel: Untersuchungen aus dem Gesammt-
gebiete der Mykologie, Heft 9, pp. 1-156, Miinster i. W., Heinrich Schoningh. C-E,
after Brefeld: Botanische Untersuchungen iiber Schimmelpilze, Heft 4, pt. 5, pp. 81-
96, Leipzig, Arthur Felix.)

sexual reproduction is left unconsidered, but they probably indicate in
general the directions that the modifications followed.

The simplest and probably the most primitive type of sporangium is
that found in the genus Mortierella. It must not be understood that this
genus is considered to be the most primitive of the Mucorales, for in its
sexual reproduction it is so much modified from the more typical repre-
sentatives of the order that it is clearly to be recognized as well advanced
in evolution. However, in its sporangial development it seems to have
retained a very primitive structure. This is merely one of the very many
cases where evolution has advanced far along certain lines of development
(in this case the manner of sexual reproduction) while remaining about
at a standstill in its mode of asexual reproduction. The sporangium is a
spherical enlargement of the apex of the sporangiophore, set off from the
latter by a cross wall at the point where the enlargement begins. In
sporangial evolution the next step appears to have been the development,
as in Mucor, of a "columella." This in reality represents a displacement of
the septum separating the sporangium from the sporangiophore so that it
arches up into the former. The columella is laid down in the position it is to

ORDER MUCORALES 153

occupy and does not represent a cross wall that subsequently bulged up
into the sporangium. This gives a much larger surface to the septum and
permits a much freer transfer of food into the sporangium. Accompanying
this more efficient food supply we find the sporangia to be larger, almost in
proportion as the columella increases in size. In these two types of spo-
rangium the numerous spores escape by the dissolution or breaking up
of the relatively thin sporangial wall. The columella often remains firm
and unchanged after the sporangium has ruptured and the spores have
been set free. Schostakowitsch (1896) reported that in Meteor proliferus
Schost. the columella forms a new sporangium by proliferation, reminding
one of the condition in Saprolegnia. (Figs. 49 A, 50A-C.)

The sporangium of Pilobolus represents a special modification of the
foregoing type. In it the apical wall of the many-spored sporangium is
very much thickened. The columella is rather small. Below the spo-
rangium the sporangiophore is enlarged into a subsporangial vesicle that
may be two or three times the diameter of the somewhat flattened
sporangium. The sporangiophore tip is sensitive to light and this leads
to the curvature of its lower part so that the sporangium is directed
toward the source of the light. As the vesicle enlarges the turgor finally
becomes so great that the apex ruptures and the sporangium is blown off,
along w4th the watery contents of the vesicle, sometimes to a distance of
over a meter. The sporangia adhere to vegetation and are eaten by
herbivorous animals through whose digestive tracts the spores pass un-
harmed. In the dung of these animals the mold grows and produces its
conspicuous fructifications. (Figs. 51A, 52.)

Returning to the genus Mucor we find that some species have un-
branched sporangiophores while in others the sporangiophores may
branch sympodially or monopodially, each branch terminating in a spo-
rangium. Usually these sporangia are approximately equal in size al-
though frequently the terminal one is slightly larger. In M. prolifer-us
and some other species, the spores produced in the lateral sporangia are
smaller than those produced in the larger, terminal sporangium. In
Thamnidium the terminal sporangium, which is often the first one formed,
is larger and possesses a well-developed columella; somewhat below it
there grow out from the sporangiophore short branches (often much
forked dichotomously), all terminating in small sporangia or some in
pointed spines. These small sporangia (sporangioles) are few-spored,
sometimes with not over two or three spores, and possess no columella.
The whole sporangiole becomes detached and distributed by air currents.
As water is absorbed the spores swell and burst the sporangial wall and
escape. In some species of Thamnidium, under certain conditions only
the sporangioles are formed. In the genus Dicranophora the sporangioles
are only one- to two-spored and their spores are much larger than those

154

PHYCOMYCETEAE

Fig. 50. Mucorales, Family Mucoraceae. Mucor mucedo (L.) Fres. (A) Mature
sporangium just before escape of spores. (B) Escape of spores upon dissolution of
sporangium wall. (C) Columella after escape of spores. (D) Sexual reproduction in
early stage, with thin-walled gametangia in contact. (E) Mature zygospore. (F)
Germinating zygospore forming sporangiophore. (After Brefeld: Botanische Unter-
suchungen uber Schimmelpilze, Heft 1, pp. 1-64, Leipzig, Arthur Felix.)

ORDER MUCORALES

155

in the terminal sporangium, which is often lacking. In Chaetodadium
the large terminal sporangium is entirely lacking. The sporangioles are
one-spored and may be indehiscent in some species or in other species
permit the spore to escape on germination. These monosporous spo-
rangioles are often called conidia. (Fig. 53C, K.)

In Blakeslea the sporangium has a large columella when the fungus
is well nourished and is smaller and few-spored without the columella

Fig. 51. Mucorales,
Family Pilobolaceae. (A,
B) Piloholus kleinii van
Tiegh. (A) Diagram of
optical longitudinal sec-
tion of sporangiophore
and sporangium. (B)
Zygospore. (C) Piloholus
crystalUnus Tode ex van
Tiegh. Heterothallic for-
mation of zygospore. (A,
after Buller : Researches
on Fungi, vol. 6, pt. 1,
pp. 1-224, London, Long-
mans, Green and Co. B,
afterjZopf •iNova^Acla Leo-
poldina, 52(7) :352-358. C,
after Krafczyk: Ber. deul.
botan. Ges., • 49(3) :141-
146.)

when poorly nourished. In addition numerous small two- to four-spored
sporangioles are formed on short sterigmata from the surface of large
rounded heads clustered at the apex of a sporangiophore. These spores
differ markedly in size and shape from those produced in the sporangia.
In Choanephora the sporangium possesses a columella and the sporangioles
are borne as in Blakeslea on round heads at the apex of the sporangiophore
of the other type. They are monosporous and indehiscent and function, as
in Chaetodadium, as conidia. In Cunninghamella the sporangia are never
found and only the heads of indehiscent sporangioles ("conidia") are
developed. In Mycotypha the stalk of the sporangiophore is septate and
the head is a little enlarged and much elongated. It is covered with in-

156 PHYCOMYCETEAE

\i Source of light

Fig. 52. Mucorales, Family Pilobolaceae. Pilobolus sp., showing growth of sporangio-
phores toward the light. (Courtesy, K. B. Raper.)

numerable ellipsoidal indehiscent one-spored sporangioles, mostl}^ in
closely crowded whorls. (Fig. 54.)

Another direction of sporangial modification is found in the Pipto-
cephalidaceae. In this family the sporangiophores or their branchlets are
somewhat swollen terminally and from these swollen portions there grow
out radially numerous cylindrical sporangia. In these the spores are
formed in a single row, the number ranging from 2 to 6 or 8, or more —
rarely, up to 30. As the spores approach maturity they may enlarge so
that the sporangium is constricted between them and breaks apart into
one-spored pieces or the whole sporangium becomes detached and the
spores escape one by one from the open base. In the former case the
earlier students of these forms interpreted the structure as a chain of
conidia. In the genera Coemansia, Kickxella, and Martensella, which are
included by Linder (1943) in a distinct family, the Kickxellaceae, these
sporangia are reduced to indehiscent, one-celled structures which are
borne on distinct sterigma-like structures arranged pectinately on one
side of lateral, several-celled sporocladia. (Figs. 55, 56.)

Sexual reproduction is typically by the union of two approximately
equal gametangia to form a so-called zygospore which usually occupies
the cavities of the two gametangia and develops thick walls. The game-
tangia are multinucleate. The zygospores germinate, usually after con-
siderable time, by the formation of a germ tube which may branch and
start new mycelium or remain unbranched and terminate in a sporangium.
The study of the sexual reproduction of the Mucorales reveals an
interesting state of affairs. In perhaps the majority of species tested in
this respect a culture started from a single spore as well as all cultures
derived from sporangia produced on this culture will, when grown sepa-
rately or in contact with each other produce no zygospores. On bringing
such cultures into contact with other cultures of the same species, origi-

ORDER MUCORALES

157

Fig. 53. Miicorales. (A-B) Family Mucoraceae. Zygorhynchus macrosporus Ling-
Young. (A) Sporangiophores and sporangia. (B) Anisogamous formation of zygo-
spores. (C-K) Family Thamnidiaceae. (C-J) Dicranophorafulva Schroet. (C) Primary
sporangium and clusters of sporangioles. (D-J) Successive stages in sexual reproduc-
tion. (K) Thamnidium elegans Link, sporangiophore showing large terminal sporan-
gium and lateral clusters of sporangioles. (A-B, after Ling- Young: Rev. gin. botan.,
42(495) :152. C-J, after Dobbs, Brit. Mycol. Soc. Trans., 21(1-2) :172, 183. K, after
Brefeld from Comparative Morphology of Fungi, by Gaumann and Dodge. McGraw-
Hill Book Company.)

nated from plants obtained in various places, it is found that sometimes
at the line of contact between two cultures very abundant zygospore
formation occurs. On their part these other plants are self-sterile or
sterile when grown in contact with each other so far as sexual reproduc-
tion is concerned. Thus A. F. Blakeslee (1904) determined that for many
species of Mucorales there are two sexes, each capable of almost indefinite
perpetuation by means of the asexual spores, but producing zygospores
only when the mycelium of one sex comes into contact with that of the
other sex. He named this phenomenon heterothallism and called such
molds heterothalHc. In contrast to these he found many species in which,
when the proper conditions of environment and nutrition were met, zygo-
spore production would occur within the mycelium, originating from a
single spore. Such molds he called homothallic. The common bread mold,
usually called Rhizopus nigricans Ehr., is a good example of a hetero-
thallic mold while Sporodinia grandis Link, a mold frequently found on
decaying mushrooms, is homothallic. Ling- Young (1930) showed that

158

PHYCOMYCETEAE

Fig 54 Mucorales, Family Choanephoraceae. (A-E) Choanephora conjunda
Couch (A) Sporangium and sporangiophore. (B) Opening sporangium showmg spores
and columella. (C) Young stage of sexual reproduction. (D) Sexual reproduction with
mature zygospore. (E) Sporangiophore with sporangioliferous heads. (F-J) Blakeslea
trispora Thaxter. (F) Single sporangioliferous head. (G) Single sporangiole. (H) Spore
from sporangiole. (I) Young zygospore. (J) Mature zygospore. (A-E, courtesy,
Couch: J. EHsha Mitchell Sci. Soc, 41(1-2) :141-150. F-J, courtesy, Weber and Wolf:
Mycologia, 19(6) :302-307.)

ORDER MUCORALES

159

Fig. 55. Mucorales, Family Piptocephalidaceae. (A-C) Syncephalis cornu van
Tiegh. (A) Sporangiophore. (B) Sporangia containing spores. (C) Germination of
zygospore. (D, E). Piptocephalis cruciata van Tiegh. (D) Branched sporangiophore.
(E) Terminal branches with sporangia borne on cruciate sterigmata. (F-I) Pipto-
cephalis freseniana de Bary, stages in zygospore formation. (J-L) Dispira americana
Thaxt. (J) General habit of fungus. (K) Head with not quite mature sporangia. (L)
Two mature sporangia on sterigma. (A-E, after van Ticghem: Ann. sci. nat. Botan.,
6me ser., 1:5-175. F-I, after Brefeld: Botanische Untersuchungen iiber Schimmel-
pilze, Heft 1, 1-64, Leipzig, Arthur Felix. J-L, after Thaxter: Botan. Gaz., 20(12) :513-
518, Univ. of Chicago Press.)

160 PHYCOMYCETEAE

when two colonies of the same sex of Phycomyces nitens (Ag.) Kze. are
grown in a culture medium rather poor in nutritive substances, as the
hyphae reach a distance apart of only 1 to 1.5 mm. their further growth
toward one another in the medium ceases while if the two colonies are
of opposite sex the hyphae intermingle and soon begin to form zygospores.
This resembles the "barrage sexuel" discovered by Vandendries and
Brodie (1933) in some Basidiomyceteae between mycelia of incompatible
sexual phases (Chapter 12). Since the majority of Mucorales do not exhibit
noticeable differences in size in the uniting gametangia it is impossible,
for most species, to decide which plant should be called male and which
female. Satina and Blakeslee (1926) have made chemical tests on the two
sexual strains of several species and found different reactions which seem
to indicate to which sex each strain belongs. Blakeslee (1920) and others
have observed that the sexual differences show different degrees of in-
tensity. A plant that is very strongly male will conjugate with plants of
all degrees of femaleness and vice versa. On the contrary a plant weakly
male will not conjugate with one weakly female. Blakeslee observed at-
tempted conjugation between hyphae of different species or even genera,
but only when opposite sexes were concerned. In this way it has been
possible to correlate the sexes of the various genera and spe'cies of Mu-
corales. Burgeff (1925) actually obtained hybrids between two species
of Phycomyces.

Lendner (1908) believed that of all the nuclei in the united game-
tangia only one pair survived to unite and become the zygote nucleus in
the homothallic species Sporodinia grandis Link. Miss Keene (1919)
studying Phycomyces nitens (Ag.) Kze., a heterothallic species, claimed
that the nuclei became reduced to six or eight pairs. Ling- Young (1930)
confirming the observations of Dangeard (1906), Moreau (1913), and
others believes that the majority of the nuclei undergo union instead of
one or a few pairs. The density of the protoplast as well as the large
number of granules of stored food and the thickness and hardness of the
zygospore wall make cytological studies of the behavior of the nuclei
very difficult. Cutter (1942a, b) reports the results of very extensive
studies which in some particulars contradict the conclusions of Lendner
and of Miss Keene and to a large degree substantiate those of Ling-
Young. He finds several types of behavior. In Mucor hiemalis Wehm.,
Blakeslea trispora Thaxt., Ahsidia spinosa Lendn., all three heterothallic,
and a number of other Mucorales, all of the nuclei entering the zygote
unite by twos within a few days and quickly undergo meiosis, the dormant
zygospore containing only haploid nuclei. In Rhizopus nigricans Ehr.,
Ahsidia glauca Hagem, and A. coerulea Bain. {Tieghemella glauca (Hagem)
Naum. and T. coerulea (Bain.) Naum., respectively), also heterothalHc,
some of the nuclei entering the zygote degenerate and some unite

ORDER MUCORALES

161

by twos. The dormant zygospore contains only diploid nuclei which
undergo meiosis shortly before their germination. In Phycomyces blakes-
leanus Burg, and P. microsporus van Tiegh., of which at least the former
is heterothallic, the nuclei increase in number in the zygote and become
gathered in groups of several nuclei each, with a few scattered single
nuclei. Shortly before germination of the zygospore some of the nuclei
unite by twos and in the sporangium arising from it are found some
diploid nuclei, some haploid nuclei, the product of meiosis, and probably

Fig. 56. Mucorales, Fam-
ily Kickxellaceae. (A, B)
Coemansia erecta Bainier.
(A) Vigorously growing coni-
diophores. (B) Sporocla-
dium. (C) Coemansia aci-
culifera Linder. Mature
sporocladium and a single
detached conidium. (Cour-
tesy, Linder: Farlowia,
l(l):49-77.)

also some haploid nuclei which represent the scattered single nuclei
which never united. In the homothallic Sporodima grandis, contrary to
Lendner's report, there is nuclear division in the zygote but no nuclear
fusion and no meiosis.

Germination generally does not occur until after a considerable time
has elapsed. Usually a stout upright sporangiophore is sent out after the
outer wall is cracked open, and numerous spores are formed in a large
sporangium. In the plant usually called Mucor mucedo (L.) Fres., a
heterothaUic species, the spores produced in this sporangium are all of
one sex, showing that the differentiation of the sexes must have occurred
in some nuclear division within the zygospore prior to its germination.
On the other hand in Phycomyces nitens spores of both sexes are found in
the sporangium produced by the germinating zygospore, as well as,

162 PHYCOMYCETEAE

occasionally, a few spores that are not yet differentiated sexually and
which produce homothallic plants. Even in these plants the spores in the
sporangia that they bear become more and more differentiated into the
two sexes. Evidently this is connected with the nuclear condition found
by Cutter to exist in the zygospores of this species. Sjowall (1945) claims
that in M. mucedo and M. hiemalis the gamete nuclei unite and undergo
meiosis early in the zygospores so that for most of the resting period the
nuclei are haploid. These nuclei undergo degeneration until finally only
a few, apparently all of the same sexual phase, enter the sporangium. In
Rhizopus the union of nuclei occurs as a zygospore is germinating and
the reduction division as the sporangium is being formed, evidently only
one diploid nucleus being concerned. According to him in the process
of meiosis three of the four nuclei formed degenerate so that only one
haploid nucleus is available for further functioning. In the sporangium
the spores start out with one nucleus each, usually, but in many cases
this divides so that the mature spores have several nuclei.

The sexual process of the Mucor type is as follows : At a point where
the hyphae come in contact (if of a heterothalhc species these two hyphae
must originate from plants of opposite sex) a swelhng occurs in each
hypha, pushing them apart. These two swellings flatten against one
another and become much enlarged, tapering down to the hyphae from
which they arose. Soon a cross wall appears in each of the two processes,
parallel to and a little distance from the flattened surface of contact,
forming the two gametangia. No significant difference in the size of the
two gametangia can be noticed except in a few species. Each contains a
rather dense mass of cytoplasm with many nuclei. Beginning at the center
the double wall separating the two gametangia dissolves away. In some
cases there is a bulging of the central part of this double wall into one
of the gametangia and when the wall is dissolved the cytoplasm and
nuclei pass into this receptive gametangium. Soon, however, the whole
intervening wall is dissolved away and the mingled cytoplasm fills out
the cavities of the two gametangia. In the very young zygospore the thin
wall represents that of the original gametangia. Within this is laid down
a layer of more or less distinct dark plates or pyramids below which a
two- or three-layered cell wall is secreted. As the zygospore enlarges the
original gametangial wall is broken into small pieces that flake off so
that finally the exterior is covered by the dark plates which may re-
semble short broad spines. Ling- Young showed that in many cases of
heterothalhc species the more vigorous strain (female) usually produces
a larger gametangium and a larger suspensor than does the male my-
celium. In Mucor hiemalis Wehm. not only is this the case but according
to this author the contents of the male gametangium pass into the female
gametangium through a small opening which then becomes closed so

ORDER MUCORALES 163

that the zygospore is formed solely from the female gametangium. (Fig.
50D-F.)

In a few species of Mucorales the gametangia and supporting sus-
pensors are very unequal so that the product of the union of the two
gametangia might be called an oospore. In general practice, however,
they are called zygospores since their mode of origin is similar to that in
isogamous species. In Dicranophora the one gametangium is many times
larger than the other. In Zygorhynchus also, the difference between the
two gametangia is marked. Both these genera are homothallic. (Fig.
53A-J.)

In Piloholus, Mortierella, Piptocephalis, Phycomyces, and other forms
two hyphae coming into contact may become attached by finger-like
processes or wind around each other once or twice and then grow parallel
and in contact. Very soon they curve away and bend so that only the
tips are in contact, much as in a pair of tongs. In Piptocephalis and Endo-
gone the zygospore does not form in the space occupied by the two
gametangia but buds out from these to form an external zygospore. (Figs.
51B-C, 54C-D, 55F-I, 59A-F.)

The zygospore remains naked in the majority of species. In Phyco-
myces nitens branches grow out from the suspensors, surrounding the
zygospore loosely with stiff, black, more or less dichotomously branched
processes. In Ahsidia glauca these protective structures are curved and
hooked. Often they arise more abundantly from the female suspensor in
both these species. In Mortierella a dense hyphal mass several layers in
thickness is formed around and closely appressed to the zygospore. (Figs.
49C-E, 57A-B, H-J.)

The order Mucorales is divided into several families. Mycologists are
not in accord as to their number or limits. Probably these differences of
opinion will remain until greater agreement is attained as to the probable
course of phylogenetic development within the order and as to the rela-
tive importance to be ascribed to the evolutionary processes involving
the sporangia and those concerning sexual reproduction. If the group is
primitively isogametangic then the heterogametangic genera represent a
later modification while if heterogamy is considered primitive then the
isogametic condition found in Mucor is a modification of this ancestral
condition. The customarily used classification is based largely upon the
asexual reproductive structures and development and is probably artificial
in many particulars. It can be replaced by a more natural system only
when the above-mentioned questions are settled satisfactorily. In this
book eight families are recognized, following Fitzpatrick (1930) in the
main. These agree in most points with the six families into which Zycha
(1935) divides this order. Naumov (1935, 1939) recognizes eight families.
He divides many of the older genera so that he includes 38 genera and

164

PHYCOMTCETEAE

Fig. 57. Mucoralcs, Family Mucoraceae. (A) Absidia septata van Tiegh. &
Le Monnier, sporangiophores and zygospore. (B) Absidia glauca Hagem, mature
zygospore. (C-D) Circinella minor Lendner. (C) Unopened sporangia. (D) Opened
sporangia showing columella. (Ej Circinella umbellata van Tiegh. & Le Monnier,
zygospore. (F-G) Sporodinia grandis Link. (F) Dichotomously branched sporangio-
phore. (G) Zygospore. (H-J) Phycomyces microsporus van Tiegh., stages in formation
of zygospore. (A-F, after Lendner: Beitr. Kryplogamenflora Schweiz, 3(1):103, 104,
110, 134, 137. G, after Ling- Young: Rev. gin. botan., 42(496) :206. H-J, after Christen-
berry, J. Elisha Mitchell Sci. Soc, 56(2):332-366.)

ORDER MUCORALES 165

340 species and subspecies, excluding the Endogonaceae and a few other
forms. Including these the number would reach 44 genera and about
370 species.

Family Mucoraceae. Sporangia relatively large and many spored,
oval or spherical, with a well developed columella. The sporangia are all
alike and are borne singly, or several may be produced on a racemosely
or otherwise branched sporangiophore. The wall of the mature spo-
rangium may dissolve in the presence of moisture or may break into
pieces and fall away, thus releasing the contained spores. The most impor-
tant genera are : Mucor, whose unbranched or branched sporangiophores
arise from the main mycelium; Sporodinia, whose sporangiophores are
repeatedly and closely dichotomously branched ; Rhizopus, in which long
stout creeping hyphae (stolons) form tufts of sporangiophores at the
points where the stolons attach themselves by rhizoids. Ahsidia, Zygor-
hynchus, and Circinella belong in this family as do several other genera.
The zygospores are naked or more or less protected and are produced by
both the Mucor type and the Piloholus type as described above. It should
be noted that some of the names used above, e.g., Mucor, Rhizopus, and
Sporodinia, do not conform to the international rules of nomenclature
requiring the earliest generic names to be used. Until some future Inter-
national Botanical Congress shall settle the status of Syzigites vs.
Sporodinia, Rhizopus vs. Mucor, Mucor vs. Hydrophora, it will perhaps be
well to use the more customary generic names. (Figs. 50, 53A, 57.)

Family Pilobolaceae. Sporangia more or less flattened vertically
with a thick dark colored apical wall which does not break or dissolve.
The sporangiophore is swollen at the base in Piloholus and has a pro-
nounced enlargement or subsporangial vesicle (in Piloholus) or a slight
one (in Pilaira) subtending the sporangium. The columella is more or less
conical or may project almost to the top of the sporangium. In Piloholus
when the maximum osmotic pressure has been attained in the subspo-
rangial vesicle it ruptures in a circular slit so that the columella and
adhering sporangium are violently expelled by the mass of liquid forced
out by the contraction of the ruptured vesicle. This sporangium turns
over in flight or on striking some object, such as a blade of grass, and
adheres by its gelatinous base. In Pilaira the elongating sporangiophore
places the sporangium in contact with some near-by object to which it
remains attached when the sporangiophore collapses. The dung-inhabiting
species of Piloholus are very easily obtained by bringing freshly dropped
horse manure into the laboratory and enclosing it in a large dish. After
a very few days the spores, which have passed unharmed through the
alimentary canal of the horse, will produce an abundant growth of my-
celium and the large conspicuous sporangiophores will appear in large
numbers. When the dish is placed in a box closed on all sides except for a

1G6 PHYCOMYCETEAE

small window, the sporangiophores can be induced to turn and discharge
their sporangia in the direction of the window, without much scattering
of aim, if the illumination is good. Miss R. F. Allen and Miss H. D. M.
Jolivette (1913) and later Miss Jolivette (1914) studied the relation of
the discharge to different colors of light qualitatively. Miss Parr (1918)
studied these relations quantitatively. The light at the violet end of the
spectrum is the most efficient, grading to the red without intermediate
maxima or minima. The presentation time, i.e., the minimum length of
exposure necessary to bring about the effect, "varies in inverse ratio to
the square root of the wave frequency" (Parr). Flint (1942) demonstrated
that an unidentified species of Piloholus gave no curvature but abundant
discharge of sporangia in random directions in red light of the range of
6576-7024 A units, and strong phototropic curvature and discharge with

blue light centering at 4600 A units.

As the sporangiophore is elongating, before the sporangium has begun
to form, it is pointed and positively phototropic, so that it points toward
the direction of the strongest light of effective wave length. With the
formation of the subsporangial vesicle the latter has been shown by
Buller (1921, 1934) to form a very effective light-perceiving organ or
"eye" which brings about the final aiming so that the sporangium is shot
off in the most favorable direction. Buller showed that in the vertical
direction the sporangium may be shot over six feet and horizontally,
eight feet.

In this family the suspensors of the gametangia are curved like a pair
of tongs.

Grove (1934) recognizes 9 species oi Piloholus with 7 additional doubt-
ful species, and 5 species of Pilaira, while Naumov (1939) recognizes
respectively 18 and 5. (Figs. 51, 52.)

Family Thamnidiaceae. The genera of this family show the begin-
ning of the differentiation into two types of sporangium, viz., the large
terminal sporangium with a columella and the small lateral separable
sporangioles which lack a columella. The latter are borne on short, var-
iously divided branches arising laterally on the main axis of the spo-
rangiophore at whose apex the large sporangium is produced, if present.
In Thamnidium the spores are alike in sporangium and sporangiole. In
some species some of the branches upon which the sporangioles are borne
are sterile at the tips and form spine-like processes. The sporangiole-
producing branches are dichotomously forked. The zygospores are formed
on approximately equal suspensors about as in Mucor. In Helicostylum
the sporangioles are borne on short circinate branchlets from the un-
forked lateral branches. In Dicranophora the sporangioles, according to
Dobbs (1938), are one- to few-spored and have a rounded columella and
thin sporangial wall. The spores are very variable in size and mostly

ORDER MUCORALES 167

bean-shaped, while they are mostly ellipsoid and more numerous and on
the whole smaller in the terminal sporangium. The gametangia are very
unequal in size. The genus Chaetodadium is parasitic upon other Muco-
rales. It shows close relationship to Thamnidium both in its sexual repro-
duction, which resembles that of Mucor, and in the production of numer-
ous sporangioles on forking branches some of which terminate in sterile
spines. The two genera differ in the complete absence of typical sporangia
in Chaetodadium as well as in the fact that the sporangioles are mono-
sporous and indehiscent, and hence are called conidia by many authors.
Other genera are recognized by some students of the group. (Fig. 53C-K.)
Family Choanephoraceae. The large sporangia are produced in
two genera and possess a columella and resemble those of Thamnidium.
The sporangioles instead of arising singly at the tips of forked branches
are found crowded on the surface of the swollen apical portion of a large
sporangiophore or of its branches. They are monosporous and indehiscent
in three genera and several spored and dehiscent in one genus, Blakeslea.
Most of the species are parasitic or saprophytic on flowers or other vege-
table matter. The proportion of sporangia to sporangiole-bearing heads
in Blakeslea has been shown by Goldring (1936) to depend considerably
upon the amount of food present in the medium. Choanephora produces
sporangia and indehiscent monosporous sporangioles, while Cunning-
hamella produces only the latter. Gaumann interprets the swollen apex
of the sporangiophore of the latter as a homologue of the sporangium and
the sporangioles clustered on its outer surface as in reality spores which
have been, as it were, pushed out so as to become external instead of
being produced internally. This view is not at all in accord with that
of the author, who considers the sporangioles to be homologous to those
of Thamnidium, i.e., reduced lateral sporangia. Possibly related to Cun-
ninghamella is the genus Mycotypha, described by Miss Fenner (1932).
The mycelium is like that of most Mucorales, coenocytic and much
branched and only occasionally septate. The sporangioles are reduced to
minute "conidia" closely covering the sides and apex of a cylindrical or
clavate enlargement of the upper portion of the eventually septate
sporangiophore. The resemblance of the head of sporangioles to the flower-
ing head of Typha suggested the name given to the organism. So far the
formation of zygospores has not been observed in Mycotypha. In
Choanephora and Blakeslea zygospore formation is much like that in
Piloholus but in Cunninghamella it resembles that of Mucor. The genus
Sigmoideomyces probably belongs to this family. Its heads of spores
(sporangioles) are borne laterally on branched septate sporangiophores
whose branches are curved more or less like the letter S. Sexual
reproduction is unknown. Possibly Thamnocephalis also is related. (Fig.
54.)

168 PHYCOMYCETEAE

Note : An additional genus of the Choanephoraceae was published too
late to appear in the text of this chapter. Cokeromyces Shanor (type
species C. recurvatus Poitras) produces numerous sporangioles on elon-
gated recurved stalks arising from the terminal capitate swelling of the
sporangiophore. Twelve to twenty spores are produced in each sporangi-
ole, lacking the striae and appendages found in other genera of the family.
No true sporangia are produced. Zygospores arise in the manner of Mucor
and Cunninghamella.

Shanor, Leland, Adrian W. Poitras, and R. K. Benjamin: A new
genus of the Choanephoraceae, Mycologia, 42(2) :27 1-278. Figs. 1-12.
1950.

Family Piptocephalidaceae. The species of this family are largely
parasitic on other Mucorales although some are saprophytic. The spo-
rangia are narrow and more or less clavate or cylindrical, with the spores
usually in one row, often appearing when mature like chains of conidia.
The number of spores formed in the sporangium varies from two (rarely
one) to 30. Usually the sporangium breaks into monosporous segments,
the spore being enclosed in the sporangial wall. In a few cases the whole
sporangium may break off and the spores escape individually from the
opening. In Syncephalis the sporangiophore is upright and unbranched,
tapering from the more thickened basal portion up to the enlarged,
usually spherical head which may be upright or nodding. The radiating
sporangia may arise directly from the surface of this head or singly or
several together upon short " sterigmata. " These sporangia may be de-
hiscent at maturity letting the contained two to several spores escape.
In Syncephalastrum the sporangiophore is mostly branched as in the
racemose type of Mucor. On the spherical heads terminating the branches
the narrow sporangia arise radially without the intervention of sterig-
mata. These break apart into monosporous segments so that they were
long considered to be true conidia and compared to the chains of conidia
formed on the swollen heads of the conidiophores of Aspergillus. In Pipto-
cephalis the sporangiophores are dichotomously branched, bearing at the
tips of the branches slightly enlarged segments ("sterigmata") of various
shapes from which arise several sporangia which break up into one-spored
pieces. In Dispira the terminal heads of the much branched sporangio-
phore bear two-celled "sterigmata" from each of which arise short spo-
rangia of from two to several spores. The genus Spinalia (Vuillemin,
1904) probably belongs in this family. It has a coenocytic mycelium and
produces heads covered by "conidia." On the spherical heads the so-
called "conidia" occur in chains of two. According to Vuillemin (1922)
the zygospores are formed by the Mucor type of sexual reproduction in
Syncephalastrum. In Piptocephalis and Syncephalis sexual reproduction

ORDER MUCORALES

169

is more like the Piloholus type except that the zygospore buds out from
the united gametangia and matures externally. (Fig. 55.)

Family Kickxellaceae. Linder (1943) erected this family to contain
Kickxella, Coemansia, and Martensella. In these the sporangia (conidia)
are indehiscent and one-celled, but in the first-named genus an incom-
plete septum indicates that perhaps it is descended from a two-spored
ancestral form. In all three genera there are special lateral sporangium-

FiG. 58. Mucorale^
Family Mortierellaceae.
Dissophora decunibens
Thaxt. (A) Young apical
portion of fertile hypha.
(B) Portion of fertile
hyphae bearing sporangi-
ophores. (C) Mature spo-
rangium. (After Thaxter:
Botan. Gaz., 58(4): 353-
366, Univ. of Chicago
Press.)

bearing sporocladia consisting of several cells. From each cell of these
except one or more terminal cells arise several sterigmata each bearing a
single conidium. The sporocladium with its conidia somewhat resembles
a comb. The nearest relationship of this family appears to be with some
members of the preceding family. Sexual reproduction has been observed
in only one species of Coemansia and resembles considerably that of
Piloholus. (Fig. 56.)

Family Mortierellaceae. The position of this family is uncertain.
In sporangial structure the absence of a columella would indicate a posi-

170 PHYCOMYCETEAE

tion below that of the Mucoraceae, but the occurrence of sporangioles
which are reduced to conidia in some forms would suggest a higher posi-
tion. This is also indicated by the formation of a dense hyphal protective
coat around the base of the sporangiophore in some species and around
the zygospores which are formed much as in Piloholus. Of the several
genera assigned to this family Moriierella is the only one that has been
fairly well studied. In addition to the spherical, many-spored sporangia
there occur in some species indehiscent, monosporous sporangioles ("con-
idia" or "stylospores"). The thirty or more species are mostly sapro-
phytic on vegetable matter or are weak parasites. In Haplosporangium
the main sporangiophores are more or less prostrate, septate and con-
stricted at the septa. From these segments arise tapering lateral branches
bearing terminally or also laterally one-spored sporangioles (in one
species) or two-spored sporangioles in H. bisporale Thaxter. No large
sporangia are produced and sexual reproduction is unknown. Dissophora
resembles a racemosely branched Mortierella. (Figs. 49, 58.)

Family Endogonaceae. The twenty-six or more species at present
included in this family are grouped together on the basis of their sporo-
carps. These are loose or moderately firm sclerotium-like bodies a few
millimeters to 2-3 centimeters in diameter and produced in humus-rich
soil, leaf mold or among mosses. The interwoven hyphae are coenocytic
with occasional septa in the old hyphae. Scattered throughout the sporo-
carp are three types of reproductive bodies, but never all three in the
same sporocarp. In two species there are numerous sporangia. They have
a transverse basal septum as in Mortierella. They contain from 4-12
spores in one species and many spores in the other. In a dozen or more
species the reproductive bodies are thick-walled chlamydospores of var-
ious shapes and sizes. Some authors consider these to be azygospores.
True sporangia are entirely lacking. In a few species only zygospores are
formed in much the same manner as in Piptocephalis, with the suspensors
parallel and the zygospore budding out of the united gametangia or out of
the larger of the two. According to Bucholtz (1912) in Endogone lactiflua
Berkeley most of the numerous nuclei in each gametangium degenerate
leaving only one privileged nucleus and these unite in the zygospore.
Atkinson (1918) reports that in E. sphagnophila Atk. many pairs of
nuclei unite. In E. fasciculata Thaxter and E. occidentalis Kanouse both
chlamydospores and zygospores are present in the sporocarp. Miss
Walker (1923) obtained E. 7nalleola Harkness in pure culture and pro-
cured mycelial growth and sporangia of the Mortierella type but no sporo-
carps were produced nor chlamydospores nor zygospores. Miss Kanouse
(1936) succeeded in obtaining a culture from some of the mycelium in a
not quite mature sporocarp of E. sphagnophila. This produced a Mucor-
like mycelium bearing numerous sporangiophores and very closely re-

OEDEB MUCORALES

171

Fig. 59. Mucorales, Family Endogonaceae. (A-F) Endogone lactiflua Berkeley.
(A) Young gametangia. (B) Gametangia in longitudinal section, showing numerous
nuclei. (C) Gametangia in longitudinal section, showing one privileged nucleus in the
right-hand gametangium, the others receding basally. (D) Gametangia set off by
septa and male nucleus entered into the female gametangium. (E) Zygospore budding
out from top of female gametangium. (F) Practically mature zygospore. (G) Endogone
pisiformis Link, mature sporangium. (After Bucholtz: Botan. Centr. Beihefte, II Abt.,
29(2):147-225.)

sembling the mycelium and sporangiophores of M. ramannianus Moller,
and like that species with a pink to rose color. Very numerous chlamydo-
spores were produced also and eventually, when grown on 1 per cent malt
agar, zygospores of the typical Endogone type. The sporangial walls
quickly break up or, when placed in water, deliquesce, letting the spores
escape and revealing a spherical columella. The species is homothallic,
for cultures from a single spore produce zygospores. Endogone occidentalis
contains in its sporocarps both chlamydospores and zygospores, but
when brought into culture by Miss Kanouse gave rise to chlamydospores
only, producing neither zygospores nor sporangia. In none of the cultures
by Miss Walker or Miss Kanouse could typical sporocarps be obtained.
Because the sporangia obtained from cultures of E. sphagnophila are
Mwcor-like, not like those of E. malleola Harkness and E. reniformis
Bresadola, Miss Kanouse segregates the latter two in a separate genus
Modicella. The two other genera assigned to this family, Sclerocystis and

172 PHYCOMYCETEAE

Glaziella, produce only chlamydospores and differ from Endogone in their
arrangement in the sporocarp. To what family of Mucorales the Endo-
gonaceae are nearest is uncertain. The lack of columella in the sporangial
species has led some mycologists to suggest that the family is related to
the Mortierellaceae, especially since in E. lactiflua the enlarging zygo-
spore becomes surrounded, according to Bucholtz, by a tight weft of
hyphae as in Mortierella. This weft is lacking in E. sphagnophila in which
species the sporangium possesses a columella. Possibly the family is in
reality not monophyletic but represents an assemblage of more or less
unrelated species which agree only in the production of a sclerotium-like
sporocarp under certain environmental conditions. The earlier mycol-
ogists placed this family in various positions; among the Ascomyceteae,
in the Protomycetaceae, close to the Ustilaginaceae, etc., but Bucholtz
(1912) by his study of zygospore formation showed its affinity to be with
the Mucorales. (Fig. 59.)

The genus Helicocephalum Thaxter (1891), with two or three species,
has the type of growth of Mucorales but whether it is related to any of
the foregoing families is uncertain until further study has been made
(see Drechsler, 1934).

Order Entomophthorales. The Entomophthorales are fungi whose
mycelium is often much reduced. Upon germination of the spore the germ
tube is usually coenocytic but sooner or later septa appear which divide
it into plurinucleate or even uninucleate segments, forming a septate
mycelium in some genera or in other genera falling apart into the so-called
"hyphal bodies." These latter may multiply by fission. Of the five or
more genera two are saprophytic or rarely parasitic, one is parasitic in
the gametophytes of ferns, one in the Desmidiaceae, and the remainder
in the bodies of insects. The latter group includes the vast majority of
the known species of the order. Asexual reproduction is by means of
reduced sporangia (probably more properly sporangioles) which may be
uninucleate or plurinucleate and are shot off singly from the apex of a
somewhat club-shaped sporangiophore, except in the genus Massospora
in which they are produced internally in the body of the insect host. These
sporangia are usually called conidia, as in the homologous structures in
the Peronosporaceae and Mucorales. In many cases such a "conidium"
may produce another " conidiophore " and shoot off a secondary conidium,
and that may produce a tertiary conidium, and so on. The ultimate ger-
mination is by a germ tube except in the genus Basidiobolus in which
the conidium shows its true sporangial nature by producing internal
spores. Sexual reproduction is by the union of mycelial segments (or of
hyphal bodies) to form zygospores which may lie in one of the uniting
gametangia (Basidiobolus), between the suspensors (Conidiobolus), or later-
ally to the fusion cell or to one of the conjugating gametangia {Enlo-

ORDER ENTOMOPHTHORALES

173

CnZh' fx\ ^^to™«Phthorales, Family Entomophthoraceae. Conidzobolns hrefeldianns
Couch. (A) Comdiophore and almost mature conidium. (B-D) Successive stages in

£1726%) :Tlt?3of ""^ ^'^''' '^ ^^^"^^ -^^^^-^'^^^- (Courtesy, Couch, LI J.

174

PHYCOMYCETEAE

mophthora). The zygospore is thick-walled. In many species similar
spores are formed parthogenetically, the azygospores.

Family Entomophthoraceae. Usually only one family is recognized
in this order: the Entomophthoraceae.

Conidiobolus is saprophytic or weakly parasitic on the fruit bodies of
higher fungi and on certain insects. Its asexual spores are conidia, not
dividing internally into sporangiospores. The branching mycelium is
divided by septa into multinucleate segments. The spherical conidia are
shot off in the manner characteristic of the family. Sexual reproduction
occurs by the union of adjacent cells of unequal size in the same hypha
or by contact of two hyphae. The contents of the smaller gametangium
are cut off from the supporting cell (or suspensor) by a septum. They pass
through a pore into the female cell within which the thick-walled zygo-
spore becomes mature. On germination it produces a conidiophore or
mycelium. (Fig. 60.)

Basidiobolus is most frequently found growing saprophytically on
the excrement of frogs, lizards, etc. It has been isolated by van Overeem

'■'A

Fig. 6]. Entomophthoralcs, Family Entomophthoraceae. Asexual reproduction.
(A-F) Basidiobolus ranarum Eidam. (A) Developing conidiophore. (B) Apex of
mature conidiophore. (C-E) Development of conidium into sporangium in stomach
of frog. (F) Spores set free from sporangium. (G, H) Entomophthora muscae (Cohn)
P'rcscnius. (G) Grouj) of conidiophores. (H) Single conidiophore. (A, B, G, H, after
Thaxtcr: Mem. Boston Soc. Natural History, 4(6):133-201. C-F, after Levisohn : JaAr6.
wiss. Botan., 66(3) :5 13-555.)

ORDER ENTOMOPHTHORALES 175

(1925) in pure culture from a fungus mass deep in a fistular abscess in
the leg of a horse in Java. On the excrement it forms sporangiophores
which shoot off the not yet divided sporangia, much as occurs in Piloholus.
These sporangia are eaten by beetles which in turn are devoured by
frogs or lizards in whose stomachs the beetles are digested, setting free
the sporangia which only then divide internally to produce the spores.
These escape from the sporangium and multiply in the alimentary canal
by fission or by budding. They are set free in the excrement and then
germinate to form mycelium on which are produced other sporangia. This
mycelium is at first coenocytic but soon septa appear and divide it into
a multiseptate branched mycelium. Zygospore formation is as follows:
Two adjacent segments of a hypha send up parallel beaks in contact,
into each of which a nucleus migrates and divides. One nucleus of each
pair remains in the apex at its beak, being cut off by a septum. One of the
two original segments enlarges considerably, an opening is dissolved
through the separating septum, and the nucleus and part of the cytoplasm
of the smaller cell pass into the larger cell. There the nuclei fuse and the
thick-walled "zygospore" is formed. Couch (1939) reported that the cell
wall responds to the chloriodide of zinc test by the cellulose reaction.
Because of this and of the uninucleate condition of the cells he advocated
placing Basidiobolus in a separate family, Basidiobolaceae. (Figs. 61A-F,
62A-E.)

Ancylistes was formerly included in the Ancylistidaceae (now called
Lagenidiaceae) because of the similarity of its vegetative structure and
sexual reproduction to Lagenidium. No zoospore production is known.
With the discovery by Miss Berdan (1938) that "conidia" similar to
those in the Entomophthorales were produced and that these were shot
off in the same manner the genus had to be transferred to the latter order.
The three species are parasitic in Desmidiaceae of the genus Closterium.
Two adjacent cells in the same filament or cells in contact in parallel
filaments may conjugate. One cell is usually smaller than the other and
the thick-walled zygospore is formed in a protuberance from the larger
cell. Its germination is unknown.

Completoria is parasitic in the cells of the gametophytes of ferns. In-
fection spreads from cell to cell of the host by means of hyphae pene-
trating the cell walls. Azygospores are formed in the host cells. The
conidiophores emerge into the air and bear conidia which are violently
discharged. The mycelial masses in the host cells are much branched and
lobed and not conspicuously septate.

Entomophthora is the largest genus of the family, containing, according
to Fitzpatrick (1930), about 40 species, all parasitic within insects. Some
species of this genus are called Empusa by some Avriters, but since this
name was used earlier for a genus of Orchids the next later name, Ento-

176

PHYCOMYCETEAB

Fig. 62. Entomophthorales, Family Entomophthoraceae. Sexual reproduction.
(A-E) Basidiobolus ranarum Eidam. (F-H) Entomophthora sepulchralis Thaxt. (I-K)
Entomophthora fresenii Now. (A-C, after Eidam: Beitr. Biol. Pflanz., 4(2):181-251.
D-E, after Fairchild: Jahrh. wiss. Botan., 30:285-296. F-K, after Thaxter: Mem.
Boston Sac. Natural History, 4(6):133-201.)

mophthora, must be used. The internal organs of the host are dissolved,
presumably by enzymes secreted by the fungus. The mycelium may be
quite extensive and filamentous with only occasional septa or may be-
come septate at frequent intervals or break apart into numerous hyphal
bodies. The clavate conidiophores emerge through the thinner parts of
the body wall of the insect and shoot off their conidia with great violence.
The conidiophores may be simple and packed together in a palisade layer
or they may be branched at the base. In the latter case the conidia are
uninucleate, in the former they are plurinucleate. Both azygospores and
zygospores are known in this genus. It is not settled yet whether only
one pair or several pairs of nuclei are functional, nor is it known at what
stage of development of the zygospore the nuclei fuse. E. muscae (Cohn)
Fresenius is the freciuent cause of the death of house flies (Musca domes-
tica) in the autumn months. The affected flies cling to window panes and
other objects and die there, and the glass immediately surrounding them
becomes whitened by a halo of discharged conidia. True zygospores are
unknown but azygospores are produced abundantly in Europe but very

ORDEB ZOOPAGALES

177

rarely in America. E. grylli Fresenius is very abundant in some seasons
in the plains states of the United States, causing the death of* immense
numbers of grasshoppers which climb up on stalks of grasses and other
plants and there die. In moist weather their abdomens are first covered
by belts of conidiophores emerging from between the segments, but in
dry weather these are not conspicuous. Azygospores are produced in
great numbers in the body cavity of the host. (Figs. 61G-H, 62F-K.)^

Several other genera have been described, including Alassospora,
parasitic in the 17-year cicada (Tibicina septendecim) . In this genus the
conidia are produced within the host, not on extruded conidiophores, and
are distributed by the gradual sloughing off of the affected parts while
the insect is still capable of creeping around. Sexual reproduction is
unknown.

Speare (1912) and Sawyer (1929) devised means of growing species
of Entomophthora in culture and thus have been able to follow out the
life histories of some species more fully than formerly.

Order Zoopagales. The fungi tentatively brought together in this
order are parasitic on soil-inhabiting and aquatic animals: amoebae,
nematodes, and insect larvae. The hyphae are coenocytic and at first
nonseptate and slender. Later they develop occasional cross walls. Sexual
reproduction is by conjugation of short or long filaments and the pro-
duction of zygospores of various forms, spherical to boat-shaped. Asexual
reproduction is by the formation of conidia, either laterally and singly
or apically, in the latter case sometimes in chains. Whether these conidia

Fig. 63. Zoopagales. (A-D) Family Harpellaceae. Harpella melusinae Leger &
Duboscq. (A) Vegetative hypha not yet septate. (B) Septate hypha bearing curved
conidia. (C, D) Formation of zygospore. (E) Family Genistellaceae. Genistella ramosa
Leger & Gauthier. Whole fungus with spikes of conidia and formation of young and
mature boat-shaped zygospores. (A-D, after Leger and Duboscq: Compt. rend.,
188(14) :951-954. E, after Leger and Gauthier: Compt. rend., 194(26) :2262-2265.)

178 PHYCOMYCETEAE

are comparable to the sporangioles of some Mucorales or to the sporangia
of Piptocephalidaceae needs further study.

Two groups of fungi placed by their discoverers (Leger and Duboscq,
1929b and Leger and Gauthier, 1932) in the families Harpellaceae and
Genistellaceae, occur as parasites (possibly only as commensals) in the
alimentary canals of various aquatic insects. They consist of slender un-
branched flexuous coenocytic hyphae (Harpellaceae) or branched coeno-
cytic hyphae (Genistellaceae). They are attached to the host tissues by
disk-like or lobed expansions, even finger-like structures in some cases,
but do not penetrate through the epidermal layer. The hyphae eventually
become septate and bear slender or stout, straight or curved, usually
uninucleate conidia, mostly one from each cell of the hypha, often only
along one side or in a crown near the apex. Leger and Gauthier (1935)
showed that these conidia bear at their point of attachment one (Stachy-
lina and Tijphella), two {Genistella) , or four (Harpella) fine filaments
which are coiled in the supporting cell and pulled out of the latter as the
conidium becomes detached. These appendages may be 3 to 6 times the
length of the conidium. Their function is not known but possibly they
have something to do with the manner of infection of the host. Sexual
reproduction occurs through the union of adjacent cells in the same fila-
ment, the thick-walled zygospore being formed in one of these cells as
in Basidiobolus or (often in the same species) by the union of conjugation
tubes from filaments lying side by side, the zygospores being formed in
the center of the enlarged connecting tube or on a stalk growing from
this point. In the Harpellaceae the zygospore is spherical while in the
Genistellaceae it is biconical or boat-shaped and surmounts a lateral
stalk from the uniting tube and is transversely perched at the summit of
the stalk. Two parallel filaments of Glotzia centroptili Gauthier may,
according to Miss Gauthier (1936), show several conjugations in a scalari-
form manner. Leger and his colleagues consider these two families to
belong in or close to the Entomophthorales. (Fig. 63.)

Drechsler (1935, 1936, 1937, etc.) has described numerous species of
fungi parasitic on terricolous amoebae and nematodes, placing these in
the family Zoopagaceae. This family he places tentatively between the
Mucorales and the Entomophthorales. These fungi have various types
of haustoria or internal mycelium within the hosts and long slender ex-
ternal aerial hyphae from which conidia arise. These external hyphae are
nonseptate and multinucleate at first. The conidia arise laterally along
the course of these filaments or are terminal and then single or in chains.
They are more or less spindle-shaped. The haustorium of Endocochlus,
parasitic in Amoeba, is a stout two or more times helicoidally wound
structure formed at the end of the infection tube which penetrated the

ORDER ZOOPAGALES 179

host from a conidiiim that adhered to the external surface of its victim
by means of a sticky substance. More rarely infection results from inges-
tion of the conidia. From this coil, after the death of the host, one or
more slender aerial unbranched or slightly branched hyphae which are
at first nonseptate emerge to the exterior. The terminal portion becomes
septate and from a few of the cells thus formed arise laterally sessile,
spindle-shaped conidia. Other short slender hyphae arising from the
haustorial coil grow parallel for a short distance and unite at the apex.
Just beyond the point of union a spherical thin-walled enlargement (called
by Drechsler, 1935a, the zygosporangium) is formed, equally upon the
apices of the two uniting hyphae or on an extension from near the apex
of one of them. Within this zygosporangium is produced an angularly
lobed, thick-walled zygospore. The zygosporangial wall may remain in-
tact and does not break up into small pieces and disappear as does the
corresponding structure in Mucorales. (Fig. 64.)

In Cochlonema the thick spiral haustorial body much resembles that
of Endocochlus but the spindle-shaped conidia are in chains on the aerial
conidiophores. They are apparently ingested by the host and thus infect
it. Sexual reproduction is much as in that genus but in some cases the
zygosporangial wall is lobed or warty to correspond to the lobes or warts
of the zygospore. In Bdellospora the spindle-shaped conidia are also pro-
duced in chains. Instead of being ingested by the amoeba and forming a
coiled haustorium in its interior the conidium adheres to the outer surface
of the host cell and sends in a slender infection tube which divides di-
chotomously into several short lobes. The external conidium enlarges
until many times its original size and from it arise the aerial conidiophores
and also the slender branches (from separate individuals) that coil
around one another many times and then conjugate at the tips to form
a zygospore as in the foregoing genera. In Zoopage the mycelium is ex-
ternal, somewhat branched and nonseptate. When an amoeba comes in
contact with such a hypha it adheres to it and a short-lobed haustorium
penetrates the host cell. The external mycelium produces short aerial
chains of elongated spindle-formed conidia. Sexual reproduction is much
as in the preceding genera. Acaulopage and Shjlopage are similar but their
conidia are single, not in chains, practically sessile in the former and on
short conidiophores in the latter. The latter genus may attack Amoebae
and Nematodes. Cystopage produces no distinct conidia but intercalary
or lateral chlamydospores in the intramatrical mycelium as well as in
the extramatrical mycelium. It attacks Nematodes and Rhizopoda.

Drechsler (1935) points out the similarity of the mycelium and catenu-
late conidia of some species of Actinomyces to those of some of the more
delicate forms in the Zoopagaceae and suggests that there may be some

180

PHYCOMYCETEAE

Fig. 64. Zoopagales, Family Zoopagaceae. (A-D) Zoopage phanera Dreohs. (A)
Parasitized amoeba with two haustoria from each of two hyphae. (B) Creeping hypha
with upright chains of conidia. (C) Early stage of sexual reproduction. (D) Mature
zygospore. (E-I) Endocochlus asteroides Drechs. (E) Large amoeba with stages of
infection and young thalli of fungus. (F) Coiled thalli of fungus producing zygospores
and sending out a long conidiiferous hypha (only basal portion drawn). (G) Portion
of fertile region of conidiiferous hypha. (H) Young stage of sexual reproduction.
(I) Mature zygospore. (Courtesy, Drechsler: Mycologia, 27(l):6-40.)

relationship here. He believes that the latter as well as the Harpellaceae
and Genistellaceae are related and have affinities with the Syncephalideae
in the Mucorales as well as with the Entomophthorales.

Order Eccrinales. Like the family Harpellaceae the members of this
order also grow as parasites attached to the wall of the alimentary canal,
stomach, intestine, or anal plates, of Arthropoda: insects, crustaceans,
myriapods, etc., both aquatic and land-inhabiting species. The genera

ORDER ECCRINALES

181

Eccrina and Enterobryus were described by Leidy (1849) but were con-
sidered by him to be endobiotic algae. Leger and Duboscq (1905) first
assembled these and other genera into a distinct order of fungi. Since
that date these authors (1916, 1929a, and other papers), Poisson (1929,
1931), Lichtenstein (1917), and others have recognized twelve or more
genera which Leger and Duboscq divide among three families. In general
these fungi consist of unbranched or not extensively branched hyphae
attached to the chitinous wall of the alimentary canal by a cup-like or
disk-like holdfast of callose. The hyphal walls of the fungi are reported
to be composed of cellulose. These hyphae are coenocytic at first and
may attain the length of 100 ^ in some species of Amoebidium up to over
10 mm. in Arundinula capitata Leg. and Dub. Reproduction occurs by
the formation of microspores, macrospores, and resting spores. The
microspores arise as follows: In the distal portion of the hypha (this
may be only a small portion or nearly the whole hypha) the nuclei arrange
themselves axially in close proximity. Cross walls are then formed pro-
ducing uninucleate microspores that are thin disks or short cylinders
rarely attaining an axial length equal to the diameter of the filament.
These usually have their walls free eventually from the hyphal wall and

Fig. 65. Eccrinales, Family Eccrinaceae. Enterobryus elegans Leidy. (A) Young,
not yet septate specimens. (B) Mature specimen, somewhat artificially coiled to save
space in drawing. (After Leidy: Smithsonian Inst. Pubs. Contribs. to Knowledge,
5(2):l-67.)

182 PHYCOMYCETEAE

by the breaking of the filament at the apex or through lateral openings
the spores escape. The macrospores are larger and may be uninucleate
at first, the septation of the hyphae being often oblique. The nuclei divide
until usually there are four nuclei to each macrospore. Their walls also
become separate from the hyphal walls and from the septa. Holes are
dissolved in the lateral wall near the apex or near the base of the con-
taining cell and the macrospores escape. The resting spores are usually
formed in the hyphae at about the time the chitinous wall of the posterior
portion of the intestine begins to become free at the molting stage of the
host. These cells may arise by the union of two naked protoplasts or
"gametes" within the segment of the filament, the resulting zygotes
becoming thick- walled {Arundinula capitata) or the segments of the
hyphae may be binucleate and the two nuclei unite. Then a thick wall is
formed around each such zygote. The cross septa of the hypha may dis-
appear leaving the numerous resting spores free in the tube. They may
escape by a distal opening {Taeniellopsis orchestiae Poisson, 1939). In
some genera of the family Taeniellaceae the resting spores possess one
nucleus at maturity. In the family Arundinulaceae^ there are two nuclei
and in the family Eccrinaceae there are four nuclei in each zygote. The
sexual process is merely surmised in most cases. The fourth family, Amoe-
bidiaceae contains the single genus Amoebidium. The tube or sac-like
hypha may divide by oblique w^alls into 4-16 "endoconidia" (Lichten-
stein, 1917) which elongate in the mother hypha, piercing the wall and
thus forming a cluster of hyphae. More often the contents of the non-
septate hyphae divide into 2-4, rarely 8 pyriform amoebae which escape
through the dissolved apex of the hypha and creep and float in the sur-
rounding water with the larger end foremost. They soon encyst as spher-
ical cells. No flagella are apparent. A third mode of reproduction is the
formation inside the hyphae by oblique walls of uninucleate fusiform
cells, their formation proceeding distally from the base. They escape by
lateral openings in the hyphal wall. After escaping the thin membrane
becomes thickened thus forming a sort of resting spore. (Fig. 65.)

The coenocytic hyphae with cellulose walls would seem to indicate
some affinity of the Eccrinales with the Phycomyceteae. The absence of
flagella even on the naked amoeboid spores of one genus makes their
definite placement out of question without further data. The doubtful
or reduced modes of sexual reproduction give little help in determining
their relationship.

Drechsler (194()b) has described a fungus Gonimochaete horridula,
which attacks and kills free-living soil nematodes and which has some of
the characteristics of the Eccrinales, but which differs in many respects.

iLeger and Duboscq (1929a) used the name Arundinula (and the family name
Arundinulaceae) instead of Arundinella, a preoccupied name.

KEYS TO THE FAMILIES AND MORE IMPORTANT GENERA OF MUCORALES 183

Only asexual reproduction has been observed so far. In hyphal outgrowths
from short one- celled thalli within the host are produced a few to numer-
ous endoconidia with definite thin walls. These are pushed out several
at a time at short intervals until all have been discharged. In germination
they produce a small sticky knob by which they become attached to the
living nematode host, into whose body a germ tube penetrates and then
divides repeatedly to form the unicellular thaUi. Possibly this fungus may
have some relationship to this order.

Keys to the Families and More Important Genera of Mucorales

Key to the Families of Order Mucorales
Asexual sporangia, sporangioles or "conidia" aerial.
All the sporangia many-spored, with a well-developed columella; sporangium
wall relatively thin and breaking or deliquescent.

Family Mucoraceae
All the sporangia many-spored, with a moderate-sized columella; sporangium
wall thickened above and not breaking up or deliquescent.

Family Pilobolaceae
Terminal primary sporangium of the sporangiophore many-spored, with a
well-developed columella; sporangium wall thin and breaking up or deli-
quescent; secondary sporangia in the form of few-celled or one-celled
sporangioles which are usually indehiscent. Primary sporangia lacking
under unfavorable conditions, and never formed in a few genera.
Sporangioles on more or less dichotomous branches formed laterally along
the main sporangiophore. (Primary sporangium lacking in the genus
Chaetocladium.) Family Thamnidiaceae

Sporangioles on the surface of rounded or elongated heads terminating
sporangiophores apart from the primary sporangiophore. Primary spo-
rangium lacking in the genera Cunninghamella, Mycotypha, etc.

Family Choanephoraceae
Sporangia spherical, many-spored, with a basal septum and no columella.

Family Mortierellaceae
Sporangia narrow, one- to several-spored, with no columella, usually more or
less capitately borne, often breaking apart into one-spored segments.

Family Piptocephalidaceae

Sporangia reduced to one-celled, indehiscent sporangioles ("conidia") borne

singly on sterigmata arranged on one side of a branch ("sporocladium")

so as to resemble a comb. Family Kickxellaceae

Sporangia, zygospores and chlamydospores in the interior of rounded masses of

hyphae; often buried in humus or soil. Family Endogonaceae

Key to the More Important Genera of Family Mucoraceae

Sporangiophores arising from the thin or thick mat of aerial mycelium, not from

stolons. Not repeatedly forked dichotomousl^y.
. Primary sporangium present at the apex of the sporangiophore.
Sporangiophore tall, dark, unbranched, with metallic appearance.

Phycomyces
Sporangiophore slender and uniformly thick its wliole length, unbranched,
or racemosely or cymosely branched— in the latter case the branches

184 PHYCOMYCETEAE

sometimes circinately curved ; not dark-colored, nor of a metallic appear-
ance. Sexual reproduction practically isogamous. Not parasitic on fungi
(but sometimes saprophytic). Mucor

Like Mucor but parasitic upon Mucorales. Parasitella

Much like Mucor but sporangiophore much broader below, tapering upward,
somewhat metallic in appearance. Parasitic upon Basidiomycetes.
Mycelium often with short lateral spine-like branches. Spinellus
Much like Mucor but sexual reproduction very strongly heterogamous;
homothallic. Zygorhynchus

Primary sporangium missing at the apex of sporangiophore which is cymosely
branched with circinately recurved branches.
Sporangia spherical, without apophysis, spores spherical. Circinella
Sporangia pyriform, with apophysis, spores ellipsoid. Pirella

Sporangiophores repeatedly forked dichotomously; homothallic, isogamous.

Sporodinia
Sporangiophores arising from stolons.

Sporangia mostly large, mostly spherical, spores dark and striate; sporangio-
phores arise at rooting nodes of the stolon. Rhizopus
Sporangia small, pyriform, spores colorless or colored; sporangiophores
arising from the summit of the arch of curving stolons which mostly
root at the nodes.
Stolons forming a series of regular, rather short arches with the sporangio-
phores arising from their summits. Absidia^
Stolons forming longer, less regular curves, the sporangiophores mostly

clustered at their summits but some scattered singly. Tieghemella
Stolons as in Tieghemella but the sporangiophores in a whorl midway along
the curve. Mycocladus

Key to the Genera of Family Pilobolaceae

Sporangiophore swollen to a subsporangial vesicle which is broader than the
sporangium, which is violently discharged. Pilobolus

Sporangiophore uniform in thickness, without subsporangial vesicle. Sporangium
not discharged. Pilaira

Key to the Genera of Family Thamnidiaceae

Sporangiophore terminated by a primary sporangium that is many-spored and
with a columella, and bearing lateral variously branched clusters of secondary
sporangiophores with dehiscent or indehiscent sporangioles.

Sporangiole-bearing branches dichotomously forked, walls of sporangia not
strongly cutinized. Thamnidium

Sporangiole-bearing branches dichotomously forked, walls strongly cutinized.
Very strongly heterogamous. Dicranophora

Sporangiole-bearing branches in whorls, many of the hyphae terminating in
awl-like extensions. Chaetostylum

Sporangiole-bearing branches circinately curved. Helicostylum

No primary spoi-angiuin formed, main axes of sporangiophore and of the branches
terminating in awl-like extensions. Chaetocladium

2 Tieghemella, Mycocladus, and three other genera, Proabsidia, Protoabsidia, and
JAchlheimia, are often all included in Absidia.

KEYS TO THE FAMILIES AND MORE IMPORTANT GENERA OF MUCORALES 185

Ke]) to the Genera of Family Choanephoraceae

Primary sporangia with columella, terminal to an upright sporangiophore.
Sporangioles borne on the surface of spherical heads clustered at the upper
end of special sporangioliferous hyphae.
Sporangioles containing 3 (to 6) spores, resembling those of the sporangium.

Blakeslea
Sporangioles one-spored, indehiscent. Sexual reproduction of the Mucor type.

Choanephora

Primary sporangia lacking, only the indehiscent one-spored sporangioles formed.

Sporangioles borne on rounded heads on short lateral branches, mostly rough.

Fertile hyphae in a dense, tangled mass of aerial mycelium and bearing the

heads of sporangioles terminally and on lateral branches without regular

order, sporangioles smooth or rough, round or ovoid. Sexual reproduction

of the Mucor type. Cunninghamella

Fertile hyphae upright, forking into S-shaped branches, sterile at the tips

but bearing heads of spherical rough sporangioles on short lateral branches.

Sigmoideomyces.
Fertile hyphae upright, supported at the base by mostly four spreading
branches and at the top dichotomously dividing several times into branches
sterile at their tips but bearing on short lateral branches the heads of
spherical rough sporangioles. Thaynnocephalis

Sporangioles small, ellipsoidal, smooth, borne on an elongated head which be-
comes septate at maturity and with the sporangioles resembling a head of
Typha. Mijcotypha

Key to the Genera of Family Mortierellaceae

Sporangiophores erect, unbranched, or branched. Mortierella

Sporangiophores creeping. Herpocladiella

Sporangiophores short and unbranched along the sides of a hypha of indeter-
minate growth. Dissophora
{Haplosporangitmi possibly belongs to this family.)

Key to the Genera of Family Piptocephalidaceae

Main sporangiophores with large heads covered by the radiating narrow spo-
rangia.
Main sporangiophore branched, sporangia arising directly, not from sterigmata.

Syncephalastrum
Main sporangiophores stout, not branched, tapering toward the terminal head.
In most cases sporangia borne on sterigmata, one or more on each.

Syncephalis
Main sporangiophores branching freely above by true or false dichotomy with
small heads or only slight enlargements from which the sterigmata arise.
Branches straight, sterigmata one-celled. Piptocephalis

Branches more or less spirally curved, sterigmata two-celled, two-celled spo-
rangia arising from the apex of each cell. Dispira
Main sporangiophores with small or medium size heads terminal to the hyphae
and their branches, bearing on all sides two-celled sporangioles. Origin of
these latter not definitely worked out and relationship uncertain.

Spinalia

186 PHYCOMYCETEAE

Key to the Genera of the Family Kickxellaceae

Sporocladia in a whorl at the apex of the sporangiophore. Kickxella

Sporocladia pleurogenous on the sporangiophore.

Sporocladia producing sporangioles (conidia) on the upper side.

Martensella

Sporocladia producing sporangioles on the lower side. Coemansia

Key to the Genera of Family Endogonaceae

Fruiting body containing reproductive cells throughout, either sporangia, or
chlamydospores or zygospores. Endogone

Fruiting body hollow, containing chlamydospores scattered irregularly in the thin
wall. GlazieUa

Fruiting body not hollow, rather firm, chlamydospores arranged in regular
layers. Sclerocystis

Key to the Genera of Order Entomophthorales

Parasitic in the gametophytes of ferns. Completoria

Parasitic in the desmid genus Closterium. Ancylistes

Saprophytic in the excrement of frogs and lizards. Basidiobolus

Saprophytic or weakly parasitic on fungi (rarely on insects) ; the septate mycelium

extensive: conidiophores elongated, gametangia unequal, the zygospore

developed in the larger, receptive gametangium. Conidiobolus

Parasitic in insects.

Conidia produced internally in the body of the insect, not discharged with

violence. Massospora

Conidia produced on conidiophores extruded through the body wall, discharged

with violence. Entoinophthora^

Key to the Families and Genera of Order Zoopagales

Saprophytes or commensal parasites, attached to, but not penetrating through,
the membrane of the alimentary canal of aquatic insects (chiefly larvae).
Fungus filament not branched, zygospores approximately spherical.

Family Harpellaceae
Conidia curved cylindrical, with 4 basal filaments. Harpella

Conidia navicular, stalked, with one basal filament. Stachylina

Conidia tubular, borne side by side on the racket-shaped terminal cell of the
filament. Opuntiella

Fungus filament branched, zygospores navicular or biconical, stalked.

Family Genistellaceae
Conidia elongate-ovoid, with 2 basal filaments, borne on one-sided spike.

Geiiistella
Conidia long stalked, curved like a banana, in a crown at apex of plant.

Orphella
Conidia terete on a one-sided spike. Stipella

Conidia rod-shaped arising laterally on a somewhat twisted stipe, with one

basal filament. Glotzia

Conidia ovoid, with single basal filament, on one-sided spikes. Plant bulbous
at base. Graminella

'The name Empusa is not valid for this, having been used previously for an
orchid.

LITERATURE CITED 187

Parasitic by large or small haustoria or internal hyphae, in soil-inhabiting Rhizo-
pods or Nematodes. Family Zoopagaceae

Main fungus body a thick spiral structure in the cell of an amoeba, conidia
borne on slender aerial hyphae.
Conidia borne singly at intervals on long aerial hyphae. Endocochlus

Conidia borne in chains from erect aerial hyphae. Cochlonema

Main fungus body a very much enlarged conidium on the outside of the body
of an amoeba, attached by dichotomously forked internal haustoria.
Conidia in long chains on erect aerial hyphae. Bdellospora

Main fungus body an effuse mycelium, adhering to Rhizopods or Nematodes
and sending lobed haustoria into them.
Conidia in vertical chains on short lateral branches. On Rhizopods.

Zoopage
Conidia single, sessile or almost so, upright on the creeping mycelium. On
Rhizopods. Acaulopage

Conidia single on upright branches from creeping mycelium. On Rhizopods

and Nematodes. Stylopage

No conidia found but asexual reproduction by means of terminal, lateral or
intercalary chlamydospores. On Rhizopods and Nematodes. Cystopage
Main fungus body an extensive mycelium in the body of Nematodes. External
hyphae formed and bearing at right angles a number of short lateral coni-
diiferous branches which produce first a terminal conidium and then suc-
cessively in basipetal direction other conidia. Euryancale

Key to the Families of Order Eccrinales

No amoeboid spores.

Resting spore at maturity with a single nucleus. Family Taeniellaceae

Resting spores at maturity with two nuclei. Family Arundinulaceae

Resting spore at maturity with four nuclei. Family Eccrinaceae

Amoeboid spores produced, also some resting spores. Family Amoebidiaceae

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Drechsler, Charles: A new species of Helicocephalum, Mycologia, 26(1) :33-37.

PI. 4. 1934.
: Some conidial Phycomycetes destructive to terricolous Amoebae, ibid.,

27(l):6-40.P/s. 1-7. 1935a.
: Some non-catenulate conidial Phycomycetes preying on terricolous

Amoebae, ibid., 27(2):176-205. Figs. 1-5. 1935b.
: A new species of conidial Phycomycetes preying on nematodes, ibid.,

27{2) -.206-215. Fig. 1. 1935c.
: A new species of Stylopage preying on nematodes, ibid., 28(3) :241-246.

Fig. 1. 1936a.
: New conidial Phycomycetes destructive to terricolous Amoebae, ibid.,

28(4) :363-389. Figs. 1-7. 1936b.
: New Zoopagaceae destructive to soil Rhizopods, ibid., 29(2) :229-249.

Figs. 1-6. 1937.
: New Zoopagaceae capturing and consuming soil amoebae, ibid., 30(2):

137-157. Figs. 1-4. 1938.
: A few new Zoopagaceae destructive to large soil rhizopods, ibid., 31(2):

128-153. Figs. 1-7. 1939a.
: Five new Zoopagaceae destructive to rhizopods and nematodes, ibid.,

31(4):388-415. Figs. 1-5. 1939b.
: Four Phycomycetes destructive to nematodes and rhizopods, ibid.,

33(3):248-269. Figs. 1-5. 1941.
: New species of Acaulopage and Cochlonema destructive to soil amoebae,

ibid., 34(3) :274-297. Figs. 1-6. 1942.
: Several additional Phycomycetes subsisting on nematodes and amoebae,

ibid., 37(1):1-31. Figs. 1-4. 1945.
: Three new Zoopagaceae subsisting on soil amoebae, ibid., 38(2): 120-143.

Pis. 1-6. 1946a.
: A nematode-destroying Phycomycete forming immotile spores in

aerial evacuation tubes. Bull. Torrey Botan. Club, 73(1):1-17. Figs. 1-36.

1946b.
: Three Zoopagaceous fungi that capture and consume soil-inhabiting

rhizopods, Mycologia, 39(3):253-281. Figs. 1-8. 1947a.

LITERATURE CITED 189

: Three new species of Zoopage predaceous on terricolous rhizopods, ibid.,

39(4) :379-408. Figs. 1-7. 1947b.
EiDAM, Eduard: Basidiobolus, eine neue Gattung der Entoraophthoraceen,

Beitr. Biol. Pflanz., 4(2):181-251. Pis. 9-12. 1887.
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Fenner, E. Aline: Mycotypha microspora, a new genus of the Mucoraceae,

Mycologia, 24(2):187-198. Pis. 2-3. Fig. 1. 1932.
FiTZPATRicK, Harry Morton: The Lower Fungi — Phycomycetes, xi + 331 pp.

112 ^^s. New York, McGraw-Hill Book Co., 1930.
Flint, Lewis H.: Note on phototropism in Pilobolus, Am. J. Botany, 29(8) :672-

674. Fig. 1. 1942.
Gauthier, Marcelle: Sur un nouvel entophyte du groupe des Harpellac^es

Leg. et Dub.; parasite des larves d'Eph^m&ides, Compt. rend., 202(12) :1096-

1098. Figs. 1-4. 1936.
Goldring, David: The effect of environment upon the production of sporangia

and sporangioles in Blakeslea trispora Thaxter, Ann. Missouri Botan.

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Hopkins, E. W.: Microchemical tests on the cell walls of certain fungi. Cellulose

and chitin. Trans. Wisconsin Acad. Sci., 24:187-196. 1929.
Jolivette, Hally D. M.: Studies on the reactions of Pilobolus to light stimuli,

Botan. Gaz., 57(2):89-121. Figs. 1-12. 1914.
Kanouse, Bessie B.: Studies of two species of Endogone in culture, Mycologia,

28(l):47-62. Figs. 1-33. 1936.
Keene, Mary Lucile: Cytological studies of the zygospores of Sporodinia

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: Studies of the zygospore formation in Phycomyces nitens Kunze, Trans.

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Krapczyk, Hans: Die Zygosporenbildung bei Pilobolus cristallinus, Ber. dexit.

botan. Ges., 49(3):141-146. Figs. 1-2. 1931.
Leger, L., et O. Duboscq: Les Eccrinides, nouveau groupe de Protophytes

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— ■ ■, ET : Eccrinoides Henneguyi n.g. et n.sp. et la systematique des

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, ET : Harpella melusinae n.g. et n.sp. Entophyte eccriniforme

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, ET Marcelle Gauthier: Endomycetes nouveaux des larves aquatiques

d'insectes, ibid., 194(26) :2262-2265. Figs. 1-3. 1932.

ET : La spore des Harpellac^es (Leger et Duboscq), champignons

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190 PHTCOMYCETEAE

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LITERATURE CITED 191

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Zycha, H. : Mucorineae, in Kryptogamenflora der Mark Brandenburg, vol. 6a,

pp. 1-264. 114 figs. Leipzig, Gebriider Borntraeger, 1935.

THE HIGHER FUNGI: CARPOMYCETEAE

THE fungi that have been the subjects of discussion in the foregoing
chapters are frequently designated as the Lower Fungi or Phyco-
myceteae (excluding the Mycetozoa) in contrast to the often much more
highly developed forms usually designated as the Higher Fungi or Carpo-
myceteae. It may be well here to contrast the two groups as to their main
points of difference, recognizing that though these distinctions are in
general valid yet there are many forms of Higher Fungi in which these
differences are not recognizable.

Structure

1. The Phycomyceteae are prevailingly tubular coenocytes, non-
septate except at regions of injury or where reproductive organs are cut
off from the rest of the mycelium. (Exceptions may be noted in old aerial
mycelium of some Mucorales and in most Entomophtlorales.) The simpler
members of this class may consist of single cells. The Higher Fungi are
cellular, i.e., their hyphae are divided by septa into true cells, usually
uninucleate or binucleate. (Exceptions are numerous. Thus old cells
sometimes become multinucleate; on the other hand, many young hyphae
from germinating spores or young ascogenous hyphae may delay the
formation of cross walls for some time. Aside from these, numerous
species are scattered throughout the whole group in which almost all the
cells are plurinucleate.)

2. In the Lower Fungi sexual reproduction leads to the formation of a
single, usually thick-walled, oospore or zygospore. In the Higher Fungi,
wherever true sexual organs or processes can be distinguished, the union
of sperm and egg, or of cells substituted for these, leads to the production
of a many-celled structure, called by Sachs (1874) a "spore-fruit," all
the cells of which may become the reproductive cells (as the ascospores
in the asci of the order Saccharomycetales) or of which only a part are
reproductive cells, the remainder being necessary components of a more
or less complicated fruit body (e.g., the sum of the ascogenous hyphae

192

PARASITISM 193

and asci arising from the oogone of Pyronema, along with the various
hyphae of vegetative origin, all of which form the apothecium, while
only the ascospores are the reproductive cells).

3. In the Phycomyceteae, except the Mucorales and the Entomoph-
thorales and allied forms, uniflagellate or biflagellate zoospores are gen-
erally produced as a means of asexual reproduction, while in these orders
the asexual reproductive structures are clearly only modifications of zoo-
sporangia. In the Higher Fungi the asexual reproductive cells are conidia,
which are separable single cells (sometimes several-celled structures) of
the mycelium with no indication of ever having had any homology with
zoosporangia.

4. The aquatic Phycomyceteae and the terrestrial order Perono-
sporales have walls composed of cellulose (or carbohydrates closely related
to cellulose). Cellulose is lacking in the Higher Fungi, or when present
has been shown by Thomas (1928, 1930) to play only a subordinate part.
In these the basis of the wall is chitin surrounded or mixed with carbo-
hydrates and more or less fatty substances. Hopkins (1929) reports that
neither chitin nor cellulose is present in several species of the class Asco-
myceteae and of the class Basidiomyceteae, although other species of
both classes tested by him showed chitin but not cellulose to be present,
as is claimed by von Wettstein (1921) to be probably universal in these
two classes. Senft (1916) reports that the ascogenous hyphae of Chryso-
thrix nolitangere Mont., a lichen, respond to the standard cellulose tests.
The same is true, according to Honey (1936), of the disjunctors in the
chains of conidia of Monilinia. Malengon (1931) has shown that the
epispore of the spores of some Agaricaceae, e.g., species of Russula and
Lactarius, is blued by iodine reagents. He believes the epispore to contain
a mixture of cellulose and callose. This was first observed by Rolland in
1887. Klihner (1936) finds a similar reaction in the outer layer of the
internal hyphae of the pileus and stipe of some species of Mycena. The
Mucorales and Entomophthorales seem to occupy an intermediate posi-
tion in that they have chitin in their cell-walls and rarely have true
cellulose, although other carbohydrates are present in the wall.

Parasitism

The Higher Fungi are with few exceptions terrestial or epiphytic, a
very few species are truly aquatic, some being parasitic upon marine
algae, but they possess none of the characteristics of a primary aquatic
habit such as zoospores, etc. One order, the Laboulbeniales, consists
exclusively of insect-infesting parasites. The remainder are mostly either
saprophytes, or parasites upon plants (exceptionally upon animals). All
degrees of parasitism are found, varying from destructive parasitism in
which the tissues of the host are killed even before the invading hyphae

194 THE HIGHER FUNGI : CARPOMYCETEAE

reach them (e.g., Sclerotinia sclerotiorum (Lib.) de By.) to extreme cases
of balanced parasitism (e.g., some of the Ustilaginales in which the
fungus and host, together grow for a long time with little apparent harm
to the latter). One large group has developed a special type of parasitism
upon terrestrial or epiphytic fresh-water algae, forming peculiar struc-
tures which enclose the algal hosts. These are the Lichens. Some of the
most harmful fungi from the economic standpoint, the Rusts and Smuts,
are found among the Higher Fungi.

Reproduction

Asexual reproduction by means of conidia is widely distributed
throughout the various classes and orders, but seems to be entirely lack-
ing in some of the groups. The conidial production may consist of the
abstriction of a single cell from the tip of a short, unbranched conidiophore
or the conidiophore may be longer and branched. The conidia may be
produced successively at the tip, separating from the conidiophore as
soon as formed or clinging together in a mucilaginous drop or remaining
attached in a chain. A chain of conidia may produce new conidia acro-
genously, i.e., each new conidium arises from the apical conidium of the
chain, so that the basal conidium is oldest and the apical one the last
formed. This is the type of conidial formation in Cladospoi'ium, Alteniaria,
etc. On the other hand the conidial formation may be basigenous, i.e.,
each new conidium is produced at the apex of the conidiophore just
below the last formed conidium (e.g., Erysiphaceae, Aspergillaceae). A
third method of formation of chains of conidia is the almost simultaneous
rounding up of the cells of a simple or branched hypha into catenulate
conidia. These conidia then fall apart almost simultaneously (e.g., Moni-
litiia, Oospora, etc.). Where the conidia formed in this manner are small
they are often called "oidia."

The conidiophores may be scattered or crowded or enclosed in a
hollow structure provided with an apical opening (i.e., produced in a
pycnidium). If the conidiophores become laterally crowded and adherent
into a column we have a coremium, such as is formed in species of Peni-
cillium under certain environmental conditions.

The conidia themselves may vary greatly in size and shape as well
as color and number and arrangement of cells. In general we may dis-
tinguish conidia destined for distribution by air currents and those that
are distributed by other means. The latter are usually surrounded by a
sticky substance when wet and cling to anything with which they come
in contact, drying down and becoming firmly attached when dry. When
wet by rain such conidia may be scattered by the currents of water
running on the surface of the substratum or may be splashed about by
the falling rain drops and distributed by the wind which carries these

STORAGE ORGANS 195

droplets about. Insects may become contaminated by the sticky spores
and thus carry the fungus from place to place.

Chlamydospores are very frequent in many of the orders of the Carpo-
myceteae. They arise by the rounding up of mycelial cells, filled with
food substances, and the production of a thick wall. They may be inter-
calary or terminal, single or two to many in a series. In some fungi they
germinate promptly, in others they are able to remain dormant a long
time.

The mycelium usually consists of long, slender branching hyphae with
terminal growth. Usually the cell is several to many times as long as wide.
The septa are centrally perforated by a pore which may be small or may
approach in size the perforations found in the septa of the Florideae. In
the order Laboulbeniales the cells are short and broad and the whole
structure does not resemble typical mycelium. In the Phycomyceteae
where septa do occur they are formed, as in the Higher Fungi, by an
annular growth on the inner surface of the longitudinal wall which pro-
gresses inward, but instead of leaving a central pore it continues its
development until the opening is obliterated.

Buller (1933) and Brodie (1942) give the diameter of these openings
in the Carpomyceteae as of the order of 1.0 to 1.5 n. The flow of cytoplasm
through the septa has been observed by Ternetz (1900) and by Buller,
sometimes to carry small vacuoles with the flowing stream. The rate of
flow through the hyphae varies but Ternetz observed the speed of 10.5
cm. per hour in Ascophanus carneus (Pers.) Boud., and Buller the speed
of 6 cm. per hour in Firnetaria fimicola (Roberge) Griffiths and Seaver.

Storage Organs

In many species in the class Basidiomyceteae peculiar structures called
"clamp connections" occur at each septum. These will be described in
connection with that class. The mycelium may be packed together
laterally into compact strands with a firm hard outer layer, forming long
"rhizomorphs" which may extend many meters. In many species storage
organs, sclerotia, are formed in which the hyphae are packed tightly
together and the short cells are pressed mutually to become polyhedral
in shape. The adjacent cells adhere to form a parenchyma-like tissue,
more properly called pseudoparenchyma. True parenchyma which is
rare in the fungi consists of moderately thin-walled ceUs, isodiametric or
at least not greatly elongated, which have arisen through the division in
various planes of larger cells or of small meristematic cells. The walls
separating these cells are the septa between two daughter cells. Pseudo-
parenchyma arises from closely crowded or interwoven hyphae divided
by cross septa into rather short cells which by their enlargement and
mutual pressure become similar in appearance to the cells of true paren-

196 THE HIGHER FUNGI : CARPOMYCETEAE

chyma. Except for the septa separating the cells in the same hypha the
cell walls are double, i.e., walls of separate cells pressed together and
usually adhering, but not the result of division of one preexisting cell.
The interior cells of a sclerotium are filled with stored food and the outer
cells are thick-walled, almost sclerenchymatous, and usually dark in
color. Pseudoparenchyma may be found also in the fruiting bodies of
some of the Higher Fungi, such as the perithecia and apothecia of some
of the Ascomyceteae. True parenchyma is apparently present in some
fungi of this group. The body of the spore fruit is more often built up of
elongated, more or less interwoven hyphae of elongated cells. This is
often, though erroneously, as Starback (1895) points out, called prosen-
chyma. In parasitic species the mycelium often sends haustoria of various
shapes into the host cells.

Fruiting Structures

Unlike the Lower Fungi whose fruiting structures are microscopic or
at most only a few millimeters or centimeters in length (some Mucorales) ,
the fruiting bodies of the Higher Fungi often attain considerable size.
Thus Calvatia gigantea (Batsch ex Pers.) Lloyd, the giant puffball, was
recorded by C. E. Bessey (1884) as producing a spore fruit 1.6 meters
long, 1.35 meters wide, and about 24 cm. high. That of Fomes officinalis
(Vill.) Fr. sometimes (rarely it is true) reaches the height of 60 cm. and
a diameter of 15 to 20 cm. Clements (1910) reported that Polyporus
squamosus (Huds.) Fr. is "said to attain a width of 7 feet and a weight of
40 pounds." Roger Heim (1936) has described three species of Boletaceae
from South Africa and Madagascar 40 to 60 cm. in pileus diameter.
Ljungh (1804) described a species of cup-fungus from Java, Peziza cacabus
(now called Geopyxis cacabus (Fr.) Sacc), with a spore fruit 3 ft. tall,
the cup being 20 in. tall and 25 in. broad, and the hollow stipe 16 in. tall
and 3 in. thick. Specimens of a form of Agaricus arvensis Fr. collected
by the author had a pileus diameter of 30 cm. Such enormous spore fruits
produce almost incredible numbers of spores. Thus Buller (1909) esti-
mated that a puffball 40 X 28 X 20 cm. would produce about 7 trillion
(7,000,000,000,000) spores. At the same rate the enormous puffball men-
tioned above would produce about 160 trillion (160,000,000,000,000)
spores. A specimen of Agaricus campestris Fr. only 8 cm. in diameter
produced over 1,800,000,000 spores at the rate of about 40,000,000 spores
per hour. At this rate the Agaricus arvensis found by the author would
have produced about 27,000,000,000 spores. A spore fruit of Ganoderma
applanatum (Pers. ex Fr.) Pat. with an area of one square foot lower surface
produced, according to White (1920), 30,000,000,000 spores a day for
about six months, or a total of over 5,000,000,000,000 spores. On the
other hand many of the Higher Fungi have microscopic spore fruits.

KEY TO THE CLASSES OF HIGHER FUNGI 197

Coloring

In the majority the vegetative mycelium is colorless and that is true
of the reproductive structures in many cases. In rhizomorphs and the
outer layer of cells of sclerotia and sometimes in individual hyphae the
color may be dark. This color seems to reside in the cell wall and is prob-
ably related chemically to melanin. Some fungi produce pigments in the
interior of the hyphae. These may be soluble in various solvents and are
sometimes variable in color depending upon the chemical reaction. Thus
a Fusarium studied by the author (1904) and parasitic on Sesamum
orientale L. produces a red- or violet-colored pigment that turns blue
when the surrounding medium becomes alkaline. It is soluble in acids
and the red form is soluble in solutions of their salts. Litmus is the product
of the cells of one of the lichen-producing fungi, as is orcein. Some hyphae
cause the coloration of the substratum in which they are growing. Thus
wood in which Chlorociboria aeruginosa (Fr.) Seaver is growing takes on
a green color, due to a pigment secreted by the mycelium. The repro-
ductive structures are in the majority of cases colored, the pigments
in some being within the cell (e.g., the bright red color of the hymenium
of some Pezizales), in others in the cell wall. The latter is mostly the case
with the light to dark brown or almost black coloration found in the
majority of apothecia and perithecia and in the teliospores of Rusts and
Smuts, the spore fruits of Tremellales, Auriculariales, etc. The fungus
pigments, especially those occurring in the lichen-producing fungi, have
been studied by various authors but no really comprehensive modern
study has been made of the subject taking advantage of the more recent
investigations in organic chemistry.

Nomenclature

The Higher Fungi were called by Charles E. Bessey (1907) the Phylum
Carpomyceteae, i.e., fruit-producing fungi, in reference to the production
of spore fruits in this phylum. The name Eumyceteae is often applied to
this group but is here discarded in view of the fact that it has also been
applied to include all the filamentous fungi in contrast to the nonfila-
mentous Mycetozoa, Chytridiales, etc. The Higher Fungi are divided into
several classes whose distinctions are based on the type of the ultimate
reproductive cells of the spore fruits.

Key to the Classes of Higher Fungi

The ultimate reproductive cells of the spore fruit are ascospores, produced mostly
eight in number within the cell called an ascus, which starts out as a di-
caryon cell in which the nuclei fuse, the resultant diploid nucleus then
dividing meiotically until usually eight nuclei are produced, around which
the ascospores are developed. Over 42,000 species have been described.

Class Ascomyceteae

198 THE HIGHER FUNGI : CARPOMYCETEAE

The ultimate reproductive cells of the spore fruit are the basidiospores or sporidia,
borne externally on a cell called the basidium which was originally a
dicaryon cell in which the two nuclei united, or on the outside of a group
of four cells produced by the division of, or as an outgrowth from, the
cell with the diploid nucleus. About 32,000 species have been described.
With considerable reluctance the author follows the practice of most
mycologists and includes the Class Teliosporeae of the first edition of
this textbook as a distinct subclass within the Class Basidiomyceteae.
The teliospore characteristic of this subclass originates as a dicaryon cell
whose nuclei unite. From this cell grows out a thin-walled, usually four-
celled, filament, the promycelium, upon whose cells are borne the sporidia.
The limits of this subclass coincide rather closely with the Hypodermii
which form Order IV of Class IV, Coniomycetes, of Elias Fries (1832).
Ascus, teliospore, and basidium appear to be liomologous structures,
originating as binucleate cells in which karyogamy occurs, followed by
reduction division of the fusion nucleus, the nuclei thus formed becoming
the nuclei of the ascospores, sporidia, or basidiospores, respectivel3^

Class Basidiomyceteae

In addition to these classes there is another class, the Fungi Imperfecti, often
called Deuteromyceteae, consisting of fungi whose vegetative structures
or mode of asexual reproduction show their relationship to the Higher
Fungi, but which lack any sexual type of reproduction or structures sub-
stituted for such sexual reproduction. Perhaps the majority are asexual
stages of Ascomyceteae though some are undoubtedly corresponding stages
of Basidiomyceteae. Until the perfect (sexual) stages can be found, their
true relationship cannot be ascertained. About 32,000 species have been
recognized. Class Fungi Imperfecti

Literature Cited

Bessey, Charles E.: An enormous puff-ball. Am. Naturalist, 18(5) :530. 1884.
: A synopsis of plant phyla, Univ. (Nebraska) Studies, 7(4):l-99. PL 1.

1907.
Bessey, Ernst A.: Die Bedingungen der Farbbildung bei Fusarium, Flora,

93(4) :301-334. 1904.
Brodie, Harold J.: Protoplasmic continuity in the powdery mildew, Erysiphe

graminis DC, Can. J. Research, C, 20:595-601. PL 1. and Figs. 1-9. 1942,
Buller, a. H. Reginald: Researches on Fungi: vol. 1, xi + 287 pp., 5 pis. 83

figs., 1909; vol. 2, xii + 492 pp., 157 ^^s., 1922; vol. 3, xii + 611 pp., 227 figs.,

1924; vol. 4, xiii + 329 pp., 4 pis., UQfigs., 1931; vol. 5, xiii -f 416 pp., 174

figs., 1933; vol. 6, xii -|- 513 pp., 231 figs., 1934. London, Longmans, Green

and Company.
Clements, Frederick E. : Minnesota Mushrooms, Minnesota Plant Studies IV,

169 pp. 124: figs. Univ. Minnesota, 1910.
Fries, Elias: Systema mycologicum, sistens-fungorum ordines, genera et species

hue usque cognitas, vol. 3, ^•iii -{- 524 pp., index 200 pp. Greifswald, Ernest

Mauritius, 1829-32.
Heim, Roger: Observations sur la flora mycologique malgache: HI. Trois bolets

gigantesques d'Afrique et de Madagascar, Rev. MycoL, N.S., 1(1):3-18. Pis.

1-4. 1936.
Honey, Edwin E.: North American species of Monilinia: I. Occurrence, grouping

and life histories. Am. J. Botany, 23(2):100-106. Figs. 1-4. 1936.

LITERATURE CITED 199

Hopkins, E. W.: Microchemical tests on the cell walls of certain fungi. Cellulose
and chitin, Trans. Wisconsin Acad. Sci., 24:187-196. 1929.

KiJHNER, Robert: Sur la reaction a I'iode des parois des hyphes des carpophores
des Mycena, Compt. rend., 203(23) :1287-1 289. 1936.

Ljungh, Sven Ingemar: Peziza Cacabus, en ny och hesynnerlig svamp fran Java,
Kgl. Svenska Vetenskapsakad. Handl., 25:39-41. PI. 1. 1804.

Malen^on, Georges: Consid^-ations sur les spores des Russules et des Lactaires,
Bull, trimestr. Soc. mijcol. France, 47:72-86. PI. 3. Figs. 1-3. 1931.

Rolland, Leon: De la coloration en bleu developee par I'iode sur divers cham-
pignons et notamment sur un agaric, Bull. Soc. mycol. France, 3:134-137.
1887.

Sachs, Julius: Lehrbuch der Botanik, vierte umgearbeitete Aufiage, xvi + 928
pp. Figs. 1-492. Leipzig, Wilhelm Engehnann, 1874.

Senft, E. : Beitrag zur Anatomie und zum Chemismus der Flechte Chrysothrix
Nolitangere Mont., Ber. deut. hotan. Ges., 34(8) :592-600. PI. 17. 1916.

Starback, K. : Disconiyceten-Studien, Bihang til Handl. Kgl. Svenska Vetenskaps-
akad., Band 21, Afd. Ill, No. 5, 1895.

Ternetz, Charlotte: Protoplasmabewegung und Fruchtkorperbildung bei
Ascophanus carneus Pers., Jahrb. wiss. Botan., 35:273-312. PI. 7. 1900.

Thomas, R. C: Composition of fungus hyphae: I. The Fusaria, Am. J. Botany,
15(9) :537-547. 1928; II. Sclerotinia, ibid., 17(8):779-788. 1930.

VON Wettstein, Fritz: Das Vorkommen von Chitin und seine Verwertung als
systematisch-phylogenetisches Merkmal im Pflanzenreich, Sitz. her. Akad.
Wiss. Wien, Math, naturw. Klasse, Abt. I, 130(1) :3-20. 1921.

White, J. H.: On the biology of Fomes applanatus (Pers.) Wallr., Trans. Roy.
Can. Inst., 12(2):133-174. Pis. 2-7. 1920.

CLASS ASCOMYCETEAE: LABOULBENIALES
AND DISCOMYCETES

Introduction

THE members of the two extremes of this class have Httle in common
beyond the production of the ascus. The type of sexual union (which
is often absent), the plan of the spore fruit, even the nature and size of
the vegetative mycelium vary tremendously. At the one extreme we find
the Yeasts (Order Saccharomycetales) in some of which the unicellular
plant becomes transformed directly into an ascus, while near the other
extreme are the "Discomycetes" (Lecanorales, Pezizales, etc.) in which
there is a well-developed mycelium and in some species a sexual union of
a nonmotile sperm with a trichogyne, leading to the production of a
well-organized apothecium with many asci.

It is therefore essential to study the ascus, as being the one structure
common to all Ascomyceteae. Omitting the apogamous forms for the
present, we find the young ascus to be a binucleate cell well supplied
with food. The two nuclei are usually considerably larger than those of
the vegetative mycelium. They fuse, forming a diploid nucleus with
double the number of chromosomes found in each of the original pair.
This nucleus usually enlarges quite considerably. It divides meiotically
to form four nuclei which in the great majority of cases divide again. In
a few species the nuclear division may be repeated until 16, 32, 64, or
more nuclei are present in the young ascus. In one species of Schizothecium
(Pleurage) the number, according to L. M. Ames,^ is 512, while in Thele-
bolus stercoreus Tode ex Fr. the number is over 1000. A part of the ascus
cytoplasm gathers around each nucleus and is soon set off from the re-
maining cytoplasm (epiplasm) by a cell wall thus forming the ascospore.
The epiplasm may assist in the formation of the outside layer of the
ascospore wall (epispore) which is often beautifully sculptured. The

1 In a letter to the author.

200

INTRODUCTION

201

cytoplasm of the ascospore builds the endospore, the inner layer of the
spore wall. The spore nucleus may subsequently divide, usually followed
by septum formation, so that the ascospore may eventually be two-celled

Fig. 66. Ascomyceteae. Variations in
ascospore number in asci of various
species. (A-C) Tuber candidum Hark.
1-spored, 2-spored, and 4-spored asci, re-
spectively. (D) Septotinia podophyllina
(E. & E.) Whetzel. 8-spored ascus. (E)
Dipodascus uninudeatus Biggs. Multi-
sporous ascus. (F) Thelebolus stercoreus
Tode ex Fr. Section through apothe-
cium showing the single ascus with 1000
or more ascospores. (A-C, after Gilkey:
Oregon State Monographs. Studies in
Botany, 1:1-63. D, after Whetzel: Myco-
logia, 29(1):128-146. E, after Biggs: My-
cologia, 29(l):34-44. F, after Ramlow:
Botan. Ztg., 64.(1) -.85-99.)

or even multicellular, though perhaps the one-celled condition is the more
frequent, as it is probably the more primitive. (Fig. 66.)

The ascus varies in shape from cylindrical or clavate in those forms
with a well-developed hymenium, to ovoid or subglobose in those in
which the asci are scattered or only loosely clustered. The ascospores
escape in various manners. In many cases, particularly in the forms with

202

CLASS ASCOMYCETEAE

Fig. 67. Ascomyceteae. Types of dehiscence of ascus. (A) Typical operculum.
(B) Bilabiate opening, modification of operculum. (C) Inoperculate opening by
softening and bursting of apex. (D-G) Contraction of external wall and expansion and
lateral rupture of inner wall and discharge of spores in Pyrenophora. (H, I) Ascus of
Myriangium duriaei Mont. & Berk. (H) Unexpanded. (I) Inner wall expanded, outer
wall contracted. (A-C, after Seaver: The North American Cup-fungi (Inoperculates).
D-G, after Atanasoff: Mycologia, 11(3):]25-128. H-I, after Fetch: Brii. Mycol. Soc.
Trans., 10:45-80.)

a typical hymenium, the asci absorb water as they reach maturity and
become considerably distended. At the apex there is an area which under-
goes softening and stretching until the wall suddenly gives way under
the pressure from within the ascus, permitting the escape of the epiplasm,
ascospores, and vacuolar liquid while the ascus wall contracts. The asco-
spores may be shot off for a distance of several centimeters, many times
the length of the ascus. In the family Pezizaceae the apex of the ascus
develops a little lid (operculum) which is forced out, often remaining
attached by one edge like a trap door. In many Ascomyceteae the whole
ascus undergoes digestion at maturity, thus setting free the ascospores in
a mucilaginous liquid. Other methods of ascus dehiscence or rupture have
been reported by Atanasoff (1919), Falck (1916, 1923), Ziegenspeck
(1926), and other investigators. (Fig. 67.)

The spore fruits of the Ascomyceteae may be classified in general
as either apothecia or perithccia or as stromatic structures not referable
to either of these forms. In addition there are structures which fit none
of these categories such as the naked asci of the order Saccharomycetales
and the spore fruits of the order Laboun)eniales.

In the typical apothecium we find a disk or saucer-shaped or even

INTRODUCTION 203

cup-like structure usually from a few millimeters up to several centimeters
in diameter. The texture is usually fleshy, fragile to tough, sometimes
leathery, and the color from pale brown to black, sometimes red, yellow,
or other colors, or even colorless. The upper surface constitutes the
hymenium, a layer of elongated cells standing at right angles to the sur-
face like a palisade. It consists of asci intermingled with supporting and
protective cells or hyphae, the paraphyses. Immediately below the
hymenium is a layer, thin or fairly thick, the hypothecium, consisting
mainly of light colored hyphae running parallel to the surface of the
hymenium and from which the asci and paraphyses arise. Often sharply
contrasted with the hypothecium but sometimes grading into it is the
excipulum which makes up the larger part of the basal portion of the
apothecium. Its tissue may be pseudoparenchymatous or may be formed
by interwoven hyphae. The outer (lower) surface may be filamentous or
may resemble an epidermis. Varying from this type we may find apothecia
borne on stipes (as in Sclerotinia) or the hymenium may be convex (as
in Pyronema) . In some cases the body of the apothecium is bent back
along the stipe so as to form a clavate structure with the upper portion
covered by an external hymenium {Geoglossum and Morchella) . In other
cases the apothecium is subterranean and variously folded internally to
form passages and chambers lined by the hymenium (various Tuberales) .
The apothecium proper is the product of the growth of the hyphae
adjacent to the ascogone, when this organ is present. It may develop
upon, underneath or within a more or less fleshy stroma or the stroma
may be entirely absent (most Pezizaceae). Corner (1929-1931) has made
a detailed study of the mode of growth and development of various types
of apothecia and concludes that their structure indicates relationship
to algal ancestors, possibly a group ancestral to the present Florideae.
(Fig. 68.)

The typical perithecium is small, usually less than a millimeter in
diameter, and more or less spherical in shape. It is more often dark-
colored and somewhat hard and brittle, though not always so. Thus in
the forms customarily included in the order Hypocreales, the perithecium
may be bright-colored and fleshy or leathery. Usually, but not always,
there is an apical opening, the ostiole, through which the ascospores
eventually escape. It may be a simple opening or may have a low lip or
be drawn out to a long slender neck. As limited by Nannfeldt (1932) the
true perithecium is lined over the whole inner surface or only in its basal
portion by a hymenium composed of thin-walled asci (sometimes thick-
ened at the apex) intermingled with true paraphyses and with periphyses
in the ostiolar region. Julian Miller (1928) has shown that the true wall
of the perithecium lies within a stromatic structure which may form
simply a thin, darker-colored external layer or may form a massive struc-

204

CLASS ASCOMYCETEAE

C imi

^U ^"^ V

^i'

W//

iH.

.^.^■^\^;i^,v:;^v^

Fig 68. Pezizales, Family Pezizaceae. (A-D) Diagram of development of angio-
carpic apothecium. (E-H) Development of gymnocarpic apothecium. (I) Apothecium
of Ascobolus stercorarins (Bull.) Schroet. (J) Apothecium of Ascophanus granuliformis
(Cr.) Boud. (Courtesy, Corner: Brit. Mycol. Soc. Trans., 14:275-291.)

INTRODUCTION 205

ture within which numerous perithecia are imbedded or on which the
perithecia are seated, each with a thin outer stromatic layer. The true
perithecial wall is colorless or light-colored and is formed from one or
more layers of hyphae arising from the supporting cell of the oogone or
antherid, producing a hollow structure surrounding the ascogonium and
the ascogenous hyphae and the asci which arise from it.

In the works of the older mycologists other structures were also called
perithecia, which they often resemble very greatly. The studies of von
Hohnel (1902-1923), Theissen (1913), Nannfeldt (1932), and others have
shown that these structures are entirely stromatic, without any true
perithecial wall, and with single asci or tufts of asci without paraphyses,
arising in cavities of the stroma. Fungi with this type of spore fruit form
the group Ascoloculares of Nannfeldt (1932) in contrast to the Asco-
hymeniales which produce apothecia or perithecia in the sense indicated
above. A third type of ascocarp with perithecium-like structures, usually
without any ostiole, and with the asci scattered throughout the interior,
neither in tufts nor forming a hymenium, is considered by some my-
cologists to be a true perithecium, by others to be of a different nature.
Fungi with this type of spore fruit form the Plectascales of Nannfeldt and
others.

What seems to be, in the author's opinion, a rather primitive but
characteristic type of sexual reproduction is that described by Higgins
(1936) in MycosphaereUa tulipiferae (Schw.) Higgins. In this species, as in
other species of the genus studied by the same investigator (1914, 1920,
1929), the male gametes are nonmotile, thin-walled sperm cells produced
usually by fours within sperm mother-cells in the interior of more or less
spherical, hollow spermogonia. The sperms are imbedded in a mucilagi-
nous mass and escape through an apical opening in the spermogonium
as the mass expands with the absorption of moisture. Within a loose mass
of hyphae an archicarp is formed consisting of a spherical or ovoid cell,
the oogone, with a single large nucleus, and a trichogyne, extending as a
slender hypha several times the length of the oogone. The loose hyphae
surrounding the archicarp grow and those at the exterior eventually
cohere into a dark-colored, firm outer wall, the interior hyphae forming
a pseudoparenchymatous mass of thin-walled colorless cells. In the mean-
time one or more sperm cells have adhered to the trichogyne which forms
a papilla at whose tip an opening is formed through which the sperm
nucleus enters and passes down into the oogone. The male nucleus gradu-
ally enlarges as it progresses and eventually the two nuclei are approxi-
mately equal in size and side by side in the oogone. The trichogyne dis-
integrates, a wall cutting it off from the oogone. The latter enlarges and
becomes more or less lobed, the two nuclei in the meantime dividing
conjugately many times. From the lobed oogone arise ascogenous hyphae

Fig 69 'Ascomyceteae. Supposedly rather primitive type of sexual reproduction
in Mycosphaerc4la tuUpiferae (Schw.) Higgins. (A) Spermogonium showing some sperm
mother-cells with several contained sperms as well as such cells with sperms emerging

{Continued on facing page.)

206

INTRODUCTION 207

containing pairs of nuclei and forming asci at their tips by means of hooks
or croziers in the manner described below in Pyronema. As the asci en-
large, the thin-walled hyaline pseudoparenchymatous cells of the peri-
thecium are destroyed except for a number of slender hyphae at the top
(periphyses) which seem to play a part in the formation of an apical
opening (ostiole) in the spore fruit, through which the mature asci pro-
trude as they discharge their ascospores. (Fig. 69.)

An appreciation of the characteristic sexual reproductive processes
in the more complex Ascomyceteae can perhaps best be obtained by a
study of the phenomena in Pyronema omphalodes (Bull, ex Fr.) Fckl.
This is by no means a very primitive form nor is it a simple structure. It
illustrates, however, most of the features that occur in this class. This
fungus is found in nature most frequently on patches of soil where there
has recently been a fire, such as the site of a camp fire. It also appears
frequently in greenhouses on flower pots that have been steamed to
sterilize the soil. At first there appears a thin whitish, moldy growth on
which arise groups of orange-colored apothecia which give the whole sur-
face of the soil an orange color lasting only a few days, after which the
fungus disappears, to be followed by other fungi. The mycelium is color-
less and septate, with its cells mostly multinucleate. Tufted branches are
produced, each bearing terminally a more or less spherical, multinucleate
oogone, from whose apex there grows out a curved hypha, also multi-
nucleate, the trichogyne. From one of the basal cells supporting the
oogone arises an obovoid or clavate multinucleate antherid. The tricho-
gyne grows to the antherid and coils upon or around its apex. An opening
is then formed from one to the other. The majority of the hundred or
more antheridial nuclei pass into the trichogyne, whose nuclei have al-
ready begun to degenerate, and then through an opening in the septum at
the base of the trichogyne into the oogone which itself contains 100 to
200 nuclei. Here they pair with the oogone nuclei. According to Harper
(1900) and to Gwynne-Vaughan and Williamson (1931) the paired
nuclei fuse, forming about half as many diploid zygote nuclei. According
to Claussen (1912) they do not fuse but merely pair closely. According
to Dangeard (1907) and his followers no opening is formed between the

Fig. 69 — {Continued)

(B) Spermogonium with sperm cells mostly discharged. (C) Very young perithecium
showing oogone with one large nucleus and short trichogyne. (D) Apical portion of
fully developed trichogyne with attached sperm (a) and the small trichogyne nucleus
(b). (E) Oogone and lower half of trichogyne showing sperm nucleus (a) and tri-
chogyne nucleus (b). (F) Oogone containing two nuclei (male and female). (G) Oogone
with eight pairs of nuclei, each in a mass of denser cytoplasm. (H) Ascogenous hyphae,
branching out from oogone. (I) Early stages of ascus formation on branching asco-
genous hypha. (J, K) Nearly mature and mature perithecia. (Courtesy, Higgins: Am.
J. Botany, 23(9):598-602.)

208 CLASS ASCOMYCETEAE

trichogyne and the antherid (which he considers to be a degenerate struc-
ture and calls the trophogone), and the nuclei in the antherid and tricho-
gyne disintegrate in situ, the oogone nuclei then arranging themselves in
pairs. Following this stage all agree that soon 10 to 20 buds appear on
the surface of the oogone and elongate to become ascogenous hyphae into
which the diploid zygote nuclei or the pairs of haploid nuclei pass until
many nuclei are present in each hypha. The nuclei probably divide in the
oogone as some of them do in the ascogenous hyphae. The latter elongate,
forking somewhat. Eventually septa are formed, producing cells that are
plurinucleate toward the base of the hypha and fewer nucleate toward
the apex where the last few cells are binucleate. In the meantime from
the cells supporting the tufts of oogones and antherids there have been
growing outward and upward numerous hyphae which intermingle with
the ascogenous hyphae derived from the oogone and also form a mass of
external hyphae.

From the terminal binucleate cell of each ascogenous hypha a lateral
branch forms just beneath the apex and the two nuclei divide simul-
taneously (conjugate division) so that the lateral cell also becomes a
dicaryon (binucleate) cell. This may be repeated. Eventually the nu-
merous terminal dicaryon cells thus produced proceed to the production of
the asci. The cell curves back upon itself like a hook, with a pair of nuclei
in the curve. The nuclei divide conjugately and cross walls are formed,
leaving two of the nuclei (a daughter nucleus of each of the two original
nuclei) at the curve and one daughter nucleus in the cell at the tip of the
hook and another daughter nucleus in the cell cut off at the base of the
hook. The two nuclei in the curve of the hook fuse while the cell elon-
gates. This is the young ascus. The basal and apical cells of the hook may
fuse and then elongate and form a new hook and a new ascus, etc. The
fusion nucleus of the young ascus is diploid or tetraploid according to
the interpretation as to the presence or absence of nuclear fusions in the
oogone. This nucleus undergoes three successive divisions to produce
eight nuclei. The first two divisions are reduction divisions (meiosis)
according to either theory, the third division being considered the final
division of a second meiosis by the advocates of the tetraploid nature
of the young ascus nucleus. By both theories the eight resulting nuclei
are haploid. From the centrosome remaining in close proximity to each
of the eight nuclei fibrillae appear to radiate and certain of these rays
curve downward around the nucleus, at some little distance from it,
apparently delimiting a mass of cytoplasm surrounding the nucleus from
the remainder of the cytoplasm of the ascus, the epiplasm. Along this
delimiting surface the ascospore wall is laid down. In the meantime the
asci have been elongating as have the vegetative hyphae surrounding
and between them. The latter become the paraphyses while the former

INTRODUCTION

209

Fig. 70. Pezizales, Family Pezizaceae. Pyronema omphalodes (Bull, ex Fr.) Fckl.
Sexual reproduction. (A) Group of antherids and oogones. (B) Section through
oogone, trichogyne, and apex of antherid, showing opening of latter into the tricho-
gyne. (C) Similar section, later stage, showing a young, forked, ascogenous hypha.
(D) Ascogenous hypha from oogone to ascus. (E) Ascogenous hypha showing two
hooks. (F) Young ascus with its single diploid nucleus. (A, B, E, F, after Harper:
Ann. Botany, 14(55) :32 1-400. C-D, after Claussen: Z. Botan., 4(l):l-63.)

210 CLASS ASCOMYCETEAE

produce the marginal tissues of the apothecium. The body of the apothe-
cium consists then of the several oogones and the branched ascogenous
hyphae which grew out of these, of the antherids, and of the vegetative
hyphae which arose from the supporting cells of the oogones and an-
therids. These latter form the main body of the apothecium as well as
its paraphyses. In the excipulum these hyphal cells by lateral enlargement
and mutual pressure form a pseudoparenchymatous tissue. (Fig. 70.)

It may seem strange that so common a species, the object of numerous
investigations by different investigators, should still be the subject of
so much disagreement. Perhaps the difficulty of staining well the rather
small nuclei and the rapidity of the progress of the sexual phenomena
are responsible for the greater part of the difficulty encountered. To this
must be added the fact that many of the stages of development, if the
exact sequence is not certain, could be interpreted differently if considered
as belonging to an earlier or later stage. Furthermore, an investigator,
with the best will possible, is apt to interpret what he sees in the light
of what appears to him to be the most logical series of events.

Using the phenomena just described for Mycosphaerella and Pyronema
as a basis for comparisons we find that sexual reproduction has been
modified in several different ways in the Ascomyceteae. Thus the antherid
when present may not be a functional organ. This is clearly the case in
the variety inigneum of Pyronema omphalodes in which W. H. Browm
(1915) has shown that there is no opening between antherid and tricho-
gyne and frequently no contact. Dangeard denies the functioning of the
antherid in the whole class except in the order Saccharomycetales. He
accounts for the pairs of nuclei in the oogone and ascogenous hyphae as a
pairing of the female nuclei, which seems to be beyond doubt the case
in the variety of Pyronema just mentioned. For Dangeard the only
nuclear fusion is that occurring in the ascus. For those following Claussen
this is the true nuclear fusion (karyogamy), but the union of antherids
and oogone is looked upon as a true sexual fusion also (cytogamy).
Harper and Gwynne-Vaughan and Williamson believed cytogamy and
karyogamy to occur one just after the other, with a second nuclear fusion
occurring in the ascus.

There is a marked tendency toward the production of a more or less
coiled series of cells, usually considerably greater in diameter than the
cells of the vegetative mycelium and often tapering to a long slender,
multicellular trichogyne. Such a structure is called an ascogonium and the
cell out of which the ascogenous hyphae bud may properly be considered
the true oogone. This may be multinucleate or uninucleate. In the Laboul-
beniales, a few genera of the Lecanorales, and apparently also in a few
of the Pezizales and Sphaeriales, minute nonmotile sperms are produced
internally or externally on short antheridial branches and upon reaching

INTRODUCTION 211

the trichogyne fuse with it, the sperm nucleus entering and passing from
cell to cell until it enters the oogone. In other cases the trichogyne fuses
with an antherid without the formation of separate sperm cells and the
nucleus or nuclei from the antherid enters the trichogyne and eventually
reaches the oogone. All degrees of reduction may be found from a multi-
cellular ascogonium with a long trichogyne to a one-celled oogone and
one-celled trichogyne as it occurs in Pyronema. In many cases the tricho-
gyne is lacking, so that the antherid comes into direct connection with
the oogone. Sometimes no antherid at all is formed. In such a case pairing
of the nuclei often does not occur in the oogone but the female nucleus
divides and the nuclei pass out in pairs into the ascogenous hyphae. In
Asco'phanus granulatus (Bull.) Speg. {Humaria granulata Quel.), which
does not possess an antherid, Gwynne-Vaughan and Williamson (1930)
report that the oogone nuclei unite by pairs and the resultant zygote
nuclei enter the ascogenous hyphae. Apparently a cell of the series of
ascogonial cells may in some cases be substituted in function for an
antherid, its nucleus taking the place of the antherid nucleus. Sometimes
no recognizable oogone or ascogonial cells can be found. Some of the
vegetative hyphae of the spore fruit become converted in a manner not
known into ascogenous hyphae with dicaryon cells. In many families of
the Ascomyceteae the terminal cell or cells of the ascogenous hypha
become asci without the formation of a hook as described above in Pyro-
nema. The two nuclei of the cell fuse and the cell enlarges terminally or
laterally, with successive nuclear divisions and ascospore development. In
the sexual species of the order Saccharomycetales two cells fuse to form
a single ascus and no ascogenous hyphae are produced. Kharbush (1927)
reports^ a similar origin of the asci in the highly developed apothecium
of Botryotinia fuckeliana (de Bary) Whetzel. In this apothecium, according
to him, there are no distinguishable ascogenous hyphae. At the base of the
hymenium the apices of adjacent hyphae unite and the nuclei fuse, thus
giving rise to the young asci, one fusion of paired hyphae for each ascus.
Greis (1940) has found a quite similar origin of asci in two species of
Morchella studied by him. In the subhymenial layer, or shortly below it,
terminal multinucleate cells of adjacent vegetative hyphae unite and
grow out to form a stouter binucleate cell, with one nucleus contributed
by each hypha. This cell may enlarge and become the ascus or the nuclei
ma> divide conjugately with the production of a short ascogenous hypha,
whose terminal cell becomes the ascus, without the formation of the
usual hook. The opposite extreme is found in some species of Taphriiia
{Exoascus) in which Miss Wieben (1927) has shown that the ascospores

2 In view of the entirely different mode of sexual reproduction reported by Drayton
(1934) for Stromatinia gladioli (Drayton) Whetzel it is evident that the process in
Botryotinia fuckeliana needs re-examination.

212 CLASS ASCOMYCETEAE

are of two opposite sexual tendencies, four of each in each ascus. These
ascospores, or the spores that bud off from them, give rise to slender germ
tubes which fuse with those from spores of opposite sexual tendency,
producing a dicaryon mycelium which becomes the vegetative mycelium
within the host. Eventually some of the cells of this mycelium enlarge,
the nuclei fuse and the asci are formed. Thus in Botryotinia fuckeliana, if
Kharbush's report is correct, the mycelium and the apothecium lack
entirely the dicaryon phase except as the cells fuse to initiate the asci,
while in Taphrina the whole vegetative mycelium is of dicaryon nature.

Some botanists suggest that the production of nonmotile sperms which
fuse with trichogynes and the budding out of ascogenous hyphae from
the oogone are indications that the Ascomyceteae may have descended
from some algae related to the Red Seaweeds (Florideae). Other botanists
(e.g., Gaumann, 1926; Atkinson, 1915; Nannfeldt, 1932) considered the
nonmotile cells that fuse with the trichogyne to be merely modified co-
nidia which have been substituted for antherids, just as certain fusions
of vegetative cells have taken the place of the union of sexual organs in
the Class Basidiomyceteae. Dangeard (1907) and the Moreaus (1926,
1928) deny any fusion of sperms and trichogyne, at least so far as any
transference of nuclei occurs. Since the author follows those that consider
the production of a trichogyne and nonmotile sperms and of an oogone
producing numerous ascogenous hyphae to be primitive characters for the
Ascomyceteae the orders will be arranged in a sequence according to that
viewpoint. This matter will be discussed in detail in Chapter 17. Those
botanists who take the opposite viewpoint and consider the occurrence
of sperms and trichogynes as representing no more than accidental con-
vergence of evolutionary development in both Ascomyceteae and Flo-
rideae would probably prefer to start with the Order Saccharomycetales.

Order Laboulbeniales. These are minute, almost microscopic para-
sites upon insects. They develop externally upon the host except for a
haustorium or "foot" that is rooted in the chitinous body wall of the
host or less often may penetrate it and form a branching hyphal growth
in the body cavity. In the more usual form the foot usually enters the
body wall at a pore and thus obtains an ample supply of food without
penetrating clear into the body cavity. The fungi vary from plants with
only a few cells in number and considerably less than 0.1 mm. in height
to forms with hundreds of cells and 2 or 3 mm. tall. The cell walls are
usually thick and firm, often dark in color. Between adjacent cells which
have arisen by the division of a common parent cell a perforation in the
septum is distinctly visible as is usual in the Class Florideae. The plant
may consist essentially of a row of cells which give off laterally some
branched filamentous appendages and a female reproductive branch. On
or near the appendages are borne the antherids. This simple type of struc-

ORDER LABOULBENIALES 213

ture may become more complex by the longitudinal division of the cells
to form a body several cells in thickness from whose sides the appendages
and the male and female organs may arise. In Zodiomyces vorticellarius
Thaxter the main plant body is multicellular and widened at the top,
bearing on the flattened upper surface many filamentous appendages and
the sexual organs. Some of the larger forms lie prostrate on the body of
the host, rooting at various points by means of rhizoids. (Fig. 71.)

In most of the genera of the order the antherids are flask-shaped
organs. The apex opens and a uninucleate sperm is pushed up into the
neck by the division of the nucleus and cytoplasm of the body of the
antherid. By successive formation of sperms in this way those previously
formed are pushed out of the neck. They are apparently naked cells, en-
tirely devoid of cilia or flagella. Sometimes several flask-shaped antherids
open into a common cavity with a single opening to the outside. In a
few genera the sperms are exogenous, being produced by the abstriction
of a terminal cell of a short, slender branch from an appendage. Such
sperms appear to possess a very thin cell wall. (Fig. 7 IE, H.)

The female reproductive branch ("archicarp" of some authors) usu-
ally consists of a row of three cells, from the base to the apex respectively
the oogone ("carpogenic cell"), trichophore, and trichogyne. The first
two are nearly or completely surrounded by, usually, a single layer of
closely adhering protective cells. The trichogyne is usually one-celled
but may divide into several cells and is simple or extensively branched.
Perhaps in most cases the sperms are brought into contact with the
trichogyne by the active movements of the insect host as it brushes
against surrounding objects or other insects. Possibly in the case of
aquatic insects, or those frequenting wet places, water currents may bring
about the transfer of sperms to the trichogyne. In the genus Zodiomyces
the elongated trichogyne seeks out and unites with the sperm. In some
species antherids are unknown and the development of the oogone is
probably apogamous, (Fig. 71A-C.)

After fertilization the oogone divides into about three cells, the bi-
nucleate middle cell of which now buds out laterally on all sides to form
numerous binucleate asci in which the two nuclei fuse and then divide
in the usual way to form eight nuclei. All eight or only four of these
nuclei serve as the centers of origin of the ascospores, in the latter case
the other four nuclei undergoing degeneration. The ascospores usually
are elongated and become two-celled. The ascus walls digest and leave
the numerous ascospores in a probably somewhat sticky gum in the cavity
of the considerably enlarged spore fruit whose walls have increased in
thickness. Eventually the ascospores are discharged between the apical
cells and because of their sticky walls adhere to objects with which they
come in contact, such as the body of another insect. In the latter case

214

CLASS ASCOMYCETEAE

Fig. 71. Laboulbeniales, Family Laboulbeniaceae. Stigmatomyces haeri (Knoch)
Peyritsch. (A) Mature plant showing at the left the appendage with sevei-al antherids
and at the right the enclosed oogone with its papillate trichogyne. (B) The fertilized
oogone has divided into two basal cells and several upper cells. (C) Asci are budding
out of some of the upper cells. (D) An ascus. (E) Two antherids with escaping sperms.
(F) Amorphomyces falagriae Thaxt., adjacent male and female plants. (G-J) Zodio-
myces vorticellarius Thaxt. (G) Mature plant. (H) Antheridial branch with exogenous
sperms. (I) Oogonial branch with trichogyne and attached sperm. (J) Mature asco-
carp. (After Thaxter: Mevi. Am. Acad. Arts Sci., 12:195-429.)

ORDEE LECANORALES (thE DISK LICHENS) 215

they germinate, one cell becoming the foot and the other developing the
remainder of the plant. The full nuclear phenomena of fertilization have
not been worked out because of the difficulty of finding the fungi in the
proper stages of development and the extremely difficult technique of
sectioning and staining.

Only a very few genera and but few species of this order were known
until the monumental work of Dr. Roland Thaxter, published in succes-
sive parts in 1895, 1908, 1924, 1926, and 1931, revealed the fact that this
order contains hundreds of species, dozens of genera, and several families.
Since the appearance of the first volume of this marvelous work other
mycologists also have added numerous species to those described by Dr.
Thaxter. Not only are these fungi found on beetles (Coleoptera) but also
on Hymenoptera, Diptera, and various other orders of insects. In view
of the fact that of the 600,000 to 1,000,000 described species (and possibly
as many or more as yet undescribed species) of insects only a few thousand
have been examined for the presence of these parasites it seems reasonable
to suppose that the number of species, genera, and even families of this
order may be greatly increased in the future.

The relationship of Laboulbeniales to the other orders of Ascomy-
ceteae is not very close. The spore fruit is unlike that of any other mem-
bers of this class and the vegetative structure finds few analogies. In the
sexual reproduction the formation of functional, separable sperm cells
is known in Collema and other lichens and in Pezizales, Sphaeriales, and
other groups. In most of these however the sperms are not produced
endogenously as naked cells as is true of most of the Laboulbeniales but
seem to have thin walls even in those cases where endogenous formation
does occur. The "archicarp" of the Laboulbeniales reminds one remark-
ably of the condition in some of the Florideae where the archicarp is
surrounded by protective cell layers with merely the trichogyne exposed.

The following orders: Lecanorales, Pezizales, Tuberales, Hysteriales,
and Taphrinales all produce spore fruits that may be considered as typical
or modified apothecia. They are often or in part included under the group
name Discomycetes.

Order Lecanorales (The Disk Lichens). These constitute a large group
of organisms which have in common the production of apothecia and
which show a specialized form of parasitism on land species of Chloro-
phyceae and Myxophyceae. The validity of the maintenance of this group
apart from the Pezizales is, to say the least, very doubtful, but until the
reproductive processes, especially the behavior of the sexual nuclei, are
better known in both groups it is perhaps better to follow custom and
consider the two orders separately. Possibly when such studies have been
carried out in all the more important genera that produce apothecia the
system of classification of both orders will have to be entirely revised.

21 G CLASS ASCOMYCETEAE

As long ago as 1887 Alfred Moller reported that he had been able to grow
various, lichens in culture media in the absence of algae, and that these
produced small thalli, up to 1.5 cm. in diameter in some cases, with
typical cortex and medulla. In some tests the development of typical
spermogonia was observed. His experiments had to be cut short after four
months so that the further development of the still growing fungi could
not be followed.

Vegetatively the Lecanorales vary from a loose branching mycelium
penetrating in all directions the gummy colony of a species of Nostoc,
the form of that colony setting the limits to the size and shape of the
lichen body {Collema), to a very complexly branched, firm thallus with
a cortical outer layer of hyphae enclosing the algal hosts and thus making
possible unlimited growth, without reference to the natural shape of the
unparasitized algal colony.

The mycelium is in general slender, light-colored, septate, and branch-
ing. The septa are centrally pierced by a rather minute perforation. In
the forms with a cortex (the majority of lichens) the mycelial cells com-
posing it are short, broad, rather thick-walled and compacted together
into a pseudoparenchyma one to several cells in thickness. In the in-
terior of the lichen the mycelium is filamentous and loose. Usually the
algal hosts are found in definite layers. In some cases definite penetration
of the algal cell by the mycelium can be observed. Such cells are even-
tually killed. Geitler (1933) reported that more often no such penetration
can be seen but that the mycelium is applied to the host cell in the manner
of an appressorium. Just how the fungus draws its nourishment from the
alga in the absence of direct penetration is a matter of conjecture. Possibly
some substance secreted by the fungus increases the permeability of the
plasma membrane of the algal cells, thus permitting sugars and other
soluble food stuffs to diffuse out from the cell to be picked up by the
fungus. The fungus furnishes a certain amount of protection to the alga
and probably gives it a more equable habitat, protecting it from the rapid
extremes of drying and moisture, sun and shade, heat and cold. In so
far as this is true it is to the advantage of the alga. On the other hand,
however, the constant tribute levied in the form of food substances diffus-
ing out from the cells must reduce their vigor somewhat. It is worthy of
note that algae whose free-living development includes zoospore forma-
tion as a normal mode of reproduction usually have this entirely sup-
pressed, reproduction being limited mainly to fission.

The earlier students of lichens consider the enclosed algal hosts to be
a part of the lichen organism, possibly reproductive in nature, hence the
term "gonidia" applied to them. By some they were considered to be
the photosynthetic organs of the lichen. Their similarity to algae was
early noted and Schwendener (18G7, 18G8) supplied the evidence that

ORDER LECANORALES (tHE DISK LICHENS) 217

they were truly algae. He and others following him were able to synthe-
size lichens from cultures of algae and lichen ascospores (Bonnier, 1889).
In spite of this evidence some lichenologists were loath to give up the old
idea and as late as 1913 Elfving, observing green bodies within the hyphae
of some lichens, maintained that they were integral parts of the lichen.
Liro (1914) confirmed the occasional presence of these green bodies within
the hyphae but beheved them to be portions of cells or whole small cells
of the host algae that had entered the hyphae through openings. Tobler
(1925) summarized the results of his own work and that of others on the
biology of the lichens, confirming the belief that they consist of fungi
parasitizing upon algae with a mutual interaction of the two organisms
to produce the characteristic structures.

The algal hosts are usually Blue-green Algae (Myxophyceae) or Green
Algae (Chlorophyceae) whose habitat consists of moist situations on land,
such as on the ground, rocks, trees, etc. More often the one-celled algae
are preferred (Chroococcus, Chlorococcum, Protococcus, etc.), but some of
the filamentous forms are also captured and made prisoners {Nostoc,
Trentepohlia, etc.). Most genera of lichens are confined to a single species
or to closely related species of algae but a few lichens are able to use as
algal hosts forms from widely diverse groups. It is noteworthy that the
thalli formed by the same species of fungus with these widely separated
algal hosts are very different, according to the host present.

Lichens may be grouped morphologically in accordance with the type
of thallus into crustose, foliose, fruticose, and pendent lichens. The first
form closely adhering crusts on the substratum to which they are held
fast by hyphal strands; the foliose lichens are flat and thin ("leaf-like")
but adhere to the substratum only at definite points; the fruticose lichens
are upright in habit and more or less branched, resembling little shrubs
{frutex means shrub). The pendent forms are attached only at localized
spots and are long and slender and branching, hanging from the twigs
or branches of the trees to which they are attached. They are often con-
sidered only a special type of fruticose lichen. It must be noted that
these distinctions are not absolute for there are gradations between these
types. Most species of Cladonia are at first crustose or foliose (but with
small "leaves"), later forming upright podetia upon which the apothecia
are developed, at which time the prostrate foliar growth may disappear.
The distinction between crustose and foliose lichens is not always clear.
(Fig. 72.) _

In tropical and subtropical regions many lichens grow on the algae
attached to the surface of leaves. The fungus hyphae may then enter
through the stomatal openings into the interior of the leaf where it
seems probable that they are somewhat parasitic upon the leaf tissues.
McWhorter (1921) showed that some lichens of the genera Cladonia and

218

CLASS ASCOMYCETEAE

Fig. 72. Morphologic types of lichens. (A) Crustose type, Lecidea platycarpa Ach.
(B) Foliose type, Gyrophora muhlenbergii Ach. (C) Fruticose type, Stereocaulon
coralloides E. Fr., fruticose podetium ("secondary thallus"). (D) Pendent type,
Usnea harbata (L.) Wigg., portion of thallus with apothecia. (A, after Reinke: Jahrb.
^c^ss. Botan., 28:70-150. B-D, after Schneider: A Text-book of General Lichenology,
Binghamton, N.Y., Willard N. Clute & Co.)

Amphiloma growing in colonies of mosses (Musci) are capable of destroy-
ing the latter by direct parasitic attack. Rock-inhabiting lichens often
penetrate the rock itself for some distance by means of their holdfast
hyphae. Miss Mellor (1922) called attention to serious damage done to
glass windows in some old churches in France where lichens attached
themselves to the glass and gradually corroded it. Probably the requisite
mineral nutrients of the lichen and enclosed algae are obtained by the
hyphae that attach the lichen to its substratum. A very few species of
lichens develop in aquatic habitats.

A great many organic acids, the so-called lichen acids, have been
found in the lichens. They were given intensive study by Zopf (1907).
Following a suggestion of his, one of his students, F. Tobler (1909) con-
ducted experiments which demonstrated that this production of lichen

ORDER LECANORALES (tHE DISKJLICHENS) 219

acids is the result of mutual physiological interaction of alga and fungus,
as they do not occur in either organism Avhen groAvn alone. From some of
these acids may be obtained brilliant pigments such as orcein, litmus,
etc. (Miss Smith, 1926).

M any lichens have no known asexual mode of reproducing themselves.
The pendent forms are frequently torn to pieces by the wind and carried
considerable distances, thus achieving distribution. A good many Hchens
produce pycnidia containing conidia. For some species these conidia have
been germinated to produce a mycelium. On a great many lichens special
asexual reproductive structures, the soredia, are produced. In barest
details a soredium arises as an outgrowth of the interior mycelium of the
thallus, carrying with it some of the algal cells. Having grown out through
the surface of the thallus this mycelial mass rounds up into a ball with
a sort of cortex, containing in its interior loose hyphae and a few host
cells. This structure breaks loose and is distributed by wind or rain to
other locations where the hyphae grow fast, thus starting a new lichen
thallus.

In their sexual reproduction the Lecanorales are rather uniform in
their end product, the apothecium, which differs in detail but not in
fundamental plan in the various families and genera. These variations
ha\e to do with shape (concave, fiat, convex); color; structure of the
paraphyses; number, color, structure, and shape of the ascospores (color-
less or brown, one-celled or divided into two or more cells, ellipsoidal,
fusoid, needle-like, etc.); structure of the hypothecium and excipulum;
size; location on the thallus, etc. Two different structures called excipula
or exciples are of importance in systematic arrangement of the families
and genera of lichens. The "thalloid exciple" is a marginal wall around
the apothecium consisting of an upgrowth of the thalloid hyphae, often
with enclosed algal cells. It does not arise from the developing apothe-
cium. The "proper exciple" is the cup-like margin at the edge of the
apothecium which is formed by the outgrowth and upgrowth of the
apothecial tissues. Sometimes both types are present but more usually
only one type, and in many lichens neither type of exciple is observable.
In contrast with most of the Pezizales the apothecia of the Lecanorales
are usually slow in development and persist for a long time, maturing a
few asci at intervals. The asci are inoperculate and usually thickened at
the apex. The details of the sexual process, particularly the behavior of
the sexual nuclei, are sadly in need of further study in almost all genera
of the order. It may be safely said that no lichen has been satisfactorily
studied from all these standpoints. The conditions in Collema and Colle-
modes will illustrate the main features of the sexual process in this order.

In its interior Collema consists of a slender, branched mycelium,
loosely penetrating and limited in outline to the shape of the Nostoc

220 CLASS ASCOMYCETEAE

colony it inhabits. There is no cortical layer. On the interior mycelium
there arise here and there somewhat thicker hyphae which are noticeable
because of their dense contents and more or less loosely coiled structure.
These are the ascogonia. The cells like those of the vegetative mycelium
are uninucleate. Each ascogonium consists of from one to three coils of
cells terminated by a filament (the trichogyne) which turns toward and
projects just through the surface of the colony. Its exposed tip is sHghtly
enlarged and covered with a somewhat thickened wall which is sticky
when Avet. In its multicellular structure it differs greatly from the tricho-
gyne of the Florideae. The male organs are branched hyphae projecting
into a conceptacle-hke spermogonium which opens at the surface of the
colony. From these branches there bud off minute, uninucleate, non-
motile sperms possessing a delicate cell wall. When wet by rain the gummy
mass filling the spermogonium swells and oozes from the opening, where
the gum dissolves and the sperm cells are floated off by the film of rain
water. Such a sperm coming in contact with the sticky tip of the tricho-
gyne adheres to it. Stahl (1877) and Baur (1899) have demonstrated that
an opening is dissolved quickly through which the sperm nucleus enters
into the apical trichogyne cell. Successive swellings and disappearance
of the septa of the trichogyne seem to indicate the passage of the sperm
nucleus down to the coils of the ascogonium. The nuclear behavior has
not been followed in detail however. From one of the ascogonial cells,
which therefore corresponds in function to the oogone of Pyronema, asco-
genous hyphae begin to grow outward and upward. The surrounding
vegetative hyphae also become actively involved in growth and produce
the vegetative part, including the paraphyses, of the apothecium. The
ascogenous hyphae produce their asci by the hook method as described
for Pyronema. It should be noted that in some genera of lichens the sperm
cells, or cells resembling them and produced in similar conceptacles, are
capable of growing in pure culture in nutrient media until normal thalli
are produced bearing similar conceptacles. This was reported by Moller
in 1887. He obtained thalli by culturing such cells from Buellia puncti-
formis Hoffm., Opegrapha suhsiderella Nyl., 0. atra Pers., Arthronia sp.,
Calicium parietinum Ach., and other species of Calicium. He also cul-
tured the conidia from the pycnidia which in some cases are present in
the same thallus. He therefore drew the conclusion that the spermogonia
as well as pycnidia were both asexual reproductive structures and that
the supposed sexual function of the spermatia was erroneous. (Fig. 73A,
B, D-G.)

Miss Bachmann (1912) erroneously identified the very similar lichen
Collemodes hachmannianum Fink with Collema pulposum (Bernh.) Ach.
She found that in Collemodes the branches which produce the sperm cells
are not produced together in spermogonia but are scattered here and

ORDER LECANORALES (tHE DISK LICHENS)

221

Fig. 73. Lecanorales, Family Collemaceae. (A, B) Collema crispum Ach. (A) Asco-
gonium and trichogyne. (B) Apical cell of trichogyne, showing sticky surface. (C)
CoUemodes hachmannianum Fink. Ascogonium and trichogyne growing to clusters of
sperm cells produced within the thallus. (D-G) Collema pulposum (Bernh.) Ach.
(D) Habit sketch. (E) Cluster of apothecia. (F) Section through apothecium. (G)
Ascus. (A-B, after Baur: Ber. dent, botan. Ges., 16(10): 363-367. C, after Bachmann:
Ann. Botany, 26(103) :747-760. D-G, after Schneider: A Text-book of General
Lichenology, Binghamton, N.Y., Willard N. Clute & Co.)

222 CLASS ASCOMYCETEAE

there in the interior of the colony. The ascogonia are similar to those in
Collema but the trichogynes are longer and do not extend to the surface.
They are attracted, apparently chemotropically, to the clusters of sperm
cells and grow toward them, coiling around and uniting with them. The
subsequent development is identical with that in Collema. (Fig. 73C.)

In a number of other genera (e.g., Physcia) spermogonia and tricho-
gynes are produced. The former are various in shape, spherical or de-
pressed globose or lobed and immersed (except for the ostiole) or partly
or almost completely emergent. The sperm cells (spermatia) are minute,
one-celled, and usually slender, rarely rounded. They are produced
apparently successively at the apices of unbranched antheridial hyphae
or on the upper portion of the cells of the multicellular simple or branched
hyphae. They were studied in great detail by Lindsay (1861, 1872) in
many genera of lichens from all parts of the world. Adherence of sperms
to trichogynes has been observed but rarely. In a few cases an opening
has been observed between sperm and trichogyne but nuclear passage
has not been seen. Probably the sperm is functional in most cases of this
sort. In those species in which ascogonia have been reported but no
spermogonia are known the structure reported above for Collemodes
should be sought for before denying any type of sexual union. In many
lichens the ascogonium produces no trichogyne and may even be but a
straight row of a few cells. In one or two such cases adjacent cells of the
ascogonium lose their intervening septa whereupon ascogenous hyphae
begin to appear. Just what the nuclei do in that case can only be sur-
mised. Sexuality seems to be on the decline in this order as throughout the
Higher Fungi. The Moreaus (1926, 1928) have studied the reproduction
of many lichens and deny any sexual function to the spermatia, consider-
ing them when present to be modified conidia. (Fig. 74.)

Because of the possession of a functional trichogyne and the produc-
tion of separate sperm cells and of asci a certain degree of relationship
between the Lecanorales and Laboulbeniales can be postulated, but they
are certainly widely divergent from any common ancestor. This may
have been an alga somewhat like some of the filamentous, freshwater
Florideae. It would require the assumption that subsequent to the acqui-
sition of the ascus-producing habit the one series developed as parasites
on insects with little modification of the protected procarp while the other
series developed as parasites on algae, presumably at first submerged
forms, later land algae. At the same time the spore fruit deviated far
from the simpler procarp typo shown in the Laboulbeniales. Here again
it must be noted that many mycologists hold that the sperm cells are
nothing but modified conidia which have taken up secondarily the sexual
function in place of an antherid. The external similarity between these
groups and the Flcjrideae would be looked upon from this viewpoint as a
case of convergence, not as an indication of true phylogenetic relationship.

ORDER LECANORALES (tHE DISK LICHENS)

223

Fig. 74. Lecanorales, Family Physciaceae. Physcia sp. (A) Spermogonium. (B) Anther-
idial filament with sperm cells. (After Kny: Botanische Wandtafeln.)

Zahlbruckner (1926) believed that the group treated here as a single
order, the Lecanorales, is really polyphyletic, i.e., derived from nonHchen
forming fungi at many different points. Nannfeldt (1932) discussed this
problem and showed that some of the perithecial lichens (Order Py-
renulales) are in reality more closely related to the Order Pseudosphae-
riales and that of the lichens with true apothecia many have their closest
relationship not with other lichens but with Pezizales that are not lichen
producing. Perhaps the ultimate logical disposition of the disk hchens
will be to distribute them among the other apothecium-forming fungi
at the points where their apothecial structure and manner of sexual repro-
duction seem to fit best. Until the enormous mass of study needed to
acquire this information has been carried out it may be best to treat them
as a single order.

The forms here included in the Lecanorales are divided by Zahlbruck-
ner into about 37 families, about 275 genera, and over 7400 species. They
occur from the tropics to the Antarctic and Arctic zones and from sea
level to the tops of the highest mountains where rock is exposed. They
furnish the chief food of the caribou, the reindeer, and the musk ox. Some
species are used for human food in famine times. Iceland moss, Cetraria
islandica (L.) Ach., is sometimes used for medicine. Litmus and orcein
are derived from Hchens. Miss Annie L. Smith's (1921, 1926) publications
on these fungi should be studied for a more complete understanding of
this extremely variable group of organisms.

224 CLASS ASCOMYCETEAE

Order Pezizales. The fungi comprising this order are mostly sapro-
phytes although a number of the more or less serious diseases of culti-
vated plants are caused by parasitic species (e.g., various species of
Sclerotinia, Pseudopeziza, etc.). Those forms that are parasitic do not
attack algae in such a manner as to produce lichen thalli. Asexual repro-
duction by means of conidia is found in a good many species but is by
no means as widely distributed as in some of the orders to be discussed
later. Conidia are formed singly on simple or branched conidiophores
or the individual cells of whole segments of mycelium may round up to
form chains of conidia. In a number of species sclerotia are produced in
abundance.

In this order we find almost all gradations in sexual reproduction. In
Stromatinia gladioli (Drayton) Whetzel, it was shown by Drayton (1932,
1934) that minute sperm cells must be brought to certain receptive organs
of the ascogonia before apothecia can be produced. In Ascobolus car-
bonarius Karst., B. O. Dodge (1912) showed that a much coiled asco-
gonium bears a long trichogyne which grows toward a structure resem-
bling a conidium and attaches itself and fuses with it. This resembles
greatly what Miss Bachmann described for Collemodes. In the genus
Ascobolus other species have an ascogonium which coils directly around
and fuses with an upright antherid (Dodge, 1920). In still other species
both Schweizer (1923) and Ramlow (1914) have indicated that antherid
and trichogyne are both absent. In Scutellinia stercorea (Fr.) Kunze,
according to Miss Fraser (1907), the oogone is rounded and multinu-
cleate, as in Pyronema, but the trichogyne is several-celled. S. G. Jones
(1930) found in Pseudopeziza trifolii (Biv.-Bernh.) Fckl. the production
within the leaf of the host (Trifolium pratense L.) of numerous ascogonial
coils made of heavily staining uninucleate cells. Around these but without
any visible cell fusion the vegetative mycelium develops into actively
growing hyphae, some of which emerge from the stomata. Jones called
these "trichogynes" but denied any reproductive function, considering
them to be "respiratory hyphae." Other hyphae become true ascogenous
hyphae with binucleate cells and still others produce the paraphyses and
other portions of the apothecium while the original ascogonium degen-
erates without having served any other function than as a center of
attraction for the surrounding hyphae. There is a nuclear fusion in each
young ascus and only the first of the following three nuclear divisions is
reduf'tional. In some Pezizales even the oogone or ascogonium is not to
be found. The eventual product is an apothecium which in its general
plan is like that of the Lecanorales but usually larger and more fleshy.
(Fig. 75.)

In this order two series of forms may be distinguished, depending upon
the mode of dehiscence of the ascus at maturity, viz., the Operculatae

ORDER PEZIZALES

225

Fig. 75. Pezizales, Family Pezizaceae. (A, B) Ascobolus carbonarius Karst. (A)
Ascogonium with trichogyne reaching out to a distant antherid. (B) Ascogenous
hyphae beginning to bud out from ascogonial cells. (C) Ascobolus magnijicus Dodge,
ascogonium coiling around antherid. (A-B, after Dodge: Bull. Torrey Botan. Club,
39(4):139-197. C, after Dodge, Mycologia, 12(3):115-134.)

and the Inoperculatae. In the former a little lid (operculum) is formed at
the apex of each ascus. This gives way when the turgor pressure reaches
a certain degree, thus allowing the escape of the ascospores and the sur-
rounding liquid. The operculum may be shot off entirely but more often
remains attached at one edge like a trap door. A modification of the
typical operculate type is apparently the bilabiate type. In the Inoper-
culatae the thickened apex of the ascus gradually softens and suddenly
yields to the internal pressure forming a pore through Avhich the ascus
contents escape.

In the Pezizales the ascospores are one-celled and ellipsoidal to sub-
spherical to a much greater extent than in the Lecanorales where many-
celled ascospores are common. The apothecia in this order vary greatly
in size. In a few species of Ascobolus and some other genera the apo-
thecium is less than a millimeter in diameter; in the larger number of
genera and species it is from 5 to 20 mm. in diameter. As mentioned in
the preceding chapter a specimen of Geopyxis cacahus (Fr.) Sacc. was
collected in Java that was nearly a meter high and about 50 cm. across.
Seaver (1942) reported that Dr. Helen M. Gilkey and Dr. S. M. Zeller
found a specimen of Daleomyces phillipsii (Massee) Seaver in Oregon
that had a diameter of 40 in. (about 1 meter). The shape is also subject
to great variation. In Pyronema and some other genera it is convex and
naked from the beginning. In more forms it is flat or cup-shaped and in
most of them the hymenial surface is at first covered with a more or less
evanescent layer. Some species have a subspherical apothecium, at first
closed and then opening at the apex by an enlarging pore or ostiole. The
apothecium is more often sessile but yet is stalked in many genera. A few
genera produce their apothecia just under the surface of the soil, opening
by a small pore at the surface when the apothecium is mature. In Cyttaria,

226 CLASS ASCOMYCETEAE

parasitic on twigs of the Southern Beech (Nothofagus), there is formed a
fleshy stroma several centimeters in diameter in whose outer half or more
the small apothecia arise. These are at first closed but at maturity open
at the surface of the stroma.

A noticeable feature of the larger apothecia of the whole order is the
simultaneous discharge of ascospores over a large portion of the hy-
menium. This is visible as a cloud, like smoke or steam. This discharge is
often accompanied by a hissing sound, as has been verified by the author.
The distance to which the spores may be discharged is remarkable, some-
times several centimeters. Falck (1916, 1923) has shown that the dis-
charge of ascospores is dependent to a considerable degree upon changes
in temperature or illumination or upon contact of some other object with
the apothecia. Even the stimulus of a gentle current of air is sufficient
to cause spore discharge in some species. Buller (1934) gave an extended
discussion of the conditions that induce this simultaneous discharge of
spores and that affect its direction. In species of Ascobolus and some other
genera the ascus at maturity becomes greatly elongated and distended
laterally by the absorption of a large amount of water. When the oper-
culum gives way the greatly enlarged ascus contracts with much violence
and the contained liquid and ascospores are ejected to an amazing dis-
tance. In Saccobolus the ascospores are massed together in a ball which
is expelled further than would be possible for separate spores.

It has been shown that differentiation into two distinct sexual strains
occurs among some species of this order. Thus Miss Green (1931) showed
for Ascobolus furfuraceus Fr. and Betts (1926) for A. carbonarius Karst.
that they will not produce apothecia when grown in culture from a single
ascospore but require the meeting of mj^'celia developed from different
ascospores and then not from any two but from two of opposite sexual
strains. On the other hand some species of Ascobolus are fertile when
grown from but a single ascospore. This is a phase of investigation that
has attracted the attention of students but much still remains to be
learned. Drayton demonstrated that in Stromatinia gladioli there are
formed on the mycelium arising from one ascospore minute cells (micro-
conidia or sperms) and certain receptive structures within which are
developed ascogonia with long trichogynes. Such a mycelium remains
without producing apothecia. The mycefia produced by the eight asco-
spores of the ascus represent two phases, four of each. The sperms of any
mycelium of one phase can fertilize the receptive bodies of any mycelium
of the other phase and vice versa. Apparently this is not true hetero-
thallism or condition of malencss and femaleness of the different strains
such as occurs among the Mucorales. It is comparable to the self-sterility
of many flowering plants to their own pollen. For example the Bartlett
pear pistil rarely develops to a fruit when pollenized by pollen from the

ORDER PEZIZALES: SUBORDER OPERCULATAE 227

same variety and the same is true for the Kieffer pear, but these two
varieties are usually fertile to each other's pollen. It is perhaps more
comparable still to the dimorphic species of Primula studied by Darwin
(1889), in which the seeds of a capsule will produce about equal numbers
of the two types of primrose plants, those with flowers possessing a long
style and low-placed stamens and those whose flowers have short styles
and stamens high in the corolla tube. Each strain is relatively sterile with
pollen from plants of its own type but fertile with pollen from plants of
the other type. Whether the condition in Ascoholus magnificus Dodge is
like the foregoing, i.e., a case of self-sterility, or is true heterothallism
(a real difference in sex) remains to be discovered by further study.

The " Discomycetes " were classified by the earlier investigators
Persoon (1801) and Fries (1822), largely on the basis of external charac-
ters. Later the ascus and ascospore characters were also taken into con-
sideration. The internal structure of the apothecium proved to be of great
importance. Durand (1900) used this as a basis for a tentative classifi-
cation. Nannfeldt (1932) has used these features in his extensive writings
on this group. Boudier (1907) pointed out that the mode of dehiscence of
the ascus, whether by a lid or by a pore, i.e., operculate or inoperculate, is
of great diagnostic value. Seaver (1928) in his volume on the Operculate
Cup-fungi recognized only two families in this suborder in place of a
larger number recognized by Schroeter and Lindau (1896) in Engler and
Prantl.

Order Pezizales : Suborder Operculatae. Family Pezizaceae. Apo-
thecia flat, convex, or concave or cup-like, sessile, or short stalked, rarely
long stalked, and then the hymenium concave or at most flat. Apothecium
pseudoparenchymatous throughout, with few exceptions. Typical repre-
sentative genera in this family are: Ascoholus, growing on animal excre-
ment or on soil, with mature asci much protruding and with mature
spores violet in color. The apothecia vary, according to species and
environment, from less than 1 mm. up to nearly 3 cm. in diameter. In
A. immersus Fr. the ascospores may attain a size of 50 to 75 ju in length
by 20 to 35 /i in thickness, almost the largest ascospores known. Pyronema,
grows on soil, especially after a fire or after steaming. The apothecia are
1 to 2 mm. in diameter and the convex hymenium is practically naked
from the first. The ascospores are hyaline. Humarina {Humaria Sacc.)
is less than 1 mm. to 1 cm. in diameter, growing on the ground, forming
white or bright-colored, mostly disk-shaped apothecia, with hyaline
ascospores, differing from Ascophanus only in that the latter grows on
dung. Patella forms disk-shaped apothecia up to 1 cm. wide, and with
the outside clothed with hairs, at least at the edge. P. scutellata (L.)
Morgan {Lachnea scutellata (L.) Gill.) forms its brilliant red disks with
a fringe of dark hairs, on rotten wood and is strikingly beautiful. Plectania

228

CLASS ASCOMYCETEAE

Fig. 76. Pezizales, Family Pezizaceae. Peziza repanda (Pers.) Fr. (Courtesy, F. C.

Strong.)

has stipitate apothecia growing on sticks lying on or buried in the ground.
Its hymenial surface is brilliantly colored. P. coccinea (Scop.) Fckl. is
very abundant in some regions in early spring forming scarlet apothecia
sometimes 3 or more cm. in diameter. The stipe is buried in the soil.
Peziza has saucer- to cup-like apothecia, usually of considerable size,
2 cm. up to 30 or 40 cm. P. hadia Fr. and P. repanda (Pers.) Fr. are quite
frequent in woods and P. vesiculosa Fr. in greenhouse soil that has been
heavily manured. Geopyxis is quite similar to Peziza except that the hemi-
spherical or acorn-cup-like apothecium is supported by a relatively slender
stalk. (Fig. 76.)

Family Helvellaceae. Apothecia stalked, convex, attached at the
apex of the stalk or grown fast to its upper portion. To be mentioned are
Helvella, with the apothecium more or less saddle-shaped, attached by the
center of the under side of the saddle. The hymenial surface may be
smooth or gyrosely folded. In the latter case a separate genus Gyromitra
is often recognized. Morchella, with the apothecium grown fast down
the side of the upper part of the stalk and with its surface thrown into
strong longitudinal and transverse folds so as to be coarsely pitted, thus
greatly increases the hymenial surface. The species of this genus are
known as Morels or Sponge Mushrooms and are among the most de-
licious edible fungi known. They grow mostly in deciduous woods, fruiting
in the spring. Verpa, the Bell Morel, has the cap shaped like a bell and
free from the stalk except where attached at the top. It may be smooth
or longitudinally ribbed. It is edible, coming a little earlier than the true
morels. (Figs. 77, 78.)

^^ ^Ip^

1 JH^

A. "

■ f^^m ^«^

■ ' f ^

■

i^iigi

Fig. 77. Pezizales, Family Helvellaceae. Gyromiira escidenUi (Pers.) Fr. (Courtesy,

F. C. Strong.)

-4^#V'' "h

Fig. 78. Pezizales, Family Helvellaceae. Morchella conica Pers. (Courtesy, F. C.

Strong.)
229

230 CLASS ASCOMYCETEAE

Order Pezizales : Suborder Inoperculatae. This suborder differs from
the Operculatae in the absence of an operculum. The apex of the ascus
is usually thicker and sometimes depressed. When the spores are mature
the ascus elongates and often becomes thicker by the increase in turgor
due to the absorption of water. The apical portion of the ascus softens
and thickens and finally gives way suddenly, permitting the violent
expulsion of the contained liquid and ascospores as the distended ascus
wall contracts. The apothecia vary greatly in size, structure, and con-
sistency. Nannfeldt (1932) divided the many genera and species into
three orders: Lecanorales (already discussed above), Ostropales, and
Helotiales. Because the life histories and inner apothecial structures are
llioroughly known in so few of the described species it seems best to the
author not to recognize the latter two orders of Nannfeldt until detailed
structural and developmental studies in the Lecanorales and in the
operculate and inoperculate Pezizales shall enable mycologists to set up
a more scientific classification of the whole group of Discomycetes.

Family Ostropaceae. The members of this family are characterized
by the narrow elongated asci with thickened apex through which runs
a slender canal almost to the surface, and by the long, thread-like asco-
spores which are septate at frequent intervals and which break up at
maturity into cylindrical pieces. The excipulum is mainly pseudoparen-
chymatous. The apothecium may be stalked (Vibrissea), superficial
without stalk {Apostemidium) , sunk in the substratum and disk-shaped
(Stictis), or rarely perithecium-like, with an ostiole (Ostropa). Among the
families which would make up Nannfeldt's order Helotiales only a few
are mentioned:

Family Dermateaceae (Including Mollisiaceae). Apothecia small
or medium sized, mostly epiphytic on woody or herbaceous plants, some-
times on the ground, parasitic or saprophytic, usually fleshy but some-
times cartilaginous or leathery, mostly not bright-colored. Excipulum
usually pseudoparenchymatous and dark-colored. The apothecia are
often formed within the host tissue, breaking out and opening at matur-
ity. Mollisia, however, produces its apothecium externally on the host
tissues. It has one-celled rather elongated ascospores. Pseudopeziza pro-
duces its apothecia out of a well-developed stroma under the epidermis
of the parasitized leaf which is ruptured at the maturity of the apo-
thecium. The ascospores are one-celled. Ps. medicaginis (Lib.) Sacc. is
sometimes the cause of yellowing of the foliage of alfalfa or lucerne
{Medicago saliva L.) and its premature leaf fall. Drepanopcziza rihis
(Kleb.) V. Hohn. {Ps. rihis Kleb.) produces its apothecia on dead leaves
of species of Ribes. The actively parasitic stage of this fungus produces
only the conidial type of reproduction formerly known as Gloeosporium
rihis (Lib.) Mont, and Desm. Diplocarpon likewise produces its apothecia

ORDER PEZIZALES: SUBORDER INOPERCULATAE

231

Fig. 79. Pezizales, Family Mollisiaceae. Diplocarpon earlianum (E. & E.) Wolf.
(A) Acervulus of Marssonina stage. (B) Section through apothecium. (Courtesy,
Wolf: /. Elisha Mitchell Sci. Soc, 39(3-4) :141-163.)

upon dead leaves, sometimes under a superficial radially arranged shield-
like stroma, sometimes not. The parasitic stage produces the conidial
form known as Adinonema when the radiating dark hyphae are present,
or Marssonina in their absence, or Entomosporium when the conidia have
hair-like appendages. On various species of Prunus there occur several
species of fungi whose conidial stage forms the genus Cylindrosporium.
It is this stage which is parasitic and causes great damage to the leaves
of plums and cherries producing the disease called "yellows" or "shot-
hole." The elongated conidia are formed subepidermally in an acervulus.
Later in the summer in the same or other acervuli small almost spherical
cells are formed, frequently called microconidia. In the stroma developing
below the acervulus Higgins (1914a) and Backus (1934) observed the
formation of numerous elongated coiled ascogonia which extend up to
the microconidial layer. Backus showed that these microconidia grew
fast to the terminal cells of the ascogonium, i.e., to the trichogyne, and
should therefore be considered as sperm cells. Subsequently ascogenous
hyphae are produced and the apothecium develops. Higgins identified
this fungus with the genus Coccomyces but Nannfeldt indicated that
this is incorrect. He therefore gave to these forms the name Higginsia.
Unfortunately this name is preoccupied and until a valid name is pro-
posed the name Coccomyces will probably continue to be used. (Fig. 79.)
Family Helotiaceae. Apothecia mostly fleshy, disk- or cup-shaped,
at first closed, often stalked, the excipulum consisting of filamentous
hyphae, sometimes grading into an outer layer of shorter, thicker cells.
Mostly saprophytic or parasitic upon plant tissues. Apothecia not origi-
nating in sclerotia. Conidial stages of reproduction usually not present
or at least rarely conspicuous. Hdotium produces sessile or almost sessile,
disk- or cup-shaped apothecia, small to several millimeters in diameter, on
plant parts, probably mostly as saprophytes. The hymenium is often

232

CLASS ASCOMYCETEAE

bright-colored. The ascospores are three- to four-celled. CMorocihoria
aeruginosa (Oed.) Seaver occurs on wood and stains it green. Tricho-
scyphella {Dasyscypha of some authors) forms mostly short-stalked apo-
thecia on the bark or cones of conifers. T. willkommii (Hart.) Nannf. is
a serious canker-producing parasite of Larix.

Family Sclerotiniaceae. Mostly parasitic but capable of prolonged
growth as saprophytes. Apothecia fleshy, stipitate, mostly cupulate,
funnel-formed or saucer-shaped, in one genus resembling Verpa; usually
some shade of brown ; arising from a definite sclerotium or from a stroma-
tized portion of the substratum. Asci inoperculate, mostly eight-spored,
ascospores ellipsoidal, often flattened on one side, usually hyaline, uni-
cellular and smooth. Spermatia globose to slightly oval, conidial forms
various, in most genera lacking (Whetzel, 1945). As in the preceding
family the main portion of the excipulum consists of entangled hyphae,
only at the outer surface being reduced to short, somewhat pseudo-
parenchymatous cells. The stroma may be a free tuberoid sclerotium
as in the genus Sclerotinia or may be formed in the tissues of the host
which are digested and replaced by the fungus hyphae, e.g., Cihorinia,
Ciboria, Monilinia, or may be plano-convex and attached to the host,
Botryotinia, Septotinia, etc., or may be indeterminate or of the substratal
type, Lamhertella, Rutstroemia, etc. In all, Whetzel recognized fifteen
genera including over ninety species. Many of them were formerly in-
cluded in the genus Sclerotinia. Among the serious pathogens may be
mentioned Sclerotinia sclerotiorum (Lib.) de Bary, Monilinia fructicola
(Winter) Honey {Sclerotinia fructicola Rehm) , the cause of the brown rot
of stone fruits in America. Its conidial stage is of the Monilia type.
Botryotinia fuckeliana (de Bary) Whetzel is the cause of a serious disease
of the grape vine in Europe. Its conidial stage is a Botrytis of the B.

Fig. 80. Pezizales, Family Sclerotiniaceae. Scleroiinin sclerotiorum (Lib.) de Bary.
(A) Apothecia growing from sclerotium. (B) Magnified section of sclerotium. (After
Brefeld: Unter. Geaammt. Mykol, 4(7):112-121.)

ORDER PEZIZALES: SUBORDER INOPERCULATAE

233

cinerea type. In Stromatinia gladioli (Drayton) Whetzel, F. L. Drayton
(1932, 1934) demonstrated that the fungus has two sexual strains, the
sperm cells of one strain fertilizing the receptive female organs of the
other strain and vice versa. The conidial stage of this genus also belongs
to the genus Botrytis. It is probable that all the genera of this family in
which "microconidia" are known are of this type of sexual reproduction.
(Fig. 80.)

Fig. 81. Pezizales, Fam-
ily Geoglossaceae. (A) Geo-
glossum glabrum Pers. ex
Fr. (B) Leotia chlorocephala
Schw. (Courtesy, Durand:
Ann. MycoL, 6(5):387-
477.)

Family Geoglossaceae. This family was formerly associated with
the Helvellaceae to form the order Helvellales, before the importance of
the type of dehiscence of the asci became apparent. The apothecium is
stipitate and the hymenium either forms a closely adherent layer to the
somewhat thickened upper part of the stipe or forms the upper surface
of a head borne at the apex of the stipe. The spore fruit may be entirely
made up of hyphae or the cortex may be pseudoparenchymatous. In spite
of the external position of the mature hymenium it is really formed endog-
enously, being covered when young as in most genera of the Pezizales

234 CLASS ASCOMYCETEAE

with a convex veil. The ascospores are elhpsoid and one- or two-celled
(Mitrula) or long elliptical to filiform and several to many septate. They
are hyaline to smoky to dark brown in color. The fruiting bodies vary
from a few millimeters to five or more centimeters in height. They are
mostly found on rotten wood, decaying leaves, moss or soil, usually where
plenty of moisture is available. Several genera have bright-colored spore
fruits (e.g., Mitrula, Spathularia, etc.) while those of others are black
(e.g., Geoglossum) . The following genera are worth mentioning: Mitrula,
spore fruit clavate, bright-colored, ascospores hyaline, ellipsoid; Micro-
glossum, similar but with ascospores elongated and many septate; Geo-
glossum, clavate, black, smooth, and dry, ascospores dark, many septate;
Gloeoglossum, similar but viscid and gelatinous; Trichoglossum, similar to
Geoglossum but beset with spines or setae; Spathularia, fan-shaped,
bright-colored, ascospores hyaline, many septate ; Leotia, spore fruit capi-
tate, gelatinous, spores narrowly ellipsoid; Cudonia, capitate, leathery,
ascospores filiform, multiseptate. Eleven genera and about forty-one
species were recognized by Durand (1908) in his excellent monograph of
the family. Nannfeldt (1932) believed that this family is related to the
stipitate Helotiaceae. (Fig. 81.)

Family Phacidiaceae. The apothecia of this family arise in a well-
developed stroma which encloses it below and above, and which is often
lenticular in vertical section. These stromata may be superficial or buried
in the tissues of the host plant. Possibly as a result of the protection
afforded by the stromatic envelope the excipulum is not strongly devel-
oped. The stromata may be rounded or elongated. In the former case
the stromatic cover often splits stellately at maturity to reveal the
apothecium, while in the elongated forms a longitudinal slit is formed.
There may be but one apothecium in each stroma or several. In the
latter case each apothecium may be elongated more or less, even when
the stroma is isodiametric. The asci are clavate, with hyaline, filamentous
paraphyses. The ascospores are shot off as in the Pezizaceae when the
mature hymenium is exposed by the opening of the stromatic cover.
They are elongated and sometimes needle-shaped, one- to many-celled,
hyaline or colored. They often have a gummy outer layer. Asexual repro-
duction is known for many species. These conidial forms usually belong
to the form family Leptostromataceae of the Fungi Imperfecti. Some
species are saprophytic but perhaps the majority are parasitic upon leaves
or twigs. The genus Hypoderma is sometimes placed in a separate family,
the Hypodermataceae, but the author follows Nannfeldt in uniting the
two families. Some authors unite these with other families to form the
order Phacidiales.

Phacidium possesses a circular, stellately dehiscing stroma with a
single apothecium. Some species are found on the needles of conifers.

ORDER PEZIZALES: SUBORDER INOPERCULATAE

235

Fig. 82. Pezizales, Family Phacidiaceae. Rhytisma acerinum (Pers.) Fr. (Courtesy,

F. C. Strong.)

Several species of Lophodermium also attack the needles of conifers, caus-
ing serious leaf fall. In this genus the stroma contains only a single
apothecium, but both stroma and apothecium are elongated and narrow,
and the dehiscence is by means of an elongated slit. Rhytisma, the cause
of the tar spot of leaves of maple (Acer) and other plants, produces a
large more or less isodiametric subcuticular stroma on the upper side
of the leaf and usually a smaller sterile stroma on the lower side. After
leaf fall the apothecia begin to develop slowly but do not reach maturity
until the folloAving spring. In a single stroma are produced numerous
elongated apothecia. These do not lie strictly parallel but are more or
less sinuately curved or sometimes radiately arranged. At maturity the
stroma forms a slit over each apothecium, under proper moisture con-
ditions pulhng back at the sides so that the hymenium is fully exposed.
At least three species occur on various species of Acer, each showing a
narrow specialization to only one host or to a group of host species. The
Red Maple {Acer ruhrum L.) is very subject to the disease in some parts
of North America. S. G. Jones (1925) has shown that in the stroma there
arise ascogonia, with at first one or two cells, which become three- to
five-celled. The cross walls become perforated and almost completely
absorbed and the nuclei pass into one of the central cells Avhich we must
conclude is the oogone. From this arise the ascogenous hyphae with

236

CLASS ASCOMYCETEAE

numerous pairs of nuclei. No nuclear fusion occurs until in the young
asci which are formed by the hook method. (Fig. 82.)

Family Cyttariaceae. The position of this family is not certain,
though it probably should be included in the Pezizales. It is not closely
related to any of the foregoing famihes. Apothecia numerous, imbedded
in a fleshy stroma produced externally on the twig of the host. Ascospores
one-celled, hyaline. One genus, Cyttaria, confined with its hosts (species
of Nothofagus, the Southern Beech) to the South Temperate Zone in
South America and Australasia. The fleshy stromata serve the natives
for food. The basal portion of the stroma of one or more species produces
organs resembling spermogonia wdth sperm cells. (Fig. 83.)

Fig. 83. Pezizales, Family Cyttari-
aceae. Cyttaria gunnii Berk. (After Lindau,
in Engler and Prantl: Die Natlirlichen
Pflanzenfamilien, Leipzig, W. Engelmann.)

Two genera of inoperculate Discomycetes from Sumatra described
by Boedijn (1934) perhaps indicate a transition to the Cyttariaceae. They
are J acobsonia on wood and Myriodiscus on bamboo stems. From a
plectenchymatic stroma there radiate in all directions closely packed,
branched stalks of apothecia forming a loose or dense ball respectively.
In Myriodiscus there are perhaps 800 to 1000 or more of these apothecia
which are rather gelatinous. The asci are cylindrical or obovoid and
multisporous. These spores are formed in this large number from the
beginning and are not the result of budding of a few original spores.
Judging from the illustration there must be several thousand of these
small ellipsoidal ascospores in a single ascus. In Jacohsonia, in which the
apothecial branches are not so tightly packed nor so numerous, the asci
are only eight-spored and the apothecia are subcoriaceous.

Order Tuberales. The fungi of this order are all terrestrial and pro-
duce subterranean spore fruits (ascocarps). Some are probably sapro-
phytic but it seems possible that certain species are perhaps parasites

ORDER TUBERALES 237

upon the roots of higher plants. The ascocarps vary in diameter from a
few milhmeters up to three or more centimeters and may be found close
to the surface of the soil or at a considerable depth. With no direct means
of bringing the ascospores into the air for distribution by air currents as
is the case in the Pezizales, the Tuberales depend for distribution largely
upon the activities of mycophagous animals, probably to a considerable
extent insects, but, for some species at least, ground-inhabiting rodents.
Thus in California the so-called ground squirrels dig out the fruiting
bodies, doubtless attracted by the odor diffusing up through the soil,
and eat them on the spot or carry them away to their burrows or other
hiding places. In these processes pieces of the ascocarps are scattered
and the ascospores find their way into the soil.

By germination of the ascospores of one species of Tuber a conidium-
bearing mycelium has been obtained but this has never been grown to the
stage where normal ascocarps were produced. Chaze and Mestas (1939)
made tissue cultures of Tuber melanosporum Vitt. on various culture
media and obtained in pure culture an extensive mycelium. Here and
there in this mycelium there developed dense dark-colored masses in
which were produced the two-spored asci and ascospores typical of the
species. However, typical fruiting bodies such as occur naturally in the
soil were not produced.

In 1863 Anton de Bary observed typical clamp connections, such as
occur in many Basidiomycetes, at the base of the asci in two species of
Tuber. De Ferry de la Bellone (1886) observed clamp connections on the
brown hyphae growing out of the ascocarps of Tuber brumale Vitt.,
T. mesentericum Vitt., T. aestivum Vitt., and T. panniferum Tul. and
figured this structure for the last species. Mattirolo (1887) found such
clamp connections in mycelium external to but connected with the spore
fruits of T. lapideum Matt. Greis (1936, 1938) showed that the ascus
hook of T. aestivum Vitt. has the structure of a clamp connection. If it
is confirmed that the external mycelium in this order bears clamp con-
nections it must be concluded that the fungus is dicaryon in nature, as
is the case with the secondary phase of mycelium in the Basidiomyceteae.

The sexual process is unknown in the Tuberales, the hypogeous habi-
tat of the fruiting body making it very unlikely that it would be observed
except by rare accident. The structure of the immature spore fruit has
been studied in a good many cases but for many species only the mature
ascocarps are known. By comparing the mature stages, taking cognizance
of the ontogeny of the ascocarp where it has been observed, Fischer (1896,
1938) and Bucholtz (1902) recognized a graded series leading from types
scarcely different from some of the Pezizales that have almost completely
subterranean apothecia to very complex forms such as those of Tuber in
which the apothecial nature of the ascocarp is almost entirely concealed.

238

CLASS ASCOMYCETEAE

Fig. 84. Tuberales, Family Tuberaceae. (A) Tuber uestivum Vitt., external \dew.
(B) Tuber rufum Pico, section through ascocarp. (C) Enlarged view of a portion of
the ascocarp, the ascospores with spiny surface. (D) T. ynagnatum Pico, ascus showing
ascospores with alveolate marking. (E) Genea hispidula Berk, vertical section through
ascocarp, showing the single cavity and opening. (After Tulasne, in Engler and
Prantl: Die Natlirlichen Pflanzenfamilien , Leipzig, W. Engelmann.)

Perhaps the simplest structure in the order, according to Miss Gilkey
(1939), is that exhibited by some species of Hydnocystis. The hypogeous
spore fruit is subspherical, with an opening which is usually partly blocked
by hairs. The single large cavity is lined by an even hymenium made up
of cylindrical, 8-spored asci. The paraphyses are of about the same length
as these and do not form an cpithecium above them. Except for its per-
manently hypogeous habit this might well be placed in the Pezizales, a
position, in fact, to which Fischer (1938) assigned it. In Genea the main
cavity may have even walls or may be thrown into folds which much
increase the hymenial surface. The paraphyses grow out beyond the asci

ORDER HYSTERIALES 239

and unite above the latter to form a thick pseudoparenchymatous epi-
thecium or "secondary cortex" which is generally not so thick as the
"primary cortex" (the excipulum and its outer layers). The folding of
the surface of the cavity may lead to the formation of canals leading to
enlarged chambers. In some of these the hymenium lines both but in
Piersonia and some other genera the asci arise only in the chambers while
the canals are lined only with rudimentary paraphyses. With this increas-
ing complexity the single external opening is lost and several openings
develop at points where the canals converge near the surface. In Tuber
the canals are filled at a very early stage by the ingrowing paraphyses
and appear as "veins" in the tissue of the sporocarp. The hy menial
chambers become obliterated by the ingrowing asci and paraphyses. In
this genus the asci are ovoid or spherical and often but few-spored. They
do not form a single hymenial layer at maturity but project into the
epithecial tissues at various levels. A number of other genera are recog-
nized with varying degrees of modification of the foregoing structural
types. Of all the species of the order a few species of the genus Tuher
(the truffle) are of economic importance. These species, especially T. ■
aestivum Vitt. and T. melanosporum Vitt., occur in the rather open forests
of Southern Europe (mainly under species of Quercus), where they are
collected by the use of trained animals (dogs or pigs) which find them
by their odor. In recent years many species of Tuberales have been dis-
covered in the Pacific Coast states where the climate is quite similar to
the regions of Europe where they are best known. Miss Gilkey (1939)
described and figured nineteen genera and fifty-seven species of North
American Tuberales. She included in the Tuberales the genera for-
merly placed by Fischer (1896) in the family Terfeziaceae in the Order
Aspergillales. (Fig. 84.)

Order Hysteriales. In this order of plant-inhabiting saprophytes and
parasites the apothecia are much reduced in size and compressed later-
ally, to elongated, often somewhat boat-shaped, structures, opening by
a long narrow slit. They are dark-colored, leathery or hard, and show a
strong contrast between the dark-colored excipulum and the light-colored
hymenial layer. The latter consists of ovoid to elongated cylindrical asci
intermingled with mostly septate paraphyses which are frequently en-
larged at the apex or branched, even forming at times a well-marked
epithecium. The apothecia may be single or in closely packed groups,
superficial or emerging from the substratum. The asci open by a pore,
not a lid. The ascospores are of very many different forms, as in the
Phacidiaceae, to which this order shows many points of relationship. They
are ellipsoid and one-celled, or several-celled, or divided both longi-
tudinally and transversely into many cells, or needle-shaped. In color
they vary from hyaline to brown. Conidial formation has been reported

240

CLASS ASCOMYCETEAE

Fig. 85. Hysteriales, Family Hysteriaceae. (A, B) Hysterographium minutuni
Lohman. (A) Apothecia and pycnidia. (B) Ascus and paraphysis. (C) Hysterium
insideiis Schw., asexual stage {Septonema spilomeum Berk.). (D, E) Lophium mijtil-
inum (Pers.) Fr. (D) Asexual stage, Papulospora mytilina (Pers.) Lohman. (E)
Pycnidial stage. (After Lohman: Papers Mich. Acad. Sci., 17:229-288.)

in a few species, usually with the conidiophores included in a pycnidium
or somewhat similar structure. Lohman (1933) has shown that some of
the conidial forms usually assigned to the genus Sporodesmium, of the
Fungi Imperfecti, are the conidial stages of several species of Hysteriales.
Besides this spore form other conidial stages observed by him represent
the genera Papulospora and Septonema of the Moniliales and various
forms of the Sphaei;opsidales. The details of sexual reproduction are
almost unknown in this group. The relationships within the order and
to other orders are still more or less problematic. Usually the more than
400 species are distributed among several families. Of these Nannfeldt
(1932) removes the genera assigned to the Hypodermataceae to the
Phacidiaceae, where they have been placed in this work. Some of the leaf
inhabiting fungi formerly ascribed to this order (e.g., the genus Parmu-
laria) are placed by the more recent students of the group in the Pseudo-
sphaeriales. This leaves as the only important family the Hysteriaceae.
Family Hysteriaceae. Apothecia external, black and carbonaceous,
single or united in a stroma. Most of the species of this family are sapro-
phytic on bark or decorticated twigs or branches. Hysterographium
fraxini (Pers. ex Fr.) de Not. is common on various species of ash (Frax-
inus). Its ascospores are dark-colored and divided by transverse and
longitudinal septa. The apothecia are boat-shaped. (Fig. 85.)

ORDER TAPHRINALES (eXOASCALES) 241

The relationship of this order is more or less in dispute. The compact
hymenium, with numerous well-developed paraphyses, and the slit-like
opening suggest great affinity to some of the Phacidiaceae, as does the
great variability of ascospore structure. The suggested relationship to
the family Lophiostomataceae in the Order Sphaeriales seems doubtful,
in spite of the slit-like ostiole, for in other respects the latter family is
typically Sphaeriaceous.

Order Taphrinales (Exoascales). This is a group of approximately 100
recognized species of fungi, all but a very few parasitic. Tw^o or more
genera are recognized in two families whose actual relationship to one
another is uncertain. Both families are characterized by the production
of a superficial hymenium with indeterminate margin and without para-
physes. This may rest upon a thin membranous hypothecium consisting
of interwoven hyphae (Ascocorticiaceae) or may burst through the epi-
dermis or cuticle of the host plant without a definite hypothecium
(Taphrinaceae). In the author's opinion these represent the ultimate
steps in the reduction and simplification of an apothecium. Together with
this simplification of apothecial structure there has arisen a marked
modification of the sexual process in the Taphrinaceae. According to
Miss Wieben (1927) the life history for Taphrina epiphylla Sadeb. and
T. klcbalmi Wieben is as follows: An ascospore, or one of the smaller
spores which the ascospores may produce by budding while still within
the ascus, germinates upon the surface of the host plant. Only when the
germ tubes from two such spores of opposite sexual phase meet and fuse
is active infection of the host possible. As a result of this conjugation a
dicaryon hypha is produced which grows intercellularly in the host tissues.
R. E. Fitzpatrick (1934) has shown that in T. deformans (Berk.) Tul. a
single spore can cause infection. The nucleus of this spore divides and
thenceforth the two nuclei divide conjugately and the vegetative my-
celium is of dicaryon nature as in the case of the species studied by
Miss Wieben.

The presence of these hyphae causes hypertrophy and hyperplasia in
the affected parts of the host. This may affect the leaves, fruits, and
shoots, often causing the formation of "witches' brooms." Eventually
dicaryon cells are formed near the surface of the host, often in a sub-
cuticular location, forming a structure resembling pavement epithehum.
These cells sometimes become somewhat thick-Avalled. In them eventually
jnuclear fusion occurs, followed by elongation of the cell in a vertical
lirection, rupturing the cuticle or emerging between epidermal cells of
bhe host. The diploid nucleus and most of the cytoplasm pass into the
[upper part of the cell leaving an empty lower part which is sometimes
[but not always cut off from the upper part by a cross wall, or the diploid
[nucleus may divide and one remain in the lower cell while the other passes

242

CLASS ASCOMYCETBAE

Fig. 86. Taphrinales, Family Taphrinaceae. Taphrina deformans (Berk.) Tul.
(A) Portion of a leaf section showing asci in various degrees of development. (B)
Mature ascus showing empty basal portion. (C) Portion of ascus showing two asco-
spores beginning to bud. (D) Subcuticular ascogenous cells seen from above. (E) In-
fection of leaf by dicaryon mycelium. (A-D, courtesy, Pierce: U.S. Dep. Agr. Vegetable
Physiol. Path. Bull., 20:1-204. E, courtesy, Fitzpatrick: Sci. Agr., 14(6):305-326.)

into the thin-walled upper portion undergoing further divisions there.
The diploid nucleus undergoes meiotic and mitotic divisions, the spindle
of the first division being transverse according to Juel (1921). Around
the eight nuclei thus formed the ascospores develop. In many species
each ascospore undergoes budding so that the ascus becomes polysporous.
Miss Wieben's germination experiments showed that four of the asco-
spores (and the spores produced from them by budding) are of one sexual
phase and four of the other. From Fitzpatrick's studies it is apparent
that in T. deformans this distinction does not occur or is not well marked.
Lohwag (1934) advances reasons for believing that Taphrina is not
a true Ascomycete but a Basidiomycete which has reverted to the ances-
tral condition. The normal basidium, in his viewpoint, represents an
ascus in which the ascospores have pushed out into projections of the
ascus wall so as to become apparently external, although really retained
within the wall. In Taphrina the transverse position of the nuclear
spindle, the habit of budding on the part of the ascospores, and the pari-

ORDER TAPHRINALES (eXOASCALES) 243

etal position assumed by the ascospore nuclei before the spores are
dehmited are all pointed to as Basidiomycetous characters.

Family Ascocorticiaceae. Ascocorticium, the only genus of the
Family Ascocorticiaceae, consists of a small number of species whose fruc-
tifications produce a gray- or pink-colored thin coating with indeterminate
growth, over the surface of bark or bare wood of dead trees. The hypo-
thecium consists of four to six layers of interwoven hyphae running par-
allel to the surface of the bark. Upon these arise the ovoid eight-spored
asci. The asci are formed by the hook or crozier method from dicaryon
ascogenous hyphae whose origin is not known. It is not known whether
there is any sexual process other than the probable union of tA\ o nuclei
in each young ascus.

Family Taphrinaceae. Ta-phrina {Exoascus, Magnusiella, or Taph-
ria) is the only genus of the Taphrinaceae. Of great economic importance
is T. deformans (Berk.) Tul., causing the disease of the peach {Amijgdalus
persica L.) known as leaf curl. The ascospores or spores arising from them
by budding lie dormant in crevices of the twigs or on the bud scales until
the following spring when they germinate and infect the young leaves
and even the fruits and young twigs. On the other hand the ascospores
may also germinate by budding on the surface of the host and grow in
the manner of a yeast, infection taking place in the spring from the yeast-
like cells. The affected leaves become much thickened and distorted as
well as discolored. The diseased fruits may show irregular bright red
patches of thickened tissues. On the diseased areas the asci are formed
subcuticularly in the late spring or early summer. In a few cases the
spores appear to germinate soon after formation so that Waite (1932)
and Poole (1932) report what may be a secondary infection occurring
the same season. Taphriria pruni Tul. and T. communis (Sadeb.) Giesenh.
infect the young fruits of Prunus and Padus, causing them to develop into
the hollow hypertrophied structures which, upon the plums, are known
as "plum pockets." Here also the asci are subcuticular. T. cerasi (Fckl.)
Sadeb. causes witches' brooms on various species of cherry {Prunus cerasus
L., P. avium L., etc.). The mycelium lives perennially in the tissues of
the host. T. robinsoniana Giesenh. causes hypertrophy of the scales of
the aments of the alder (Alnus). In T. potentillae (Farl.) Johans., the
mycelium lives beneath the epidermis and sends up, between the epi-
dermal cells, branches which bear separate asci. (Fig. 86.)

In this family the whole vegetative mycelium consists of dicaryon cells
and is therefore comparable to the ascogenous hyphae of those forms
with definite sexual organs and Avell-developed apothecia, such as Pyro-
nema, etc. Many authors consider this family to be primitive, but it seems
more logical to consider it as a much simplified offshoot of the Pezizales,
the Ascocorticiaceae being a possible intermediate stage. Miss Catherine

244 CLASS ASCOMTCETEAE

Roberts (1946) describes experiments and observations with TapJy^ina
deformans and Torulopsis pulcherrima (Lindner) Sacc, which demon-
strate many points of similarity in cultural behavior and in structure of
the organisms in culture. Both grow in culture in the manner of the
budding yeasts (Saccharomycetaceae). Torulopsis frequently produces
large thick-walled cells which bud out a small thin-walled two- to three-
spored ascus. A similar phenomenon was observed in the cultures of
Taphrina by Miss Roberts and also by Mix (1924, 1935). This would
seem to indicate the possibility that Torulopsis is a yeast-like derivative
of the Taphrinales. It is well know^n that very many widely unrelated
fungi may adopt the yeast manner of growth. It must be noted however
that Anton de Bary (1884) believed the yeasts and the Taphrinales to
be primitive, closely related Ascomycetes, a viewpoint not shared by the
author of this textbook.

Key to the More Important Families of Order Laboulbeniales

{Modified from Thaxter, 1908)

Antherids producing naked sperms endogenously.

Antherids opening by separate tubes. Family Laboulbeniaceae

12 or more subfamilies. Characteristic genera Laboulbenia, Stigmata my ces, etc.
Antherids compound, i.e., opening into a common chamber which has one
opening to the outside. Family Peyritschiellaceae

20 or more genera. Peyritschiella, Dimorphomyces etc.
Antherids, more or less undifferentiated cells of the appendages or of their
branches with thin walls.
Forming massive multicellular plants. Family Zodiomycetaceae

3 or more genera. Zodiomyces, etc.

Not forming massive multicellular plants. Family Ceratomycetaceae

4 or more genera. Ceratomyces, etc.

(Many of the groups named by Thaxter should perhaps be made into distinct
families.)

Key to the Orders of Apothecial Fungi (Discomycetes)

Parasitic upon fresh-water algae which they enclose in a definite "lichen" thallus.

Order Lecanorales
Not parasitic upon algae or if so not forming a definite "lichen" thallus.

Apothecia fleshy or leathery, external from the first or emerging more or less
from the substratum; round in outline, less often narrowed, sessile or
stalked. Usually not in a stroma (except Cyttaria). Ascospores discharged
into the air. Order Pezizales

Asci opening by an approximately apical lid (operculum).

Suborder Operculatae
Asci without an operculum. Suborder Inoperculatae

Apothecia fleshy, permanently subterranean, with a simple cavity Uned with
hymenium or the cavity divided by folds and ridges into chambers and
passageways. Ascospores not discharged into the air.

Order Tuberales

KEY TO THE FAMILIES AND IMPOETANT GENERA 245

Apothecia small, laterally compressed, with a narrow, elongated opening; dark-
colored and hard. Mostly bark- and wood-inhabiting.

Order Hysteriales
Apothecia with no definite limiting border, the asci forming a hymenium with-
out paraphyses, usually with very limited hypothecial tissues.

Order Taphrinales

Key to the Families and Important Genera of Apothecial Lichens Occurring in

the United States (Lecanorales)

{Based on Fink and Hedrick, 1935)

Asci disintegrating and the spores forming a powdery coat (mazaedium) on the

surface of the hymenium.
Thallus crustose, without cortex. On trees or decorticated wood.
Apothecia borne on stipes. Family Caliciaceae

Ascospores one septate; not parasitic on other lichens. Calicium
Ascospores not septate, not parasitic on other lichens. Chaenotheca
(Besides the foregoing, five other small genera, two parasitic on other
lichens.)
Apothecia not borne on stipes. Family Cypheliaceae

Ascospores one septate. Cyphelium

(Besides the foregoing two other small genera.)
Thallus foliose or bushy, with cortex. Family Sphaerophoraceae

One genus in the U.S., mainly on soil or rocks, Sphaerophorus

Asci persistent; no mazaedium.

Apothecia irregular, linear or oblong.

Thallus crustose, well-developed "proper exciple" lacking.

Family Arthoniaceae
Ascospores transversely one to several septate. Arthonia

Ascospores both transversely and longitudinally septate.

Arthothelium
Thallus crustose, "proper exciple" present.

Apothecia not seated in a stroma. Family Graphidaceae

Ascospores hyaline or at most light brown.

Ascospores not septate; on old wood. Xylographa

Ascospores septate, paraphyses not branched.

Cells of ascospores cylindrical or cubical. Melaspilea

Cells of ascospores lenticular.

Ascospores only transversely septate. Ghraphis

Ascospores both transversely and longitudinally septate.

Graphina
Ascospores septate, paraphyses branched and interwoven.

Cells of ascospores cylindrical or cubical. Not longitudinally septate.

Opegrapha
Ascospores both transversely and longitudinally septate.

Helminthocarpon
Ascospores brown.

Transversely septate. Phaeographis

Transversely and longitudinally septate. Phaeographina

Apothecia seated in a stroma. Family Chiodectonaceae

Six genera in the U.S.

246 CLASS ASCOMYCETEAE

Thallus fruticose, "proper exciple" present. California.

Family Roccellaceae
Cortical hyphae longitudinal. Dendrographa

Cortical hyphae transverse.

Spores hyaline. Roccella

Spores brown. Schizopelte

Apothecia more or less round or cup-like.
Host algae Myxophyceae.

Thallus taking its form from that of the host.

Thallus squamulose to foUose; on Nostoc. Family Collemaceae
Spermatia produced within thallus. Collemodes

Spermatia produced in spermogonia opening externally.

Ascospores only transversely septate. Synechoblastus

Ascospores both transversely and longitudinally septate.
Definite cortex lacking. Collema

Thin definite cortex present. Leptogmm

Algal host Scytonema or Stigonema. Family Ephebaceae

Seven small genera in the U.S.
Thallus rarely tak'ng its form from that of the algal host.
Thallus large, plainly foliose.

Apothecia without distinct exciple. Family Peltigeraceae

Thallus with cortex above only; apothecium on under surface of

the thallus lobes. Nephroma

Thallus with cortex above and below; apothecia on upper surface.
Ascospores hyaline. Peltigera

Ascospores brownish to brown. Solorina

Apothecia with distinct exciple. Family Stictaceae

Single genus in the U.S. Sticta

Thallus small, crustose to somewhat foliose or dwarf fruticose.

Several small families, mostly growing on soil or rocks: Pyrenop-
sidaceae, Lichinaceae, Heppiaceae, Pannariaceae.
Host algae Chlorophyceae.^

Apothecia without exciple or this only rudimentary (fairly well developed
in Lecanactis) .
Thallus crustose, ascospores nonseptate or transversely septate.

Family Lecanactidaceae
Three small genera, mostly on trees, but some species on rocks or
soil. Not widely spread over the U.S.
Thallus crustose, ascospores transversely and longitudintJly septate.

Family Ectolechiaceae
Only one species, of the genus Lopadiopsis, in the U.S. ^

Apothecia with well-developed exciple.

Both proper and thalloid exciple present, the latter sometimes dis-
appearing.
Apothecia more or less deeply immersed in the thallus.

Family Thelotremaceae
Five genera in the U.S., mostly Southern.
Apothecia superficial or not deeply immersed.

Proper exciple dark. Family Diploschistaceae

' A few species of the foregoing families that are mostly parasitic on Myxophyceae
have Chlorophyceae for their host; see especially Peltigeraceae and Stictaceae.

KEY TO THE FAMILIES AND IMPORTANT GENERA 247

Two small genera in the U.S. of widely distributed species.
Proper exciple hyaline. Family Gyalectaceae

Five small genera in the U.S., from various parts of the country.
Either proper or thalloid exciple present, very rarely both.
Apothecia with proper exciple.

Thallus taking its form from that of the algal host.

Family Coenogoniaceae
Two small genera in the U.S.
Thallus not taking its form from that of the algal host.

Thallus twofold, a primary crustose, squamulose or foliose por-
tion out of which grow the upright podetia.

Family Cladoniaceae
Podetia short, unbranched, hypothecium hyaline.

Baeomyces
Podetia short to long, more or less branched.

Podetia mostly hollow, spores nonseptate. Cladonia
Podetia solid; spores 3 or more septate. Stereocaulon
Thallus foliose, apothecia not on podetia.

Family Gyrophoraceae
Spores nonseptate. Gyrophora

Spores transversely and longitudinally septate.

Umbilicaria
Spores only transversely septate. Derniatiscium

Thallus crustose to squamulose.

Ascospores hyaline, rarely brown, apothecia rarely yellowish,
spores nonseptate to septate, with rectangular or
cubical cells. Family Lecideaceae

Ascospores nonseptate, very large, thick-walled.

Mycoblastus
Ascospores nonseptate, not large.

Thallus crustose. Lecidea

Thallus squamulose. Psora

Ascospores one septate, very large, thick-walled.

Megalospora
Ascospores one septate, small. Catillaria

Ascospores 1 to 3 to more septate, acicular.

Thallus crustose. Bacidia

Thallus squamulose. Tonina

Ascospores 3 or more septate, fusiform. Bilimbia
Ascospores transversely and longitudinally septate.
Ascospores hyaline. Lopadium

Ascospores brown, Rhizocarpon

Ascospores hyaline, nonseptate to septate, with lenticular cells.
Apothecia usually yellowish. Thalloid exciple in one
genus. Family Caloplacaceae

Apothecium with proper exciple. Blastenia

Apothecia with thalloid exciple. Caloplaca

Two or more genera present in the U.S.
Ascospores brown. Thalloid exciple in one genus.

Family Buelliaceae
Apothecia with proper exciple, spores one septate.

Buellia

248

CLASS ASCOMYCETEAE

Apothecia with thalloid exciple. Rinodina

Three more genera present in the U.S.
Apothecia with thalloid exciple. (See also Caloplaca and Rinodina,
above.)
Thallus foliose or rarely somewhat fruticose.

Thallus more or less yellow in color, ascospores hyaline.

Family Teloschistaceae
One genus in the U.S., foliose or some species a little fruticose.

Teloschistes
Thallus greenish gray to ashy or darker, ascospores brown.

Family Physciaceae
Upper cortex plectenchymatous.

Exciple colored like the thallus. Physcia

Exciple becoming black. Pyxine

Upper cortex not plectenchymatous. Anaptychia

Thallus definitely foliose, sometimes also slightly fruticose.
Spores nonseptate, hyaline. Family Parmeliaceae

Thallus yellow, spores 16 to 32 in each ascus.

Candelaria
Thallus rarely yellow, spores 8 in each ascus.

Thallus flat, apothecia not marginal. Parrnelia

Thallus more or less upright, apothecia marginal.

Cetraria
(Besides the foregoing only one other small genus in the
U.S.)
Spores septate.

One genus in the U.S.
Thallus plainly fruticose or pendent.
Ascospores nonseptate.
Thallus dorsiventral.
Thallus upright or pendent.
Greenish gray or straw-colored.
Brownish to brown.
Ascospores 1 to 3 septate.
Thallus crustose to squamulose.
Spores minute, many in each ascus.

Apothecia with thalloid exciple, immersed. Acarospora
Apothecium with proper exciple, immersed to sessile.

Biatorella
Two other small genera in the U.S.
Ascospores usually 8 or less in each ascus. Here may also be
sought some genera in Families Caloplacaceae and Buel-
liaceae (see above).
Ascospores large, with thick wall.

Single genus in the U.S.
Ascospores with thin wall.
Ascospores nf)nseptate, small.
Ascospores 1 to 3 septate. Thallus crustose.

Lecania
Ascospores transversely and longitudinally septate.

Phlydis
(About six other small genera in the U.S.)

Family Stictaceae

Sticta
Family Usneaceae

Evernia

Usnea

Alectoria

Ranialina

Family Acarosporaceae

Family Pertusariaceae
Pertusaria

Family Lecanoraceae
Lecanora

KEY TO THE FAMILIES AND IMPORTANT GENERA 249

Key to the Families and Important Genera of Order Pezizales,

Suborder Operculatae

(Based in part on Seaver, 1928)

Apothecia cup-shaped or discoid, sometimes convex, sessile or stipitate, never
with a pileate structure. Family Pezizaceae

Ascospores globose, hyaline to pale brown. Tribe Sphaerosporeae

Plants growing only on the dung of animals, mature ascospores becoming

pale brown to blackish. Ascodesmis

Plants not restricted to dung.
Apothecia with well-developed bristles or flexuous hairs.
Apothecia red, brown, or greenish, soft and fleshy or waxy,

Sphaerospora
Apothecia black, brownish, or orange, tough or cartilaginous, mostly
stipitate. Pseudopledania

Apothecia not clothed with well-developed hairs.

Apothecia everted to form subglobose ascocarps clothed externally with
the hymenium, somewhat cartilaginous.

Sphaerosoma
Apothecia discoid to convex, with the hymenium limited to the upper
surface and sides; fleshy or waxy.
On the ground or on humus, hymenium strongly convex.

Boudiera
On the ground or on humus, plane or sUghtly convex.

Lamprospora
On living or dead foliage of Conifers, tough.

Pithya
Ascospores ellipsoid to fusoid, rarely subglobose, becoming violet, later brown
or blackish. Tribe Ascoboleae

Spores free in the ascus. Ascobolus

Spores united into a ball in the ascus. Saccobolus

Ascospores ellipsoid to fusoid, hyaline or pale brown.
Ascospores becoming reticulate at maturity.

Tribe Aleurieae
Apothecia clothed with colored hairs. Melastiza

Apothecia without colored hairs, hymenium bright orange.

Aleuria
Apothecia without colored hairs, hymenium dark brown.

Aleurina
Ascospores smooth or sculptured, but never reticulate.

Apothecia small, up to 1 cm. in diameter, not conspicuously hairy or
setose. Tribe Humarieae

Apothecia attached to the substratum clear to their edges, mostly on

wood. Psilopezia

Apothecia attached to the substratum by the center only, mostly on
soil or humus or dung of animals.
Vegetative mycelium superficial, mostly only on burnt places.

Pyronenia
Vegetative mycelium immersed in the substratum.
Asci 8-spored, apothecia fleshy, coprophilous.

Ascophanus

250 CLASS ASCOMYCETEAE

Asci 8-spored, apothecia fleshy, humicolous.

Humarina
Asci 8-spored, apothecia tough. Pseudombrophila

Asci more than 8-spored, bilabiate. Streptotheca
Asci more than 8-spored, with circular operculum.
With small spores. Ryparobius

With large spores. Thecotheus

Apothecia small- to medium-sized, conspicuously setose or hairy.

Tribe Lachneeae
Apothecia partially to entirely buried in the ground.

Sepultaria
Apothecia superficial on substratum, on soil, wood, humus, or dung.
Densely clothed with brick-red hairs. Perrotia
Clothed with hyaline or brown hairs.

Hairs not septate. Lasiobolus

Hairs septate, usually erect or bristle-like.

Patella
Apothecia medium to large, strongly unequal-sided or split on one side.

Tribe Otideeae
Springing in clusters from a buried sclerotium.

Wynnea
Usually isolated, not springing from a sclerotium.
Spores striate with light and dark bands.

PhiUipsia
Spores not striate. ScodeUina

Apothecia medium to large, symmetrical, stipitate, densely hairy or
tomentose, usually tough or leathery.

Tribe Sarcoscypheae
Hymenium bright yellow to scarlet, spores striate.

Cookeina
Hymenium as above, spores not striate. Plectania
Hymenium gray or brown, apothecium externally black or brown.
With a thick gelatinous hypothecium. Bulgaria
Hypothecium not gelatinous.

Opening more or less stellately, attached to sticks, tough.

Vrnula
Not opening stellately, on the ground, fleshy to tough.

Paxina
Apothecia medium to large, sessile to short stipitate, not densely hairy
or tomentose, fleshy and brittle.

Tribe Pezizeae
Apothecium buried in the ground when young.

Sarcosphaera
Apothecium superficial.

Medium-sized, "shaped like an acorn-cup, with a short slender stem"
(but see reference on page 225 to (7. cacabus (Fr.) Sacc.)

(rcopyxis
Medium-sized to large, sessile or with a stout stem-like base.

Ascospores apiculate, apothecium attached by numerous root-like

processes. Rhizina

Ascospores apiculate, apothecia centrally attached.

Discina
Ascospores not apiculate. Peziza

KEY TO THE FAMILIES AND IMPORTANT GENERA 251

Apothecia pileate, subglobose or columnar, pileus always supported by a distinct
stem, never cup-shaped or discoid.

Family Helvellaceae
Pileus costate, the ribs anastomosing to form irregular pits.
Sterile stem and fertile head distinct. Morchella

Fertile part extending to the base, the ascocarp forming a more or less de-
pressed globose structure. Daleomyces {Durandiomyces)
Upper side of the pileus smooth or lacunose but not truly costate.
Pileus bell-shaped. Verpa
Pileus saddle-shaped or irregularly subglobose.

Helvella
Pileus gyrosely convolute (perhaps to be included in Helvella).

Gyromitra
Pileus columnar. Underwoodia

Key to the Families and Important Genera of Order Pezizales,
Suborder Inoperculatae

(Based upon Nannfeldt, 1932, with additions)

Apothecia numerous, opening at the surface of a tuberoid fleshy stroma growing on
the living twigs of Nothofagus. South America and Australasia.

Family Cyttariaceae
Single genus. Cyttaria

Apothecia not buried in a tuberoid fleshy stroma.

Asci cylindrical, the apical thickening hemispherical, with a long slender canal.
Ascospores hyaline, thread-like, often septate and falling apart
into cylindrical cells. Saprophytes. Apothecia sessile, or stalked
(Vibrissea), or immersed and somewhat perithecium-like.
(Order Ostropales of Nannfeldt.)

Family Ostropaceae
Apothecium-like.

Stalked with convex hymenium. Vibrissea

Not stalked, superficial. Apostemidium

Not stalked, emerging from within the substratum. Paraphyses little if

at all branched. Stictis

Not stalked, emerging from wdthin the substratum. Paraphyses much
branched, forming a strong epithecium.

Schizoxylon
Perithecium-like, buried in the substratum.

With short conical neck. Ostropa

With long, upright or horizontal, then upcurved, neck.

Robergea
Asci clavate to ovoid, apex not thickened or only slightly so. Ascospores
spherical to oblong or needle-shaped, never elongated, thread-
like, and falling apart into segments (but see some genera of
Family Geoglossaceae).
Hypothecium and underlying tissues pseudoparenchymatous.

Apothecia in an often lens-shaped, externally almost black, often carbo-
naceous stroma. Excipulum weakly developed. Ascospores
oblong to needle-shaped. Family Phacidiaceae

Apothecia single in each stroma, more or less round, opening mostly by
stellate lobes.

252 CLASS ASCOMYCETEAE

Ascospores ellipsoid, one-celled, paraphyses not forming an epi-

thecium. Phacidium

Ascospores ellipsoid, one-celled, paraphyses forming an epithecium.

Trochila
Ascospores thread- or needle-like, one- to many-celled.

Cocconiyces
Several apothecia in each stroma, opening by elongated slits.
Ascospores ellipsoid. Pseudorhytisma

Ascospores thread- or needle-like. Rhytisma

Apothecia single in elongated stromata, opening by elongated slits.
Ascospores fusiform or rod-shaped, eventually two-celled.

Hypoderma
Ascospores thread-like, one-celled. Lophodermium
Apothecia cartilaginous, leathery or fleshy, superficial or emerging from
the substratum; sessile or tapering to a central stipe-like base.
Stroma if present not dark and hard. Ascospores oblong to
needle-shaped, mostly hyaline. Parasites or saprophytes.

Family Dermateaceae

(including Mollisiaceae)
Apothecia long-lived, cartilaginous or leathery, narrowed at base and
formed on a leathery or cartilaginous stroma which breaks out
through the bark. Ascospores 8 in number, one-celled, later
more-celled. Dermatea (Dermea)

Similar to Dermatea but the ascospores budding in the ascus into innu-
merable tiny spores. Tympanis
Apothecia soft fleshy, no stroma present. Asci small. Ascospores 8 per
ascus, ellipsoid, hyaline. Paraphyses slender, branching some-
what near the tips. Conidial stages Hainesia and Pilidium.
Parasitic on herbaceous tissues.

Discohainesia
Apothecia long-lived, leathery or cartilaginous, not dark-colored, break-
ing out early from the host tissue. Asci large, ascospores
hyaline, simple or septate. Paraphyses thick, often cemented
together by their enlarged tips. Parasitic on woody tissues.

Pezicula
Apothecia waxy or fleshy, breaking out of the host tissue or attached
only by the narrow base. Dark- or light-colored, often hairy
externally. Asci large. Ascospores hyaline one- to several-celled.
Saprophytes or parasites.
Apothecia superficial or attached by the narrow base, rolling together
when dry. Ascospores one- to two-celled.

Mollisia
Apothecia immersed in the host tissues, breaking out at maturity.
Clothed with dark .septate hairs. Pirottaea
Apothecia hairless except the hyaline marginal hairs. On herbaceous
plants except grasses or grass-like species.

Pyrenopeziza
Apothecia formed on the dead stems or leaves of the host plants, the
asexual stages actively parasitic. Exciple not strongly devel-
oped.
Apothecia depressed globose, conidial stage Cylindrosporium.

'^ Higginsia" Nannf.

KEY TO THE FAMILIES AND IMPOKTANT GENERA 253

Apothecia somewhat obconical.
Ascospores one-celled. Conidial stage Gloeosporium-Mke.

Drepanopeziza
Ascospores two-celled. Conidial stage Marssonina or Entommpor-
ium. Diplocarpon

Apothecia developed on the living host tissues and also on the over-
wintered dead tissues, soft fleshy, with better developed stroma
for the overwintered apothecia. Conidial stage Sporonema or
Ramularia or wanting. Ascospore one-celled.

Pseudopeziza
Apothecia superficial, attached to the woody substratum by a narrow stalk,
flat or saucer-shaped, soft fleshy, mostly bright-colored or
hyaline. Asci cyhndrical, small; ascospores 8, one-celled,
hyaline, small. Paraphyses swollen at the apex with a waxy
substance which unites them. Saprophytes.

Family Orbiliaceae
(Nannfeldt recognizes three genera, Orbilia, Patinella, and Hyalinia.)
Apothecia superficial or rarely emerging from the substratum, short- or
long-stalked or sessile, soft, light-colored, often hairy exter-
nally, Ascospores oblong to needle-shaped, hyaline, one- to sev-
eral-celled. Paraphyses thread-like or lance-like. Saprophytes.

Family Hyaloscyphaceae
Apothecia relatively large, almost sessile to long-stalked, mostly clothed
with long rough hairs. Paraphyses usually lance-like, longer
than the asci. Lachnum

Apothecia small, sessile or tapered at base. Paraphyses not lance-like.
Hairs with enlarged base and tapering to a fine point.

Hyaloscypha
(Nannfeldt recognizes some other small genera with small apothecia.)

• Hypothecium and most of the epithecial tissues hyphal (not pseudoparen-
5 chymatous) .

j Apothecia superficial or rarely breaking out of substratum, long- or short-

stalked, rarely sessile, cartilaginous, leathery or fleshy, mostly

• medium-sized or large, hght-colored or dark, asci clavate,

ascospores oblong, hyaline or dark-colored, simple or septate.

f Not rising from definite stromatic masses.

Family Helotiaceae

' Apothecia light-colored, fleshy, almost sessile to long-stalked, outer

,'• surface of apothecia not hairy. Saprophytes.

s Helotium

i Apothecia similar, growing on wood, outer surface clothed with long,

i cylindrical, and somewhat crisped hairs. Parasites or sapro-

• phytes.

Trichoscyphella

(Dasyscijpha in part)
(Nannfeldt recognizes many other genera, some sessile, most stalked.)
Apothecia arising from a definite sclerotium or a stromatized portion of
the substratum, stipitate, asci cylindrical-clavate, 8-spored;
ascospores ellipsoid, hyahne (except two genera), smooth, one-
celled. Mostly parasitic. (Key to genera modified from Whetzel,
1945.) Family Sclerotiniaceae

Stroma a sclerotium of more or less definite and characteristic form.

254 CLASS ASCOMYCETEAE

Apothecia arising from a tuberoid sclerotium that was formed on the
aerial mycelium or in cavities of the host. No conidia known.

Sderotinia
Apothecia arising from a thin effuse, subcuticular sclerotium sur-
rounding the affected portion of the host. Small black micro-
sclerotia also formed on the aerial mycelium. No conidia
known. Stromatinia

Stroma formed in the tissues of the host which it digests and replaces
by the dense sclerotial tissues, except for the remnants of the
host tissues. No conidia known.
Sclerotium of the discoid type, foliicolous.
Apothecia cupulate or saucer-shaped.

Ciborinia
Apothecia verpoid (i.e., curved downward at the apex of the
stalk) . Verpatinia

Stroma destroying the male or female flowers and forming a
sclerotium. Ciboria

Stroma developing in the infected fruits of the hosts, of the hollow
spheroid type or pseudosclerotial. Conidial stage of the Monilia
type. Monilinia

Stroma a typical plano-convex sclerotium formed on or just beneath
the cuticle or epidermis and firmly attached to it. Conidial
stage of the Botrytis type, ascospores hyaline.
Conidial branches not twisted. Botryotinia

Conidial branches strikingly twisted.

Streptotinia
Conidial stage unknown, ascospores olive-brown.

"Martinia" Whetzel
Stroma a more or less angular sclerotium formed in the dead fallen
leaves of the host.
Conidia in clusters, elongated, 2 to 3 septate, on host leaf.

Septotinia
Conidia large, obovoid, produced singly, on host leaf.

Ovulinia
Stroma of the substratal type, indeterminate, not a definite scle-
rotium.
Ascospores one-celled, brown. No conidia known.

Lambertella
Ascospores sometimes septate at maturity, hyaHne. No conidial
stage known. Stroma sometimes rudimentary or wanting.

Rutstroemia Rehm, not Karst.
Ascospores hyaline, not septate. Conidial stage of the Botrytis type,

Seaverinia
Spore fruits club-shaped or stalked with pileus. Asci clavate. Ascospores
oblong to needle-shaped, one-celled or many times trans-
versely septate, hyaline or brown. Conidial stages not known.
Saprophytic. Family Geoglossaceae"

Clavate, the ascigerous portion more or less compressed.
Spores small, one-celled. Plants bright-colored.

Mitrula

* Vibrissea and Apostemidium, placed in this family by Durand (1908), are con-
sidered by Nannfeldt (1932) to belong to the Ostropaceae (see above).

KEY TO THE MORE IMPORTANT GENERA OF ORDER TUBERALES 255

Spores long ellipsoid, 3 to many septate. Plants bright-colored,

Microglossu7n
Spores as in Microglossum, hyaline, plants blackish.

Corynetes
Spores as in Microglossum, but fuliginous or brown.
Hymenium without spines or setae, viscid gelatinous.

Gloeoglossum
No spines or setae, not viscid gelatinous.

Geoglossum
Hymenium with spines or setae. Trichoglossum
Spatulate or fan-shaped, the ascigerous portion decurrent on opposite

sides of the stem. Spathularia

Pileate, i.e., with stalk and rounded head, not black or blackish.
Spores hyaline, ellipsoid-fusiform, eventually 3 to 5 septate.

Leotia
Spores hyaline, clavate filiform, multiseptate.

Cudonia

Key to the More Important Genera of Order Tuberales

{Based mainly on Miss Gilkey, 1939)

Asci and paraphyses in a palisade. Ascocarp hollow, lined with hymenium.
Cavity often reduced by folds or projections to canals or chambers
(in some cases filled with hyphae).
Paraphyses not forming a pseudoparenchymatous epithecium beyond the asci.
Cavity simple, closed or with an opening to the surface of the ascocarp.
Ascospores smooth. Hydnocystis

Ascospores with knobs or minutely verrucose (some species).

Hydnotrya
Cavity divided into canals or chambers, or if not so divided, then with several
openings.
Ascocarp turbinate, narrowed into a short stipe-like structure, see under

Family Helvellaceae. Daleomyces

Ascocarp not narrowed to a stipe.
Ascospores smooth.

Canals converging at one conspicuous opening. Barssia

Canals not converging; openings absent or at least not conspicuous.

Geopora
Ascospores knobby or minutely verrucose (some species)

Hydnotrya
Paraphyses forming a more or less distinct pseudoparenchymatous "secondary
cortex" beyond the asci.
Hymenial areas or canals continuous.
If- Ascospores smooth or minutely granular. Petchiomyces

Ascospores papillose, verrucose, or spinose. Genea

Hymenial areas nest-like, embedded in pseudoparenchyma.

Genabea
Hymenial canals or cavities lined by asci but filled with hyphae.
Canals opening at surface of ascocarp.

Converging at apex. Pachyphloeus

Not converging at apex, fertile only at the dilated blind ends.

Piersonia

256 CLASS ASCOMYCETEAE

Canals not opening to surface, fertile their whole length. Choiromyces
Asci not in a palisade but irregularly arranged.

Ascocarp containing empty canals or chambers or hypha-filled "venae ex-
ternae" (i.e., canals reaching toward the surface but filled with
hyphae).
Cavities empty, not opening to surface. Balsamia

Cavities mostly hypha-filled, opening to surface.

Ascospores smooth. Pseudohalsamia

Ascospores variously sculptured.

Venae externae with parallel venae internae. Tuber

No venae internae.
Asci 8-spored. Hydnobolites

Asci 1- to 4-spored. Delastreopsis

Interior of ascocarp divided by sterile veins into nest-like fruiting areas; no
venae externae or empty canals or chambers.
Ascospores smooth, very large. Picoa

Ascospores sculptured.

Asci 8-spored. Terfezia

Asci 2- to 4-spored. Delastria

fThe genera Hydnobolites, Delastreopsis, Picoa, Terfezia, and Delastria are placed
in some works in Family Terfeziaceae, Order Aspergillales, Fischer,
1896b).

Key to the More Important Genera of Order Hysteriales

{Modified from von Hohnel, 1918)

Ascocarp superficial in subiculum or becoming superficial by emerging from the
substratum, elongated or linear, not strongly compressed nor with strongly
developed keel.
With brown subiculum, ascospores 2-celled, hyaline. Glonium

Without subiculum.

Ascospores 2-celled, hyaline. Psiloglonium

Ascospores with 2 or more cross septa, hyaline. Gloniella

Ascospores similar but colored. Hysterium

Ascospores muriform, hyaline. Gloniopsis

Ascospores muriform, colored. Hysterographium

Ascocarp entirely superficial, strongly compressed laterally and with a well-
developed keel, mussel-shaped.
Ascospores brown, 2-celled. Bulliardiella

Ascospores brown, with 2 or more cross septa. Mytilidion

Ascospores brown, large, fusiform, with many cross septa. Ostreion
Ascospores hyaline, thread-formed. Lophium

The genera with permanently immersed ascocarps, are to be found in the Phacid-
iaceae; Lophodermium, Hypoderma, etc.

Key to the Genera of Order Taphrinales

Asci forming an indeterminate hymenium, on a hyphal hypothecium several
cells thick on the surface of the woody substratum.

Family Ascocorticiaceae
Only one genus. Ascocorticium

LITERATURE CITED 257

Asci forming an indeterminate subcuticular hymenium or forming tufts of asci
emerging from between epidermal cells. Always on living green tissues or
fruits. Mycelium sometimes perennial in the host tissues.

Family Taphrinaceae
Only one genus. Taphrina

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258 CLASS ASCOMYCETEAE

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Elfving, Fr.: Untersuchungen iiber die Flechtengonidien, Acta Soc. Sci. Fennicae,

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Falck, R. : Ueber die Sporenverbreitung bei den Ascomyceten: I. Die radio-

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1934.
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Fischer, 1907.

CLASS ASCOMYCETEAE: THE "PYRENOMYCETES"

WITH the exception of the Order Laboulbeniales the Ascomyceteae so
far considered have been those whose spore fruits are apothecia or
possible modifications of apothecia. In this chapter are discussed those
orders which produce true perithecia with ostioles, or stromata containing
cavities within which the asci are developed. Because the life histories of
all but a few of these thousands of species are entirely unknown the rela-
tionships of these orders to one another and of the genera within the
orders are admittedly very uncertain. Indeed, with further study it is
almost certain that the limits of the orders will have to undergo radical
revision. For this reason a rather conservative attitude is taken as to
these groups. The author does not believe that the true relationships are
shown by the arrangement chosen but this is due to the ignorance of the
ontogeny of the species.

Whether the perithecial forms should be treated after or before the
apothecial forms is a matter of convenience rather than of expression of
phylogenetic history. It seems likely that very early in the development
of the Ascomyceteae three tendencies began to be emphasized. In the one
the female sexual branch bore an oogone surrounded with a protective
layer, as in many Florideae; this group led to the Laboulbeniales. In the
two other series the female sexual organs were at first naked. In one of
these the number of ascogenous hyphae and their branches was great and
a rather wide-spreading spore fruit developed; the apothecial series of
fungi. In the third the ascogenous hyphae were shorter and usually few,
little branching if at all, and the protective structure formed about them
(the perithecium) remained relatively small and was arched over them
and closed or with only a comparatively small opening, the ostiole. So
far we have followed the first two series of development, we now return
to the relatively primitive forms again and start to follow the third series.
Because the spore fruits of this series were more often hard-walled and
small and like the stones of some small fruits the name " Pyrenomycetes "
was applied to include fungi of this nature (from the Greek pyren, the

262

CLASS ASCOMTCETEAE 263

stone of a stone fruit or raspberry, etc., and mykes, a fungus). It should be
noted that the basal groups of each of the three series include forms with
free, nonmotile sperms which come in contact with some receptive struc-
ture (trichogyne) and thus bring about fertilization. In the apothecial
series as well as in the perithecial series this gives place quickly to the
direct union of trichogyne and antherid and finally to the entire dis-
appearance of the trichogyne, and in many cases of the antherid also.

Asexual reproduction probably reaches its highest degree of develop-
ment in this group of orders which may collectively be called the Pyreno-
mycetes, without necessarily indicating that they are at all closely related.
A very large proportion of the species have asexual reproduction by means
of conidia. The types of conidial production are most varied. Often sexual
reproduction is relatively rare, the conidia serving to maintain the species.
In many of the parasitic species the asexual mode of reproduction alone
occurs on the living host and the sexual stage is formed only on the dead
host tissues. The conidia may be produced on free conidiophores or on
conidiophores crowded in an acervulus or on conidiophores enclosed
within a pycnidium. In some cases all of these forms occur in the same
species of fungus at different stages of growth. Judging by the types of
asexual reproduction perhaps the majority of the so-called Imperfect
Fungi (Class Fungi Imperfecti) are probably conidial stages of the Py-
renomycetes, although possibly representing in many cases species that
have permanently lost their power of sexual reproduction.

Studies have been made of the sexual reproduction in various forms of
the group but not only are there great numbers of genera in which studies
are lacking, but for many families no such studies have been made. The
results are conflicting for some of the supposedly related forms. The
stages of sexual reproduction are often difficult to find in a closely con-
nected series of development and it is usually with difficulty that they
can be satisfactorily sectioned and stained. This makes the interpretation
of the structures observed not at all easy and their interpretation is
naturally influenced by previously held views. So in describing the sexual
reproduction of various species the author has had to report the findings
as described and interpreted by the respective investigators although
this course involves some contradictions because of the different inter-
pretations by the various authors. Until many more forms have been
investigated, representing all of the more important genera and families,
it wnll be impossible to determine to what extent weight must be given
to this process in the classification of these fungi.

A number of cases have been selected that show the various types of
structures concerned in sexual reproduction in the Pyrenomycetes, as
well as some of the special modifications of sexual behavior. In many of
these fungi spermogonium-like structures are known within which are

264

CLASS ASCOMYCETEAE

Fig. 87. Sphaeriales, Family Fimetariaceae. Schizotheciujn anserinum (Rabenh.).
(A) Mature perithecium. (B) Longitudinal section of perithecium (paraphyses not
shown). (C) Ascus and paraphyses. (D) Ascospore showing functional dark cell and
empty hyaline cell, and gelatinous appendages. (E) Ascus with three normal asco-
spores and two smaller ones (only the functional cell shows). (F) Ascogonium with
trichogyne. (G) Portion of trichogyne with attached sperm cell. (H) Antherid with
emerging sperm. (I) Sperms clustered at tips of antherids. (A, C, D, after Griffiths:
Mem. Torrey Botan. Club, 11:1-134. B, E-I, after Ames: Mycologia, 26(5):392-414.)

produced sperm cells, like those of Collema and Physcia in the Leca-
norales. In other forms minute sperm-like bodies (often called micro-
conidia) have been recorded but are not produced in definite spermogonia.
In a great many no such structures have been reported.

Ames (1934) found that in Schizothecium anserinum (Rabenh.)
(Pleurage anserina (Rabenh.) Kuntze) small flask-shaped antherids are
formed out of whose necks the small sperm cells are successively pushed,
much as occurs in most of the Laboulbeniales. The ascogonia, produced
from the same mycelium, consist of coiled structures terminating in
slender trichogynes. When a suitable sperm comes into contact with a

CLASS ASCOMYCETEAE 265

trichogyne, often aided perhaps by insects or mites crawling over the
growing mycehum, it adheres to it and soon there occur growth and
multipHcation of the cells and in about four days the perithecium is
mature. Similar structures have been studied in a number of other fungi.
Higgins (1936) studied the development of Mycosphaerella tulipiferae
(Schw.) Higgins which forms its conidial stage (Cercospora liriodendri
E. and H.) on the living leaves oi Liriodendron tulipijera L. and its asciger-
ous stage in the fallen leaves. In the small dark spots formed by the
conidial stage ascogonia may be found, each consisting of an ovoid, uni-
nucleate oogone with a slender uninucleate trichogyne several times its
length. This trichogyne projects from the surface of the leaf. Ostiolate
spermogonia are also produced and within these are many sperm mother
cells in each of which are formed four non-motile, thin-walled sperm cells.
Under the influence of moisture the gum contained in the spermogonia
swells and the sperm cells are extruded and carried passively in the
surface film of water to the trichogynes to which they adhere. Usually
near the tip of the trichogyne a sperm grows fast to a small papilla and
the nucleus enters through the latter and passes down to the oogone,
enlarging as it progresses. The sperm and egg nuclei do not unite in the
oogone but divide conjugately until eight or more pairs are formed, each
pair enclosed in a mass of more deeply staining cytoplasm. Short ascogen-
ous hyphae then arise and the paired nuclei pass into them. Asci are
formed by the crozier method and in these asci the nuclear fusion occurs,
followed by three divisions which thus give rise to the nuclei of the eight
ascospores. When the latter are formed another nuclear division occurs
and each ascospore becomes two-celled. The ascocarp walls and internal
pseudoparenchymatous contents arise as more or less parallel hyphae
growing upward beneath the epidermis before the ascogonium is initiated.
By their growth and branching a pseudoparenchymatous stroma is formed
with a firm black external wall around the developing ascogonium and
ascogenous hyphae. The developing asci press aside or dissolve the color-
less thin-walled cells of the interior of the stroma and eventually an
ostiole is produced through which the mature asci one by one stretch to
the exterior and discharge their ascospores. (Figs. 68, 87.)

In two other species of Mycosphaerella, one with Septoria as its conid-
ial stage and the other with a Cercospora stage Higgins has observed the
formation of trichogyne and sperm cells. According to Wolf (1943) only
about 12 species out of the 500 species of Cercospora listed by Miss
Lieneman (1929) as occurring in North America have their perfect stages
known, all belonging to the genus Mycosphaerella.

The formation of sperm cells is done away with in Venturia inaequalis
(Cke.) Wint., fertilization being effected by the direct passage of several
male nuclei from the antherid through a pore into the tip of the tricho-

266

CLASS ASCOMYCETEAE

Fig. 88. Pseudosphaeriales, Family Pleosporaceae. Venturia inaequalis (Cke.)
Wint. Stages in sexual reproduction. (A) The forked antherid is in contact with the
exserted trichogyne. (B) The sperm nuclei are passing from the antherid (above) into
the trichogyne. (C) The ascogonial cells have become multinucleate and are beginning
to form lobes; the cross walls of the trichogyne have dissolved. (After Killian: Z.
Botan., 9:353-398.)

gyne, as described by Killian (1917) and by Frey (1924). In Fimetaria
{Sordaria) fimicola (Rob.) Griffiths and Seaver it was shown by Greis
(1936) that the tip of a slender antherid fuses with the apex of a coiled
ascogonium. The antherid may arise from adjacent cells or from separate
hyphae. In the lack of an antherid any cell of the ascogonium may send
out an extension, a "pseudotrichogyne," to some nearby cell from which
a nucleus is obtained. This species is self -fertile, i.e., there are not two
sexual phases as in Schizothecium anserinum. Elliott (1925) reported that

CLASS ASCOMYCETEAE 267

in Ophiostoma fimbriatum (E. & H.) Nannf. (Ceratostomella) there arise
uninucleate oogones on short stalks and drawn out into a trichogyne with-
out nucleus or separating septum. From the same or from a nearby hypha
arises an antheridial branch which coils around the oogone and tricho-
gyne. The uninucleate upper cell makes an opening into the trichogyne
through which the male nucleus passes and proceeds downward to the
body of the oogone, enlarging as it goes. As a result of numerous nuclear
divisions occurring conjugately many pairs of nuclei arise which pass out
into nonseptate ascogenous hyphae. In the meantime the hyphae below
the oogone have growm outward and upward to form a closed perithecium
with a thin dark-colored external wall and a mass of hyaline, thin-walled
cells to which the ascogenous hyphae attach themselves and from which
they grow into the perithecial cavity and produce their asci, apparently
without crozier formation, scattered at random through the central hol-
low. Eight ascospores arise in each ascus and ascus walls, ascogenous
hyphae and the remainder of the "nurse cells" digest. In the meantime
the long perithecial neck has developed and finally opened at the end
permitting the mucilaginous mass of ascospores and digested cell walls to
exude as a drop. Gertrud Mittmann (1932) investigating the same fungus,
contrary to Elliott's report failed to find any antherid. According to her
the ascogonium begins as a single, uninucleate somewhat curved terminal
cell of a short lateral branch. The cell elongates and coils into several
turns, dividing into three to five uninucleate cells. From the supporting
cell, enveloping hyphae begin to grow and eventually give rise to the
perithecial wall. Miss Mittmann suggests that the antherid reported by
Elliott was one of these enveloping hyphae. When this envelope is several
cells in thickness the ascogonial cells enlarge and separate somewhat. One,
or sometimes two, of these enlarged cells becomes multinucleate, the
number of nuclei usually being eight. This cell (the oogone) divides into
several cells which also become eight-nucleate and then divide into bi-
nucleate cells which thus form a richly branched system of crowded cells
in no definite order. These cells enlarge laterally and their nuclei unite,
thus forming the young asci which are nourished by the plasma-rich
projecting cells of the inner layer of the perithecial wall. The ascospores
are surrounded by a slime layer. 0. coeruleum (Miinch) H. and P. Sydow
(C. coerulea Miinch) and 0. pluriannulatum (Hedge.) H. and P. Sydow
(C pluriannulata Hedge.) are reported by Miss Mittmann to be quite
similar in development but each falls into two sexual strains, both of
which must be present in order that perithecia shall be formed. If she is
correct in her statements that no antherid is present a union of hyphae at
some other point seems to be indicated. Sartoris (1927) found no antherid
near the coiled ascogonium of 0. adiposum (Butl.) Nannf., studied by
him. Varitchak (1931) found in 0. piceae (Miinch) H. and P. Sydow a

268 CLASS ASCOMYCETEAE

nonfunctional antherid ("trophogone") around which the young ascogo-
nium coiled without uniting with it. (Fig. 89.)

In Chaetomium kunzeanum Zopf and C. bostrychoides Zopf a cell of an
ascogonial coil is fertilized, according to Greis (1941), by an antherid at
the tip of a slender hypha arising near by or at some distance. A coil of
thick binucleate cells results. In the second species there grow up around
this coil, from below, the hyphae that produce the perithecial wall, while
other hyphae grow upward between the ascogenous hyphae arising from
the ascogonium, to form the paraphyses. The terminal cell of each as-
cogenous hypha becomes an ascus, without crozier development. In C.
kunzeanum the perithecium may develop in the foregoing manner, but
usually also there grows out from the thickened ascogonium a stout
creeping, branching extension, up to several centimeters in length, from
which arise, laterally, coils giving rise to ascogenous hyphae, typical
perithecia being formed around them. Thus from one act of fertilization
(cytogamy) there may arise, radiating from this point, up to 10 perithecia.
This is suggestive of the condition in the genus Dudresnaya in the Florid-
eae (Red Seaweeds).

In Xylaria, in which numerous perithecia are produced in a stroma,
coiled ascogonial hyphae were observed by Brown (1913) in the stroma
at points where the future perithecia were due to arise, but their further
development has not been reported. Somewhat similar structures were
reported by Miss Lupo (1922) in Hypoxylon.

Many of the Pyrenomycetes lend themselves readily to cultivation on
various media. In some cases cultures from single ascospores will produce
perithecia, but very often they have not been produced in such cultures.
Edgerton (1914) studied a strain of Glomerella cingidata (St.) Spaul. and
von Sch. in which a scattering development of perithecia was produced on
cultures from a single ascospore but yet when two such cultures were
allowed to grow in contact with each other a great mass of perithecia
appeared along the meeting line in about half the cases, i.e., when the
opposite sexual phases met.

Miss Dowding (1931) and Ames (1932, 1934) have shown for Schizo-
thecium anserinum, as B. 0. Dodge (1927) had shown previously for
Neurospora tetrasperma Shear and Dodge, that the ascospores are usually
four in number and binucleate. From single spore cultures of such spores
perithecia are produced in abundance. Occasionally in place of a bi-
nucleate spore two uninucleate spores are produced. These, Ames has
shown, produce two mutually compatible and self- sterile mycelia while
the binucleate spores produce fertile mycelium, each type of mycelium
bearing both male and female organs. By cutting off and transplanting
hyphal tips containing one or a very few nuclei Ames determined that in
the mycelium from the binucleate spores there are two sorts of nuclei

CLASS ASCOMYCETEAE

269

which when separated by the hyphal tip planting are shown to belong to
the tw^o different strains. On this mycelium with both kinds of nuclei some
branches produce both male and female organs of one phase, self-sterile,
and other branches produce similar organs, also self-sterile, but the two
sorts are inter-fertile. Thus it is clear that genes for incompatibility are
present in the diploid nucleus of the young ascus and during meiosis in

CULWQC/I

BUT

ceossrcKTiLE
IN eiTHCK oiHEcrm

iM TO<}8

HCRWPHROPITIC UirSTPil.r

Fig. 89. Sphaeriales, Family Fimetariaceae. Schizo-
thecium anserinum (Rabenh). Diagrammatic repre-
sentation of the sexual condition. (Courtesy, Ames:
Bull. Torrey Botan. Club, 59(6):341-345.)

the ascus two sorts of nuclei arise, mutually compatible, but self-incom-
patible. There is however no segregation of sex as both kinds of nuclei
carry the potentiality for the production of male and female organs.
(Fig. 89.)

In the genus Neurospora, as well as in Gelasinospora, Dodge, Shear,
Aronescu, Wilcox, and Lindegren (1927-1938) have made an exhaustive
study of the sexual reproduction and of the genetics of the segregation of
self-incompatibility and mutual compatibility as well as of various other
genetic factors. In A'^. sitophila Shear and Dodge and some other species

270 CLASS ASCOMYCETEAE

there are formed eight ascospores, each with a single nucleus. Four of these
represent one sexual strain and the other four another sexual strain. As
with the uninucleate ascospores of A^. tetrasperma and of Schizothecium
anserinum the mycelia formed by their germination will form perithecial
primordia and "microspores" (sperm cells) which are self incompatible
but mutually fertile if both sexual strains are present. In N. tetrasperma
and G. tetrasperma Dowding the ascospores normally contain two nuclei
which with rare exceptions represent both sexual strains. As a result when
the spores germinate the mycelial cells contain both types of nuclei and
at the formation of the perithecial primordium no receptive hyphae or
trichogynes are produced and no fertilization of these by microspores or
sperm cells can occur. On mycelia developed from the occasional uni-
nucleate ascospores of these species only one kind of nucleus is present
and the contact of the mycelium or spores from two mycelia of opposite
phase is necessary. In N. tetrasperma this may occur in any of the three
following ways: (1) Microspores (sperm cells) from the mycelium of one
phase are brought into contact with trichogynes that have grown out of
the perithecial primordia of the other phase and fertilize these. This is
the same method by which the perithecia of A^. sitophila are brought to
production. (2) The large conidia or mycelium from them may similarly
fertilize the trichogynes. (3) The mycelia of the two sexual phases may
come into contact and fuse and mutually diploidize one another so that
all the cells of either mycelium come to contain nuclei of both types. Then
as the perithecial primordia arise both types of nuclei being present fer-
tilization from outside is unnecessary. In Gelasinospora tetrasperma this
third method seems to be the usual one if mycelia from uninucleate asco-
spores are concerned as there are no microspores (sperms) nor conidia
(Dowding, 1933). In this species Miss Dowding and A. H. R. Buller (1940)
have demonstrated that the nuclei pass from cell to cell in the mycelium
through the central pore found in every septum. These proceed at the
rate of 4 to 5 mm. per hour which is nearly twice to over twice the rate at
which the mycelium itself grows. Normally in Schizothecium anserinum
the first method is the usual one even when the mycelium grew from a bi-
nucleate spore for the perithecial primordia and the antherids often arise
from hyphal branches that contain only one kind of nucleus. Dodge (1936)
showed that in this species there are various races some of which produce
no sperm cells. When cultures of the two sexual phases of these are mated
perithecia are formed indicating that probably the third method was the
one that was effective in perithecial formation.

Keitt and Palmeter (1938) and Keitt andLangford (1941) have shown
that also in Vcnturia inaequalis (Cke.) Winter there are two sexual phases,
four of the ascospores representing one phase and four the other. Segrega-
tion of the genes that determine the two phases may occur in the first

CLASS ASCOMYCETEAE 271

meiotic division in the ascus, in which case the four spores of similar phase
are placed in succession followed by the four of the other phase. Segrega-
tion may occur in the second meiotic division in which case the nuclei are
distributed otherwise so that two of the four at either end of the row of
eight ascospores will be of one sexual phase and the other two will be of
the other phase.

Lindegren (1932) reported the results of experiments with Neurospora
crassa Shear and Dodge, a species in which each ascus contains eight
uninucleate ascospores. He picked out the individual ascospores in order
one by one from the ascus and obtained cultures which were then mated
to determine their sexual phase. He showed that in about 85 per cent of
the asci the two strains are segregated in the first nuclear division and in
about 15 per cent in the second. No evidence was obtained to indicate
that this segregation ever occurs in the third nuclear division. In M.
sitophila, with eight ascospores per ascus, and N. tetrasperma, with
normally four binucleate ascospores, the segregation also may occur in
either the first or second division, more often in the first.

From the foregoing it seems possible that the frequent failure to obtain
perithecial development in pure cultures from a single ascospore may in
some cases be due to the occurrence of two sexual phases in that species.

The groups of families included in the collective term "Pyreno-
mycetes" differ from those collectively called " Discomycetes " in the
nature of the spore fruit. In the latter group, described in the preceding
chapter, the spore fruit is an apothecium or a modification of it, in which
the asci are in a more or less extensive hymenium which is eventually, in
the typical forms, exposed to the air with numerous paraphyses separating
and supporting the asci. In the Pyrenomycetes the spore fruits are mostly
much smaller, thicker-walled, and rarely opening wide, the ascospores
escaping through a small opening, the ostiole, or by the rupture of the
whole structure. Formerly these small closed or ostiolate structures were
all called perithecia, in contrast to the open apothecia of the Discomy-
cetes. More recent studies in the last forty years, especially by von Hohnel
(1918), Theissen (1913), Arnaud (1925), Nannfeldt (1932)^ and others,
have shown that three distinct structural types have been included under
the name perithecium. Of the thousands of species with so-called peri-
thecia a great many still remain to be studied carefully to determine to
which of these three types they belong.

The first group, called by Nannfeldt the Ascohymeniales, includes the
apothecial orders Pezizales, Lecanorales, and others included in Chapter
9, and three orders with true perithecia, Sphaeriales, Hypocreales, and
Pyrenulales, discussed in this chapter. The second group, called by Nann-
feldt the Ascoloculares, includes what the author considers to constitute
the three orders Pseudosphaeriales, Dothideales, and Hemisphaeriales,

272 CLASS ASCOMYCETEAE

also included in this chapter. The order Erysiphales, with nonostiolate
perithecia, probably contains a mixture of forms with true perithecia, and
therefore more closely related to the Sphaeriales, and possibly some
Pseudosphaeriales or Hemisphaeriales as well as some Aspergillales. The
latter order corresponds to what Nannfeldt and others have called Plecta-
scales. It represents the third group. In it the asci are scattered through-
out the tissues of the more or less perithecium-like structure, not forming
in any manner what might be considered a hymenium. The Erysiphales,
Aspergillales, Myriangiales, and the very simple fungi forming the Sac-
charomycetales form the subject of Chapter 11.

A typical perithecium consists of a more or less hollow structure whose
wall arises from below the sexual organs or at least from below the asco-
gonium, growing outward and around it and the developing ascogenous
hyphae, closing in at the top. Within this there is formed from the
ascogenous hyphae a hymenium of asci lining the perithecial wall or
forming a cluster at the base. Usually paraphyses arising from the
vegetative mycelium which produced the perithecial wall are intermingled
with the asci. Nearer the apex of the perithecial cavity there arise the
periphyses, also of vegetative origin, which converge and drive upward,
helping to produce the ostiole. Apparently in many cases, even in the
perithecia developing apart from one another, the vegetative mycelium
growing up over the perithecial wall forms a more or less distinct
stromatic layer, in which case the true perithecial wall may remain
colorless. This stroma may sometimes be massive, enclosing the whole or
the lower part of the perithecium.

In general the Sphaeriales, Pyrenulales, and Hypocreales are parallel
groups that in all probability will not be maintained separately after life
history studies have shown the true kinships of the fungi which are now
included in them. They all agree in having (with very few exceptions)
ostiolate perithecia. Those of the Sphaeriales are dark-colored and with
fairly firm to hard perithecial walls or surrounding stroma. In the Hypo-
creales the perithecia or enclosing stroma are colorless or bright-colored
and are usually softer. The members of both these groups are saprophytic
or parasitic, usually on vascular plants, and in the rare cases where they
grow upon algae they do not form a lichen thallus. The Pyrenulales are
parasitic upon algae which they enclose in a typical lichen thallus, their
chief distinction from the Lecanorales being the production of perithecia
instead of apothecia.

Julian Miller (1941), for many years a student of the Pyrenomycetes,
has had the courage to combine the Sphaeriales and Hypocreales into one
order under the former name, treating the latter merely as a family, the
Hypocreaceae, in this order.

Many of the species, a number of genera, and several families, that

ORDER SPHAERIALES 273

under the older systems of classification were confidently placed in one or
another of these three orders, have been found to possess structures that
are not true perithecia and so have had to be removed to other orders.
Many more have not been studied from this standpoint and are still kept
in their customary position but some of these, too, will probably have to
be transferred. In the following, where it seems probable that whole
families should be transferred, that has been done but in cases of doubt
where only a few genera have been carefully studied the old classification
(that of Lindau in Engler and Prantl, 1897) is retained.

The term paraphysis has been used differently by various students of
the Pyrenomycetes. Theissen, Sydow, and perhaps the majority of
mycologists, including the author, define as paraphyses those slender
hyphae that develop in the hymenium, growing up from below and ending
free above. In between them arise the asci to which they give protection
and perhaps nourishment. In some groups of Discomycetes, e.g., various
genera of Tuberales and Lecanorales, these paraphyses may enlarge and
branch above the asci and grow fast to the enlargement or branches of
other paraphyses to form a more or less continuous pseudoparenchyma-
tous layer over the tips of the asci, the epithecium. In the Pseudosphae-
riales the sheets of more or less crushed stromatic tissue that remain
between the developing asci and which may be reduced to interascal
hyphae, attached to the base of the hymenium and the roof of the asco-
carp are called by Petrak (1923) true paraphyses while he uses the term
metaphyses for the structures called paraphyses by Theissen, etc. The
term paraphysoid is used with various meanings by different mycologists.
By some it is applied to the filamentous often deliquescent remains of the
stromatic tissue between the asci (the true paraphyses of Petrak) while
by others it is applied to quickly evanescent true paraphyses.

Order Sphaeriales.^ This order is described first not from any convic-
tion that it is more primitive than the other two but because it offers a
better field in which to point out the various developmental directions in
the modification of the perithecia. The perithecial wall is in general dark-
colored, at least in its outer stromatic layer, pseudoparenchymatous in
structure and free from the enclosed asci which arise from its base or part
way up its sides. Sometimes the asci form a dense hymenium but fre-
quently they are more loosely arranged. Paraphyses may be present but
are usually delicate and evanescent and not very numerous. When true
paraphyses are absent the suspicion is aroused that perhaps these organ-

' It should be noted that this name and that of the family Sphaeriaceae are not
properly valid since the generic name Sphaeria has been entirely abandoned because
of its former application indiscriminately to fungi with conidia (i.e., without asci)
and perithecia. Rather than to propose a new set of names the old familiar ones are
retained here.

274 CLASS ASCOMTCETEAE

isms do not rightly belong here but in the Pseudosphaeriales. Periphyses,
i.e., paraphysis-like threads at the edge of the hymenium but not inter-
mingled with the asci, are much more frequent. The asci may reach
maturity at different ages within the perithecium a phenomenon not rare
in the apothecial forms also. The ascospores may be expelled violently
through a pore (not an operculum) at the apex of the ascus or the osmotic
pressure within may rupture the ascus at its middle so that the upper half
is forced off, or the outer layer may break near the apex and contract,
allowing the inner layer to expand and finally burst, or the asci, paraph-
yses, etc., may be digested within the perithecium leaving the ascospores
embedded in the gummy mass. This absorbs water during a rain and
swells, emerging from the ostiole, where the gum is dissolved away and
the spores carried off by the currents in the film of rain water or splashed
off in the droplets caused by the striking rain drops. In some genera only
the ascus stalk digests, the resultant gummy mass containing the un-
changed bodies of the asci. Nannfeldt and some others before him have
suggested that in the true Sphaeriales the asci discharge their ascospores
through an apical pore and mostly do not undergo autodigestion and
that forms in which the latter occurs and no paraphyses are present
belong in the Aspergillales or some other order.

The asci vary from cylindrical to club-shaped or obovate, sometimes
being drawn out below into a narrow stalk-like portion. The ascospores
may lie in one or more rows in the ascus or in a ball-like cluster. The num-
ber is mostly eight but in a few cases only four or even fewer spores are
formed and in others 16 to 32 or even as many as 256 or 512 are reported.
They vary exceedingly in size, shape, structure, and color. They may be
colored or hyaline, 1-celled, 2-celled, several-celled in one row (phrag-
mosporous), many-celled by both cross and longitudinal walls (muriform
or dictyosporous), long and slender, or even tetrahedral. The recurrence
of certain spore and ascus types in what the current classifications con-
sider to be widely distant families has led (and probably rightly) some
mycologists (e.g., Vincens, 1918, 1921; Julian Miller, 1928; von Hohnel,
1918; Wehmeyer, 1926; and others) to attempt to amend the classifica-
tion so as to bring together those forms with similar asci and spores.

The current classifications place first in this order those families in
which the perithecia stand separately upon the surface of the substratum
or but slightly sunken in it. Two of these families have very thin peri-
thecial walls. These are the Firaetariaceae (Sordariaceae) with naked or
almost naked perithecia and the Chaetomiaceae with the i)erithecia
covered with long hairs and with a special tuft of much longer hairs about
the ostiole. The fungi in both families grow on dung or decaying plant
tissues. They are distinguished further by the fact that the asci of the
former expel the ascospores through the ostiole while in the latter the

ORDER SPHAERIALES 275

asci are digested and the ascospores escape in a mass of slime. Nannfeldt
(1932) therefore places the Chaetomiaceae in the group he calls Plecta-
scales, in this book called Aspergillales. His reason for so doing is that
there are no paraphyses or periphyses and that the asci dissolve into
slime. Closely related to the Fimetariaceae is the family Melanosporaceae,
also with thin perithecial walls, often but not always with a more or less
well-developed neck, and with ascospores mainly inclined to be dark-
colored and lemon-shaped as in most of the species of the two other
families. Neurospora and Gelasinospora probably belong here or in the
Fimetariaceae. The Melanosporaceae are often placed in the order
Hj^pocreales because of the almost colorless or light brown perithecia but
it seems best to place them here since the chief difference is only their
sometimes lighter color. (Fig. 90 C, D.)

Family Fimetariaceae. The chief genera are Fimetaria {Sordaria)
and Schizothecium {Pleurage or Podospora) . Both are mostly found on the
dung of various animals. Some species have four-spored asci and others
asci with eight spores, while in a few species the number of spores is
much larger. In Fimetaria the spores are surrounded by a layer of water-
soluble slime on all sides except a small spot on one end. In Schizothecium
the spores are two-celled, one cell slender and empty and the other en-
larged and dark-colored. There is a long gelatinous appendage at each
end, sometimes several at one of the ends. Hypocopra is like Fimetaria
but the perithecia are immersed in a stroma, an exceptional case for this
group of families. (Fig. 87.)

Family Chaetomiaceae. Chaetomium is the characteristic and most
frequently found genus of the Chaetomiaceae. Its long ostiolar hairs may
be stiff and straight, or wavy, or loosely or tightly coiled, depending upon
the species. The spores are mostly lemoru-shaped and dark-colored and
one-celled. The species are very numerous on damp straw, pasteboard,
etc., as well as on manure. Some species are destructive to cloth and other
vegetable fabrics, especially in the warmer and more humid portions of
the world. (Fig. 90 A, B.)

Family Sphaeriaceae.^ Perithecia with firmer wall and with simple
ostiole or at most with a low papilla. Perithecia superficial on the sub-
stratum or sometimes on a felty mass of mycelium (subiculum). The
twenty or more genera in the family are mostly of little economic interest
except the genus Rosellinia. This deserves mention because of the wide-
spread occurrence of its species (over 200 are known from all parts of the
world), mostly on wood and bark. Several species are dangerous parasites,
e.g., R. necatrix (Hart.) Berl. on the roots and underground portions of
the stems of the grape (Vitis). The perithecia of Rosellinia are nearly

^ See footnote on page 273.

276

CLASS ASCOMYCETEAE

Fig 90. Sphaeriales. Various types. (A, B) Family Chaetomiaceae. Chaetomium
aterrimum E. & E. (A) Perithecium (showing the straight body hairs, the coiled oral
hairs, and the mass of exuded ascospores. (B) Terminal portion of an oral hair. (C, D)
Family Melanosporaceae. Melanospora chionea (Fr.) Corda. (C) Several perithecia
on decaying leaf. (D) Vertical section through perithecium. (E, F) Family Sphaeri-
aceae. Kosellinia aquila (Fr.) de Not. (E) Vertical section through perithecmm and

(Continued on facing page.)

ORDER SPHAERIALES 277

spherical and have a small ostiolar papilla. They sit externally on the
host with their bases sunk in a more or less well-developed subiculum.
The ellipsoidal, colored, one-celled ascospores are eight to each ascus.
Filamentous paraphyses are present. Because of the ascospore characters,
Vincens (1921), Wehmeyer (1926), and Miller (1928) suggest that this
genus belongs more properly near the Xylariaceae. Other genera in the
family vary as to color and number of cells in the ascospores as well as to
the hairiness of the perithecia. In many genera the asci ripen successively
and project one or two at a time from the ostiole to discharge their spores,
the emptied asci contracting back into the perithecium and giving place
to the next maturing asci. (Fig. 90 E, F.)

Family Ceratostomataceae. In this family the perithecia have a
distinct, sometimes fairly long neck, otherwise much like the Sphaeri-
aceae. The perithecial walls are mostly leathery rather than brittle. The
asci are mostly accompanied with paraphyses and do not digest as in the
next family. The ascospores are one-celled, two-celled, phragmosporous
or muriform, and hyaline or brown. The species are mostly saprophytic,
growing on wood, bark, or sometimes on stems of herbaceous plants. The
genus Ceratostomella, more properly called Ophiostoma, has been segre-
gated to form the following family, which is discussed here although
Nannfeldt (1932) places it in Order Aspergillales.

Family Ophiostomataceae. The perithecia have very long necks.
In the genus Ophiostoma (Ceratostomella) the neck is often several times as
long as the diameter of the perithecium. It has a thin perithecial wall and
the roundish asci are scattered throughout the perithecial cavity, without
paraphyses. The ascus walls undergo autodigestion and with the absorp-
tion of water the resultant gummy mass swells and escapes from the apex
of the neck as a hyaline drop containing thousands of the one-celled
hyaline spores. Because of these characters Nannfeldt placed this family
in the Aspergillales. Several species of Ophiostoma grow on the wood of
various trees whose sapwood takes on a blue color, the so-called "sap
stain," owing to the presence of the mycelium in the wood cells. The
conidial stages of the various species of this genus have been described
under a number of names in accordance with the type of conidia and
conidiophores. Chalara or Thielaviopsis produce their conidia endoge-
nously. In Cephalosporium the conidia are produced externally on separate

Fig. 90 — (Continued)

subiculum. (F) Asci, mature and young, and paraphyses. (G) Family Ophiostoma-
taceae. Ophiostoma ulmi (Buis.) Nannf. Perithecium with emerging mucilaginous
mass of ascospores. (A-B, after Greathouse and Ames: Mycologia, 37(1):138-155.
C-D, after Ellis and Everhart: The North American Pyrenomycetes. E-F, after
Berlese: Riv. patol. vegetale, 1(1):5-17; (2):33-46. G, after Buisman: Tijdschrift over
Plantenziekten, 38(1) :l-5.)

278 CLASS ASCOMTCETEAE

conidiophores. In Graphium similar conidia are produced but the conidio-
phores are united together in a tall dark-colored stalk (technically a
synnema). In most cases the conidia are embedded in slime as are the
ascospores and are distributed mainly by insects, which bore in the wood
or bark, to whose bodies the spores adhere. The dreaded "Dutch Elm
Disease" has for its perfect stage 0. ulmi (Buis.) Nannf., Graphium ulmi
Buis. being the conidial stage. This as well as several other species has
two sexual strains which must be brought into contact before perithecia
will appear. Dade (1928) found that this was also true of 0. paradoxum
(Dade) Nannf. This has for its conidial stage Thielaviopsis. (Fig. 90 G.)

Richard and Olga Falck (1947) create another "class" of the "Asco-
mycetales," which they call "Class Haerangiomycetes." In this class they
place those species of Melanospora and Ceratostomeila (Ophiostoma) in
which the ascus does not possess a definite cell wall but merely a plasma
membrane or where the ascus wall is almost immediately dissolved after
its formation. In these fungi, therefore, the ascus does not exercise its
normal function of ascospore dispersion but these spores are carried out
through the ostiole in a mass of "mucus" and rest in a drop in the funnel-
like "haerangium" formed by filaments diverging from the edge of the
ostiole. In the ascus the spores are formed, eight in number, shaped like
the segments of an orange and arranged in a similar manner. This struc-
ture the authors call an "octophore." This "class" is considered to be an
evolutionary development from Sphaeriales in which definite, functional
asci occur.

Family Cucurbit ariaceae. This family formerly included in this
order undoubtedly belongs in the Order Pseudosphaeriales under which
it is discussed.

Family Lophiostomataceae. Except for the base which is partly
sunk in the substratum and the laterally compressed ostiolar papilla
this family differs very little from the Sphaeriaceae. The ostiole com-
pressed into a slit resembles somewhat that of the Hysteriales. This
resemblance is only superficial for the lateral compression is confined
mainly to the ostiole and ostiolar papilla in this family while in the Hys-
teriales the whole'spore fruit is laterally compressed. Lophiostoma is the
largest genus of the family. The fungi of this family are mostly saprophy-
tic on bark, wood, or dead herbaceous stems while only a few species
are possibly parasitic. Conidial stages are known for only a few forms.
(Fig. 91 A.")

Family Amphisphaeriaoeae. In general appearance except for its
circular ostiole this resembles the preceding family, but the study of the
development and the inner structure of the perithecia indicate that it
probably may belong to the order Pseudosphaeriales.

Usually placed next are the families with perithecia entirely sunken in

OKDER SPHAERIALES

279

Fig. 91. Sphaeriales. Various types. (A) Family Lophiostomataceae. Schizostonia
montellicum Sacc. Perithecium. (B-D) Family Gnomoniaceae. Glomerella cingulata
(Stone.) Spauld. & von Schr. (B) Acervulus with conidia. (C) Perithecia in a stro-
matic base. (D) Ascus. (E, F) Family Allantosphaeriaceae. Diatrype virescens (Schw.)
Cke. (E) Vertical section through emergent stroma. (F) Ascus. (A, after Berlese, from
Engler and Prantl: Die Naturlichen Pflanzenfamilien, Leipzig, W. Engelmann.
B-D, courtesy, Stoneman: Botan. Gaz., 26(2):69-120, Univ. Chicago Press. E-F,
from Berlese: Icones Fungorum, 3:1-120.)

280 CLASS ASCOMYCETEAE

the host tissues except for the projecting ostiole. A crust-hke stroma
(clypeus) may in some cases connect the upper portions of the ascocarps.

Family Gnomoniaceae. These are parasitic or saprophytic in the
leaves, stems, and other portions of vascular plants. The asci are usually
thickened above and with a distinctly visible pore. The necks of the
sunken perithecia project well above the surface. Many of the species are
parasitic and produce the conidia on the living tissues of the host but the
perithecia are produced only on the dead tissues. The conidia are usually
produced in acervuli, in gummy masses which are distributed by rain and
insects when wet but harden into a horny mass when dry. InGnomonia the
perithecia are not in a stroma. G. veneta (Sacc. & Speg.) Kleb. on the
plane tree or sycamore (Platanus) has various types of conidial forms
that have been described in the following form genera: Gloeosjporium,
Discula, Sporonenia, and Fusicoccum. This causes leaf scorch and leaf fall
and kills the twigs and sometimes the larger branches. Glomerella is like
Gnomonia except that the perithecia are embedded in a stroma. Gl.
cingulata (Stone.) Spauld. & von Schr. is found on a large number of
hosts and has for its conidial stage forms that have been described as
CoUetotrichum and Gloeosporium, depending upon the presence or absence
respectively of setae around the edge of the acervulus. This species
causes various forms of diseases: bitter rot of the apple {Malus), withertip
of the twigs and tearstain of the fruits of orange (Citrus), anthracnose of
mango (Mangifera) and avocado (Persea), etc. (Fig. 91 B-D.)

The two families Pleosporaceae and Mycosphaerellaceae are in the
older classifications placed next to the foregoing. Their asci are not
thickened at the apex nor provided with a pore. The structure of the
ascocarp is such that these fungi must be transferred to the order Pseudo-
sphaeriales, where they are given consideration.

In contrast to the foregoing families in which in the main the peri-
thecia are not immersed in the fungus stroma there is found a group of
fungi with varying degrees of stromatic development. Lindau (1897), in
Engler and Prantl's "Die Natiirlichen Pflanzenfamilien," divides these
organisms into five families: Valsaceae, Melanconidaceae, Diatrypaceae,
Melogrammataceae, and Xylariaceae. The modern mycologists are in-
clined to reduce the first four to two, thus recognizing only three families.
The most extensive recent work on this group is a series of studies by
Wehmeyer (1926, 1933) on the life histories of these fungi. He points out
that these stromatic forms exhibit a gradual transition from fungi in
which the stroma is vague in outline and not very definite in structure to
those with a highly organized stroma. In the simplest type of stroma the
surface of the substratum is blackened by the coloring of the mycelium.
Wehmeyer states: "The next step in stromatic development comes about
by the proliferation of the mycelium within the substratum. As this

OEDER SPHAEEIALES 281

formation of mycelium increases it usually becomes more or less localized
about the forming perithecia." Wehmeyer distinguishes between ecto-
stroma and entostroma thus: "Ectostroma is that portion of the stroma
which is formed on the surface of the bark, beneath or within the peri-
derm, and which consists typically of fungous tissue only, except that
when it is developed within the periderm it may contain the remnants of
the periderm cells, but never of the bark cortex cells. An entostroma is
that portion of the stroma which develops within the cortical or woody
tissue of the host or substratum and is made up of components of both
fungous and host tissues or substratum tissues." Concerning the develop-
ment of the entostroma in the progressive specialization of the stroma
Wehmeyer remarks: "There is usually correlated, very often beneath a
differentiated ectostroma, a clustering of the perithecia." This ento-
stroma is often delimited from the surrounding tissues by a thin zone of
blackened tissue, forming the black line visible on cutting through the
host tissue. Among the variations found in this group of organisms may
be noted the following: In a fruiting area (i.e., the region where the
perithecia or clusters of perithecia are formed) the perithecia may be
scattered or clustered, with or without an entostromatic mycelium about
them. If present, according to Wehmeyer, the "entostromatic area may
or may not be surrounded by a darkened unorganized zone." It may be
lighter in color than the surrounding bark tissue. The ostioles of the
perithecia may be separately or collectively erumpent, even clustered
perithecia not necessarily being collectively erumpent. A stroma may be
effused, i.e., containing numerous separately erumpent perithecia or
several clusters of perithecia, or isolated, when it contains only one cluster
of perithecia. The portion of the stroma which is erumpent through the
periderm or epidermis is the disk. It may be conical or cushion shaped
and well distinguished from the entostroma or grading into it. In con-
trast to the foregoing are those fungi in which the stroma is compact and
composed entirely of fungous tissues and very early becoming external to
the substratum.

The three families of the stromatic Sphaeriales may be distinguished
as follows:

AUantosphaeriaceae : stroma showing all degrees of development described
above, but not entirely of fungal structure. Asci with more or less elongated
tapering stalks, forming a persistent hymenial layer. Paraphyses mostly
evanescent at maturity. Ascospores mostly allantoid, yellowish hyaline,
sometimes inequilaterally ellipsoid and brown. Conidia long cylindrical to
filiform.

Diaporthaceae: stroma as in the foregoing family. Asci with short or long
evanescent stalks soluble in water so that at maturity the free asci and
spores form a loose central mass. Paraphyses present. Ascospores ellipsoid,
fusoid, less commonly allantoid, or long cylindrical, hyaline or colored.

282 CLASS ASCOMYCETEAE

Conidia of two types: short cylindrical to filiform and ellipsoid to long
cylindrical.
Xylariaceae: stroma well developed, entirely fungal, almost always external,
at least eventually, and covered at first by a conidial layer. Asci long, cylin-
drical, ascospores one-celled, inequilaterally ellipsoid, dark brown, para-
physes filiform.

To these stromatic families probably should be added:

Phyllachoraceae (formerly included in Order Dothideales) : the members of
this family are leaf parasites with the stroma extending from the upper to
the lower surface or between cuticle and epidermis or between epidermis and
palisade layer. The perithecial walls are present, and true paraphyses are
produced.

Family Allantosphaeriaceae. The following genera may be men-
tioned as they are mostly very frequent: In the Allantosphaeriaceae
one of the commonest genera is Diatrype with stroma effuse or isolated,
ectostroma deciduous, exposing a widely erumpent entostromatic disk.
Perithecia parallel, separately erumpent. Eight ascospores, allantoid.
Many species. Saprophytes or weak parasites on twigs and branches. Dia-
trypella is similar in many respects but the ascospores are numerous in the
ascus. Eidypella is much the same as Diatrype but the perithecia are
clustered and collectively erumpent. Anthostoma has stroma effuse or iso-
lated, perithecia separately or collectively erumpent. Asci, in contrast to
the foregoing genera, cylindrical, short-stalked, the eight ascospores in-
equilaterally ellipsoid and dark brown. A transitional form in ascus and
spore structure to the Xylariaceae, and perhaps more properly placed in
that family. (Fig. 91 E, F.)

Family Diaporthaceae. In this family the genus Diaporthe with 600
or more species is the largest or almost so. Stroma effuse or isolated and
entostroma light-colored with a dark border zone. Ascospores ellipsoid or
fusoid, hyaline, tw^o-celled. Imperfect stage belonging to the form genus
Phomopsis. In the genus Valsa the stromata are isolated and the perithecia
clustered in the unaltered bark tissues beneath a distinct conical ecto-
stroma. No marginal zone. Eight ascospores, allantoid, one-celled, hy-
aline. Imperfect stage belonging to the form genus Cytospora. In Leuco-
stoma the stromata are isolated or confluent with a dark marginal zone
about each perithecial cluster. Asci and spores as in Valsa. Valsella
resembles Leucostoma but the asci are polysporous. In Endothia the
stromata are isolated or confluent with strongly developed, colored ento-
stroma. Eight ascospores, allantoid to ellipsoid, one- or two-celled. E.
parasitica (Murr.) And. & And. is the fungus which has destroyed nearly
all the trees of the American chestnut {Castanea dentata (Marsh.) Borkh.)
since the fungus was introduced from Eastern Asia on nursery stock
about 1900 or a little earlier. (Fig. 92 A-E.)

Fig. 92. Sphaeriales. Various types. (A-E) Family Diaporthaceae. Diaporthe
ardii (Lasch) Nit. (A) Section through stroma with perithecia. (B) Section through
stroma with pycnidium. (C) Ascospores. (D) Alpha conidia. (E) Beta conidia. (F-I)
Family Xylariaceae. (F, G) Hypoxylon marginatum (Schw.) Berk. (F) Vertical section
through stroma showing perithecia. (G) Asci of various ages. (H, I) Xylaria suh-
terranea (Schw.) Sacc. (H) Stromata growing from piece of wood. (I) Section of
stroma enlarged to show perithecia. (A-E, courtesy, Wehmeyer; The Genus Diaporthe
Nitschke and Its Segregates, Ann Arbor, Univ. Michigan Press. F-I, from Ellis and
Everhart: The North American Pyrenomycetes.)

283

284 CLASS ASCOMYCETEAE

Family Xylariaceae. Because of the similarity of ascus and asco-
spore structure in the Xylariaceae and in Anthostoma of the Allanto-
sphaeriaceae and of some species of RoseUinia in the Sphaeriaceae it has
been suggested that a more natural classification would group these two
genera with the Xylariaceae. Among the genera undoubtedly belonging
here is Hypoxylon with broadly cushion-shaped to almost spherical
stroma. In this genus as well as in the following genera the conidial layer
is external on the young stromata. A number of species on logs, stumps,
branches, etc. Some of these are 1 cm. or more in diameter and may be
bright red. Daldinia has large rounded stromata with pronounced con-
centric zones visible in vertical section. The fungus grows on dead trunks
and branches and sometimes reaches a diameter of 3 or 4 cm. Its color is
black, sometimes almost varnished in appearance. Daldinia concentrica
(Fr.) Ces. & De Not. shows excellently the distribution of the ascospores.
These are sometimes expelled from the ostioles to a distance of several
millimeters. Near large specimens of this species the dead limbs may be
blackened for a distance of 10 cm. or more. In Xylaria the stroma is up-
right, slender or stout, simple or branched. X. polymoj'pha (Fr.) Grev.
forms thick black clubs usually growing on buried wood. These are
5-8 cm. or more tall and 1-2 cm. thick, rounded at the apex and velvety
at the base. The interior of the stroma is firm and white, the numerous
perithecia forming a distinct layer just beneath the surface. In X. hypo-
xylon (Fr.) Grev. the basal and apical portions of the slender, usually
more or less forked stroma are sterile. This or a closely related species is
parasitic upon the roots of the apple. (Fig. 92 F-I.)

Family Phyllachoraceae. In the great majority of species making
up the subfamily Phyllachorineae, the stroma is endophyllous, frequently
well organized and firm only near the leaf surfaces, the mesophyllic
stroma consisting of less densely compacted hyphae intermingled with
remains of the host cells. Perithecial wall distinct. In some species elon-
gated conidia are produced in subepidermal pycnidial cavities (Bessey,
1919). In the subfamily Trabutiineae the stroma lies between the cuticle
and epidermis and in the Scirrhiineae between the epidermis and the
palisade layer. There were over 40 genera and more than 500 species
recognized by Theissen and Sydow (1915). All were considered to lack
true perithecial walls and therefore to belong to the order Dothideales.
In the genus Phyllachora and a number of other genera definite perithecial
walls are present (Orton, 1924; Petrak, 1924). This character and the
presence of true paraphyses in the vast majority of the species justify
placing the family in the Sphaeriales, although some of the smaller genera
may still have to be retained in the Dothideales. The greater number of
the genera are tropical. Phyllachora graminis (Fr.) Fckl. and other species
are very frequent parasites of various grasses in the United States and

ORDER HYPOCREALES 285

temperate Eurasia. The elongated black stromata resemble unopened
rust sori.

Order Pyrenulales. The perithecial lichens make up a group of about
15 families, over 80 genera, and more than 2000 species. They form typical
lichen thalli in combination with various algal hosts. Most of the species
are crustose or foliose, in only two genera fruticose. The order does not
seem to form a compact monophyletic group but its various families seem
rather to show relationship to different families of the Sphaeriales and
perhaps to some of the Pseudosphaeriales. The perithecia may be sunk
singly in the thallus or may be produced in a stroma strongly resembling
that of Diatrype. Among the common genera may be mentioned Ver-
rucaria, with nearly 300 species forming crustose growths on rocks into
which the hyphae may penetrate to a considerable distance. The peri-
thecia are black and sunken in the thallus. The eight ascospores are one-
celled, ellipsoidal, and hyaline or brown. The algal host is Protococcus or
Palmella. Pyrenula includes about 175 species usually on bark, growing
on the alga Trentepohlia. The perithecia resemble those of Verrucaria,
but the ascospores are several-celled. Long slender conidia are produced
in pycnidia. Trypethelium consists of about 75 bark-inhabiting species,
mostly tropical and subtropical, whose perithecia are produced in a
cushion-like stroma. The 37 or more species of Astrothelium are also
tropical or subtropical, on bark. Their perithecia are arranged radially
ill the stroma with their long necks approximated or joining into a com-
mon ostiole. Practically nothing is known as to the sexual reproduction
of the plants assigned to this order. Spermogonia are known in many
species and may function as they are known to do elsewhere.

Order Hypocreales. This order shows a close parallelism with the
Sphaeriales as to perithecial form and arrangement. The two orders are
customarily distinguished from one another by the consistency and color
of the perithecia. In the latter order the perithecia are dark-colored and
leathery or brittle while in the former they are bright-colored (rarely
dark) and fleshy to leathery. There are border forms such as the genus
Melanospora which has sometimes been placed in one and sometimes in
the other order but which has been treated under the Sphaeriales in this
work. Miller (1941) did not recognize the validity of separating these
groups of fungi as distinct orders and included the family Hypocreaceae
as a distinct family in the Sphaeriales, but in a more recent paper (1949)
concludes that until further studies have been made on the structure and
development of the perithecia the Order Hypocreales should be retained,
except for Family Clavicipitaceae which definitely should be placed in the
Sphaeriales. The genera may be arranged with first the forms with
scattered superficial perithecia, then those with perithecia crowded on the
surface of a stroma, and those with perithecia buried in the substratum

286 CLASS ASCOMYCETEAE

or in a stroma. A further group includes forms in which the perithecia
buried in the stroma do not have well-developed walls of their own but
represent perithecial cavities in the stroma. Only this last type does not
have its counterpart in the Sphaeriales. The ascospores, as in the latter
order, vary from ellipsoidal and one-celled to two-celled, phragmosporous,
muriform, or even thread-like. Some are brown but the majority are hya-
line or bright-colored. Conidial fructifications are rather widespread in
this order. The conidiophores may be separate and external or they may
be packed closely together side by side or may be enclosed in a pycnidium
or united into a stalked head {Stilhella type). Many of the approximately
1000 species are saprophytic; others are parasitic in the leaves, stems, and
roots or other portions of higher plants; still others are parasitic on fungi
or upon insects.

The course of sexual reproduction has been worked out completely in
a few forms, but, as mentioned for the Sphaeriales, only enough is known
to make certain that vastly more must be found out before the knowledge
may be used to modify the current system of classification. The latter,
as in Sphaeriales, is largely based upon the characters of the mature
perithecium.

In a number of Hypocreales a coiled ascogonium and antherid are
known. In Polystigma ruhrum (Fr.) DC, parasitic in the leaves of the
plum (Prunus domestica L.), Blackman and Welsford (.1912) and Nienburg
(1914) have shown that the ascogonium is a stout hypha with several
coils of mostly plurinucleate cells and tapering into a slender, sometimes
branched, trichogyne which may extend through a stoma but apparently
more frequently does not do so. Organs exist which have been called
spermogonia. Whether they really are properly so called remains in
doubt. They usually appear some time after the ascogonia, and their
spores are long and slender and curved like some of the conidia of the
Diaporthaceae. No connection between one of these spores and a tricho-
gyne has been observed. Eventually, according to Nienburg, the wall
breaks down between a multinucleate ascogonial cell and the large uni-
nucleate oogone cell next to it and one nucleus passes into the oogone.
Later ascogenous hyphae are sent out from the latter and eventually give
rise to asci. Blackman and Welsford disagree with the foregoing and claim
that the ascogonium degenerates and that the ascogenous hyphae arise
from near-by vegetative hyphae. In Claviceps purpurea (Fr.) Tul., ergot,
the germinating sclerotia give rise to stalked heads in which arise the
perithecial primordia. This consists for each perithecium, according to
Killian (1919), of a multinucleate rounded oogone from whose base
branch out one or two antherids which also have many nuclei. One of
these antherids comes into contact with the oogone at its tip and an
opening is formed through which the male nuclei enter. This gives rise,

ORDER HYPOCREALES 287

how is not known because certain stages were missed in the investigation,
to a series of binucleate cells which develop into ascogenous hyphae and
form asci by the hook method.

The 60 or more genera making up the order are variously assigned to
one family, to three families, or to still more. The basis of distinction is
the presence or absence of stromata, the location of the perithecia, and
the type of the ascospores. A number of genera with perithecial cavities
in a stroma but without well-developed perithecial walls, and w^th long
slender ascospores and lacking paraphyses, seem to constitute a natural
well-defined family, the Clavicipitaceae. Those with ascospores not of this
type and with distinct perithecia buried in the stroma are usually called
the Hypocreaceae, while those with perithecia external, with or without a
stroma, are placed in the Nectriaceae. The following genera should be
noted : Nectria has perithecia external to the substratum or to an external
stroma on the substratum, round with short ostiolar papilla or none,
usually light-colored, asci in a tuft at the base of the perithecial cavity,
ascospores two-celled, hyaline. Often parasitic on twigs or other plant
tissues. In some species, e.g., N. cinnabarina Fr., there first arises a cush-
ion-like pseudoparenchymatous stroma which bears on its outer surface
a dense layer of slender conidiophores, each bearing a small ellipsoidal
spore {Tuhercularia stage). Later around the base and eventually all over
the stroma arise the round, rather thick-walled perithecia from whose
ostioles escape the ascospores. Over 250 species of Nectria have been de-
scribed. The presence or absence of the stroma has been used by some
mycologists to distinguish two genera. Hypomyces, with 50 or so species,
produces a felt-like stroma or subiculum over the surface of various species
of Agaricaceae, Polyporaceae, etc. The perithecia are essentially like those
of Nectria except for the presence of the stroma or subiculum. Gihherella
produces its blue- or violet-colored perithecia on the stems, grains, etc., of
various plants. Its ascospores vary from two to several cells. G. zeae
(Schw.) Fetch {G. sauhinetii Oud.) is the cause of scab and root rot of
wheat and other cereal grasses and of the root rot of maize. Its asexual
reproduction is by the abundant production of several-celled, sickle-
shaped conidia {Fusarium stage). Subsequently the perithecia appear.
Sphaei'ostilbe has perithecia and ascospores as in Nectria but these arise
around the base of a stalked conidial head of the Stilbella type. It is mostly
parasitic on scale insects and other insects infesting the twigs or leaves
upon which the fungus occurs. Polystigma develops its perithecia in
stromata within the host leaf. Hypocrea has perithecia much like those of
Nectria, but buried in the bright-colored stroma Avhich resembles in many
ways that of Hypoxylon of the Xylariaceae. Over 110 species are known.
In the Clavicipitaceae may be mentioned Epichloe whose stroma develops
as a thick white band around the stems of various grasses. In this white

288

CLASS ASCOMYCETEAE

Fig. 93. (See legend on facing page.)

ORDER HYPOCREALES 289

Stroma the orange-colored perithecia develop, projecting from the surface
by their short ostioles. Cordyceps consists of many species mostly parasitic
on insects but with one or two species growing on subterranean fungi.
They produce stout or slender stalks bearing a round or more often elon-
gated, usually pointed, stromatic head in which arise the numerous
perithecia. The latter may be almost completely buried in the stroma or
they may project from it so as to be almost free. The well-known "vege-
table caterpillar" is a species of this genus. This fungus attacks a cater-
pillar which has entered the ground to pupate and from its body a stalk
several inches high emerges into the air bearing the perithecia in the
stroma in its upper part. Claviceps, the ergot fungus, with a dozen or more
species, produces its purple sclerotia in the spikelets of grasses and related
plants. On the ground, usually after overwintering, they send out stalked
stromatic heads in which the perithecia arise. The ascospores infect the
flower heads of the host species where an external conidia-bearing layer is
produced (Sphacelia stage) . These conidia are borne by insects or rain to
other grasses. Eventually the ovaries are completely filled, or all except a
small portion of the upper end, with a firm stromatic mass which may be
many times as large as the normal ovary. C. purpurea (Fr.) Tul. is the
commonest species of ergot. It occurs in cultivated rye, less often in wheat,
and in many other grasses. The fresh sclerotia have considerable medicinal
value. They are poisonous when eaten in large quantity as often happens
in time of famine when highly ergotized rye or wheat is consumed by the
underfed populace. Pastures in which this fungus is abundant sometimes
cause serious diseased conditions to develop in the animals feeding there.
(Figs. 93, 94.)

The author has long contended that the color and consistency of the
perithecium or stroma are not at all satisfactory as a basis of distinction
of the Sphaeriales and Hypocreales. The structure and mode of develop-
ment of the perithecium are of far greater value in determining the true
relationships. Many of the Hypocreales should be placed in the Sphae-
riales and perhaps some in the Pseudosphaeriales. The fact that many,
perhaps the majority, of the species of the Hypocreales are described as
lacking paraphyses suggests the need of more intensive research upon the

Fig. 93. Hypocreales. (A-C) Family Nectriaceae. (A, B) Nectria cinnabarina Fr.
(A) Portion of tree branch with conidial stromata (Tubercularia stage) with and with-
out surrounding perithecia. (B) Section through stroma with two perithecia at one
side, the remainder still conidiiferous. (C) Sphaerostilbe gracilipes Tul. Section through
stroma showing two perithecia and a synnema and conidial head of the Stilbella stage.
(D-G) Family Clavicipitaceae. (D) Cordyceps militaris Link. Caterpillar with several
stalked stromata. (E) Cordyceps ophioglossoides Link. Asci containing ascospores.
(F, G) Claviceps purpurea (Fr.) Tul. (F) Section of stromatic head. (G) Section through
a single perithecium. (A-E, after L. R. and C. T. Tulasne: Selecta fungorum carpo-
logia, vol. 3, pp. 1-221. F-G, after Tulasne from Engler and Prantl: Die Naturli-
chen Pflanzenfamilien, Leipzig, W. Engelmann.)

290

CLASS ASCOMYCETEAE

Fig. 94. Hypocreales, Family Clavicipitaceae. Claviceps purpurea (Fr.) Tul. Stromata
growing from sclerotium. (Courtesy, F. C. Strong.)

development of the ascocarp to determine whether it is a true peritheciiim
or stromatic in structure.

Order Dothideales. In contrast to the undoubted Sphaeriales, i.e., those
forms possessing true ostiolate perithecia with asci arising from the bot-
tom and sides of the perithecial wall in a common perithecial cavity, and
with periphyses near the ostiole and paraphyses among the asci, are a
large number of genera formerly more or less closely associated with that
order. Of these the Order Dothideales is recognized as a distinct order by
most mycologists. It is mostly defined as consisting of fungi parasitic
usually on leaves, producing endophyllous or epiphyllous stromata within
which arise perithecial cavities which lack definite perithecial walls.
Apparently in this order the ascogenous hyphae arise in the center or base
of the stroma and, spreading outward and upward through the stromatic
tissues, dissolve out cavities within which the numerous asci are produced.
Most of the species are tropical or subtropical, but a few forms included
in the order reach the north temperate zone. Theissen and Sydow (1915)
in their monograph of this order included four families of which all except
the Dothideaceae have been found to have their closer relationship with
other orders: the Phyllachoraceae with the Sphaeriales, the Polystomel-

ORDER DOTHIDEALES 291

laceae with the Hemisphaeriales and the Montagnellaceae with the
Pseiidosphaeriales. This leaves but the single family Dothideaceae with
about 34 genera and over 100 species.

The two orders Hemisphaeriales and Pseudosphaeriales are stromatic
forms, without true perithecia. In these the asci arise from ascogenous
hyphae developed in the midst of a more or less pseudoparenchymatous
stromatic tissue. These ascogenous hyphae spread through this stroma
making their way by pressure and by dissolution of the tissues, eventually
forming cavities within which single asci are produced, separated from one
another by a thinner or thicker remnant of the original stromatic tissue.
In some cases these stromatic remnants entirely disappear so that a clus-
ter of asci arises in the cavity dissolved by their actions in the stroma, as
in the Dothideales. The chief difference from the latter is that but one
such locule is formed in a perithecium-like stroma instead of many locules
in a more massive stroma. The intervening stromatic tissue, where the
asci are close together in monascous cavities, was in many cases formerly
mistaken for paraphyses but can be distinguished by the fact that it is
fastened above as well as below, and often laterally. Those forms in which
the "paraphyses" are described as attached reticulately to each other
belong in this series as do those where their tips form a continuous pseudo-
parenchyma above the apices of the asci. Such structures must be sharply
distinguished from the epithecium found in many Tuberales and Lecano-
rales and in some other groups of fungi in which true paraphyses overtop
the asci and fuse with one another above the latter. The asci are never
operculate. They are usually much thickened, at least upward, and are
obovoid or clavate, rarely slender and cylindrical. The ascospores vary
from one-celled and hyaline to phragmosporous or muriform, sometimes
hyaline and sometimes colored. The stromata may resemble simple
perithecia, the central portion of the apex breaking away as a pseudo-
ostiole. The asci may arise parallel in a row at the bottom of the cavity or
they may arise in a fan-shaped cluster from a raised "placenta" at the
center of the base. In the latter type there are usually no paraphysis-like
remnants of the stromatic tissue, this having been dissolved or pushed
back as the asci grew, while in the former type these fragments usually
persist until the maturity of the asci. Petrak (1923), Gaumann (1928), and
others are inclined to consider all perithecium-like structures with a
spreading, nonparaphysate basal cluster of asci as pseudosphaeriaceous,
even though the spore fruit may appear to possess a true perithecial wall
and a typically developed ostiole lined with periphyses. Whether such
forms are in reality intermediate, as they believe, between the Pseudo-
sphaeriales and the Sphaeriales or not, will require much further investi-
gation to determine. It seems clear that the typical structure is very
different in the two orders. Perhaps Nannfeldt (1932) is right in question-

292 CLASS ASCOMTCETEAE

ing the validity of the assumption of such intermediate forms, drawing the
distinction not so much on the external and structural characters of the
wall as on the nature of the hymenium. If true paraphyses (the meta-
physes of Petrak) arise from the floor of the perithecium among the slen-
der mostly parallel asci, ending free above, these are to be considered as
true Sphaeriales, while the spreading broader asci which entirely crowd
hack the stromatic tissues, or the rather broad, thick-walled asci which
arise in monascous cavities, leaving paraphysis-like threads attached both
above and below, indicate their connection to the Pseudosphaeriales, even
if the external structures are similar. Unless the latter position is taken
we have the anomaly of one genus, Leptosphaeria, with some species con-
sidered as Pseudosphaeriales, others as intermediate forms, and still
others as true Sphaeriales (Petrak, 1923, Gaumann, 1928). The forms
with more massive stromata which have at maturity several cavities
would find their place better in the Dothideales. Where these perithecium-
like stromatic structures arise separately or crowded in clusters we have
the Pseudosphaeriales. Where the stromata have been developed super-
ficially on the leaves of the host they are mostly small, hemispherical or
disk-like, with the upper surface firmer and the lower portion less firmly
developed. They open at the top by a more or less tearing of the tissue.
In the interior there may be separate monascous cavities or the central
tissues may be dissolved, as in some of the Pseudosphaeriales, leaving a
cavity with a cluster of aparaphysate asci. Such fungi form the Order
Hemisphaeriales, perhaps the majority of which were formerly included
in the old Order Perisporiales. Von Hohnel considers some of the Micro-
thyriaceae to be related to Meliola in this latter order.

The more massive stromatic Myriangiales do not show such similar-
ities to the Sphaeriales as either of the foregoing orders. Studies by Miller
(1938) demonstrate that this order formerly associated with the Pseudo-
sphaeriales does not belong here but has its closest relationship with the
Aspergillales next to which order they are given consideration.

Order Hemisphaeriales. The older interpretation of the fruit bodies
was a perithecium with the basal portion poorly developed while a more
or less shield-shaped perithecial wall formed the upper half. It is now
interpreted as a stroma with a hyphal or pseudoparenchymatous basal
portion and a firmer upper part. At the apex, by breaking of the tissues, a
pseudo-ostiole is often formed. Another view which once received strong
support on the part of a number of mycologists was to consider this struc-
ture as a small apothecium with a poorly developed hypothecium and
with a more or less permanent and rather late-opening cover, as in the
Phacidiales, to which by this theory the order was believed to be related.
Theissen and Sydow (1917) held this viewpoint and von Hohnel (1919)
believed that certain Microthyriaceae are forms transitional to the
Discomyceteae. The order consists of fungi which are entirely superficial

ORDER HEMISPHAERIALES 293

or subcuticular or which have a hypodermal stroma connected with an
epiphyllous stroma by strands of hyphae emerging through the stomata
or other openings. Asexual reproduction is known in a few forms and
consists of the formation of conidia from some of the external hyphae or
in pycnidial structures. With few exceptions the HI or more genera and
over 300 species are leaf parasites, largely tropical but represented in the
temperate zones by a number of genera. Arnaud (1930) has studied the
structures of many of these fungi, especially in relation to the host
tissues.

Kilhan (1922) studied the sexual reproduction in Stigmatea rohertiani
Fr., a form previously included in the Family Mycosphaerellaceae of the
Sphaeriales. A subcuticular pseudoparenchymatous stromatic layer is
produced and in the thicker central portion of this appear several short
cells one of which becomes a binucleate oogone with a receptive papilla
and another a binucleate antherid. After the fusion of these cells there
follow several nuclear divisions and a fusion of male and female nuclei.
The resultant diploid nuclei pass out into the ascogenous hyphae which
give rise to asci arising from the floor of the ascocarp between the loose
stromatic hyphae. This hypothecial floor is pseudoparenchymatous and
the top is hemispherical, of radially arranged hyphae which break to leave
a central ostiole. Several functional oogones and antherids may be found
in each stroma as occurs in formation of the apothecium in Pyronema.
The aberrant nuclear behavior described by Killian suggests that this
process should be reinvestigated.

Luttrell (1940) studied the reproduction in Morenoella quercina (Ell.
& Mart.) Theissen of the Family Microthyriaceae. This species grows on
the leaves of various species of oaks (Quercus). The mycelium is superficial
and forms a network of fine dark hyphae. By the division and radial
growth of several cells of a superficial hypha an elongated shield, one cell
in thickness, is produced. On the under side of this shield is developed a
plectenchymatous layer, two or three cells thick, of hyaline cells. In this
appear ascogenous hyphae of binucleate cells whose terminal cells become
the asci. As these enlarge the roof is broken open along a longitudinal
slit, and eventually the intervening stromatic tissues are destroyed so
that finally the asci stand side by side. While these ascocarps are develop-
ing circular spermogonia are produced, with a single top layer of dark
cells and a central ostiole and a floor of ovoid hyaline cells which break off
successively small rod-shaped spermatia. No union of these spermatia to
other cells was observed.

Theissen and Sydow (1917) recognized five families in this order as
follows:

Family Stigmateaceae. Upper surface of radially arranged hyphae,
arising subcuticularly, vegetative mycelium lacking or almost so. Eleven
genera of which Stigmatea is the type genus of the family.

294

CLASS ASCOMYCETEAE

Fig. 95. Hemisphaeriales, Family Microthyriaceae. Asterina camelliae Syd. &
Butl. Perithecium and mycelium. (After Theissen and Sydow: Ann. Mycol, 15(6) :389-
491.)

Family Polystomellaceae. Stromata with radial structure as in the
preceding, but external to the cuticle and arising from an internal myce-
lium ("hypostroma") from which emerge strands through the epidermis
at various points to give rise to the stromata. There are 39 genera of which
20 form very narrow perithecia which formerly led to their being placed
in the Hysteriales. Parmidaria (perhaps more correctly named Schneepia)
belongs here. Polystomella is the type genus of the family.

Family Microthyriaceae. Stromata with radial structure, vegeta-
tive mycelium and stromata entirely superficial. There are 36 genera with
over 150 species; all but a few are leaf parasites. Here and there, on the
more or less reticulately arranged coarse brown vegetative mycelium
(whi(;h is lacking in a few genera), appear the almost lens-shaped stro-
mata. In each, under the radial centrally ostiolate cover, is a hymenium of
vertically standing asci intermingled with conspicuous or inconspicuous
(rarely lacking) paraphysis-like remnants of the stromatic tissues, which
in a few cases form a definite epithecium-like layer. The stromata are
mostly round but are in some cases laterally compressed. Among the
forms without vegetative mycelium is the genus Microthyrium in which
the stromata appear as little black superficial dots on the leaves or stems
of various plants. Asterina forms small round stromata and Lembosia linear

ORDER PSEUDOSPHAERIALES 295

stromata in the brown vegetative mycelium on the surface of the host.
(Fig. 95.)

Family Trichopeltaceae. The conspicuous mycehum is radial in
arrangement, or forms sterile parallel hyphae. The cover of the stroma
appears to be merely a local thickening of the vegetative mycelium.
Paraphy sis-like threads are lacking. Six genera are known and from 10 to
15 or more species, all tropical.

Family Hemisphaeriaceae. Mycelium lacking or reticulate, super-
ficial. Cover of the stroma not radial in structure. Under the cover there
may be a single hymenium with or without paraphysis-like threads or
several smaller hymenia may be produced under one cover. In some cases
these are reduced to a considerable number of "monascous hymenia,"
i.e., embedded in the hypothecium are scattered single asci. Nineteen
genera of mostly tropical fungi. Micropeltis occurs in the Old and New
World tropics on leaves. Its stroma contains a single hymenium with
many asci and with paraphysis-like structures and with hyaline asco-
spores of four or more cells.

Order Pseudosphaeriales. The fungi included in this order have
been segregated from the Sphaeriales, Perisporiales, and Dothideales,
mainly, and even from the Pezizales. Their true relationship is not known;
indeed it is doubtful whether all the genera assigned to this order are
really related. They are largely tropical, but many occur in temperate
regions. They may be parasitic on plants or even on insects, or sapro-
phytic. The fruiting bodies resemble superficially the perithecia or stro-
mata of the Sphaeriales or some Dothideales. They are almost external or
at least their tops become external by rupture of the host tissues. They are
distinguished from all the preceding orders (except a few species of the
Hemisphaeriaceae with "monascous hymenia") by the mode of occur-
rence of the asci. These arise in separate stromatic cavities, one ascus to
each cavity. In the majority of genera they are oblong or nearly spherical
and mostly eight-spored. They appear to develop somewhat as follows:
Within a pseudoparenchymatous stromatic structure arise branching
ascogenous hyphae, probably — in many cases, if not in all — from an-
ascogonium. These hyphae grow out into the stromatic tissue, dissolving
it so that eventually each terminal ascus lies in a cavity of the original
sterile tissue. These asci may be separated rather widely or the separating
tissue may be but a thin sheet of cells. The developing asci may arise in a
fan-shaped cluster destroying the stromatic tissue as they enlarge. The
ascospores vary from hyaline to brown and from one-celled to many-
celled, in many genera being muriform. The fact that the many-celled
type of ascospore is the most typical for the order, the one-celled hyaline
spore being found only in two genera of the supposed transitional family
Dothioraceae, casts doubt upon the idea that the Pseudosphaeriales are

296 CLASS ASCOMYCETEAE

a somewhat primitive order from which have arisen the Sphaeriales. The
asci become exposed by the weathering away of the outer part of the
stroma or of its apical portion. Rarely the central apical tissues dissolve
away to form an ostiole. In the forms with a small perithecium-like
stroma the tissues between the ascus locules break or dissolve away leav-
ing shreds that have been taken for paraphyses, so that the numerous asci
appear to stand in a true perithecial cavity. Careful investigation has
shown that many species formerly assigned to the genera Pleospora and
Leptosphaeria (Family Pleosporaceae, Order Sphaeriales) have the fore-
going structure, hence must be transferred to this order. Just how far
this may apply to the many remaining species of these and other genera
can be determined only by careful study of the development of the young
ascocarps. To say, as does Gaumann, that these represent a transition
from one order to the other may represent the truth but the author pre-
fers to reserve judgment until further light is thrown on the subject by
ontogenetic investigations throughout the various genera of the Sphae-
riales. It is possible that these are transitional forms but that evolution
has progressed in the contrary direction, from the Sphaeriales to the
Pseudosphaeriales. Sexual reproduction has not been investigated in
enough undoubted members of the order to enable the information gained
to be used in classification. Conidia are produced in a number of families.

Theissen and Sydow recognize several families. The most important
are the following:

Family Pseudosphaeriaceae. The stromata resemble separate peri-
thecia of the Sphaeriales and open at the apex either by crumbling of the
tissues or by their dissolution to produce an ostiole. The asci are clustered
at the base, separated by paraphysis-like hyphae which are attached at
the top as well as bottom. Or it may be that the interior tissues and part
or all of the interlocular tissues may dissolve, leaving a cluster of asci at
the base of a hollow perithecium-like structure so that with mature asco-
carps it can not be determined easily whether the fungus belongs to the
Pseudosphaeriales or to the Sphaeriales. Among the fungi assigned here
with more or less confidence are some species of Pyrenophora, of which P.
teres (Died.) Drechsl. (P. irichostoma) is the perfect stage of Helnimtho-
sporium teres Sacc, the cause of net blotch of barley. In this genus the
ascocarps are hairy and the phragmosporous or muriform ascospores
colored. Whether Dothiora should be included here or in a separate family
is doubtful. In its general structure it resembles somewhat the foregoing
but the perithecium-like stromata are broader and flatter and the muri-
form ascospores hyaline. (Fig. 96.)

Family Mycosphaerellaceae. In the Mycosphaerellaceae the
genera Guignardia and Mycosphaerella deserve attention. G. hidwellii
(Ellis) V. & R. is the cause of the very destructive black rot of the grape.

ORDER PSEUDOSPHAERIALES

297

Native to North America it has been introduced into Europe where it
has caused great damage to the more susceptible Vilis vinifera L. The
fungus attacks the leaves on which it produces brown dead spots and the
fruits which become dark-colored and shrunken. In these leaf spots and

Fig. 96. Pseudosphaeriales, Family Pseudo-
sphaeriaceae. Pyrenophora teres (Died.) Drechsl. (A)
Conidiophores and conidia {Helminthosporium teres
Sacc). (B) Perithecium. (C) Asci in cavities dissolved
in the stromatic tissue. (Courtesy, Drechsler: /. Agr.
Research, 24(8):641-740.)

shriveled berries are produced numerous pycnidia in which arise hyaline,
ellipsoidal spores {Phoma stage of the fungus). In the leaves and berries
which overwinter on the ground there develop perithecium-like structures
in which are produced clusters of eight-spored asci. Paraphyses are not
present. Each hyaline ascospore just at maturity forms a septum dividing
it into two quite unequal parts. The formation in pycnidium-like struc-
tures of "microconidia" which do not appear to be capable of germination

298 CLASS ASCOMYCETEAE

leads to the buggestion that we should search here for fertilization of a
trichogyne by sperm cells. The genus Mycosphaerella contains over 1000
species, many of them parasites of great economic importance. The asco-
spores are hyaline or pale green with a nearly median septum. The conidial
forms are of several types. M. fragariae (Schw.) Lind., which causes the
leaf spot of strawberry {Fragaria), has as its conidial stage Ramularia
tulasnei Sacc. The brown conidiophores project through the stomata of
the diseased spots and produce terminally short chains of cylindrical or
rod-shaped hyaline conidia which are produced in acropetal order. In
M. sentina (Fr.) Schroet., on the leaves of the pear, the conidial stage was
formerly known as Septoria piricola Desm. Here the very long slender
hyaline conidia are produced in pycnidia in the leaf spots. The form
known as Cercospora cerasella Sacc. is the conidial stage of M. cerasella
Aderh. on cherry leaves. The brown conidiophores emerge from the
stomata in the leaf spots and bear terminally or almost so single elongated
several-celled conidia, tapering somewhat toward the apical end. When
one falls off the conidiophore elongates slightly in a sympodial manner
and produces another conidium, and so on, until eventually an old conidi-
ophore may show the scars of attachment of a number of conidia. Myco-
sphaerella pinodes (B. & Bl.) Stone, on the pea, produces pycnidia con-
taining two-celled hyaline conidia {Ascochyta pisi Lib.) while M. tahifica
(P. & D.) Johns., of the beet, has as its conidial stage Phoma hetae Fr., in
which the pycnidia contain hyaline ellipsoidal one-celled conidia. In all
these species and others like them the conidial stage is the destructive
stage while the perithecia are produced in the dead overwintering tissues.
The different types of conidial production have been used by Klebahn
(1918) as a basis for segregating the genus into several subgenera, e.g.,
Ramularisphaerella, Septorisphaerella, Cercosphaerella, etc.

Family Pleosporaceae. In the Family Pleosporaceae the genus
Physalospora has hyaline or pale brown ellipsoidal ascospores. In P.
cydoniae Arnaud, which forms its perithecial stage only on dead twigs, the
conidial stage is the destructive Sphaeropsis malorum Pk., which causes
the black rot of the fruit and the twig blight and canker of the apple and
quince. The conidia are produced in pycnidia and are large, ellipsoidal
and dark-colored when mature, sometimes becoming uniseptate when old.
Venturia inarqualis (Cke.) Wint., the cause of the scab on apple leaves
and fruits, forms its perithecia in the overwintered leaves infected the
previous summer. Its two-celled, slightly colored ascospores are expelled
sometimes to a height of 15 mm. The conidial stage {Fusicladium dendrit-
icum (Wallr.) Fckl.) develops subcuticularly on the leaves and fruit. In this
species both Killian (1917) and Frey (1924) have shown that a well-
developed antherid unites with the trichogyne which terminates the coiled
ascogonium and several male nuclei pass into it and by successive disso-

KEY TO THE ORDERS OF OSTIOLATE "PYRENOMYCETES" 299

lution of the intervening septa reach the oogone cell. Subsequently asco-
genous hyphae are produced. Pleospora has dark-colored muriform asco-
spores. In some species the conidia belong to the form genus Stemphylium
in which the muriform conidia are produced singly and in other species
the conidia are produced in acropetal chains belonging to the form genus
Alternaria.

Family Botryosphaeriaceae. Probably also to be included in the
Order Pseudosphaeriales are the families Botryosphaeriaceae and Cucur-
bitariaceae. These differ from the foregoing families by possessing a more
massive stromatic ascocarp. The ascocarps of these two families possess a
subcortical hypostroma the upper part of which bursts through the bark
and develops as a more or less spherical, perithecium-like body. In the
Botryosphaeriaceae the thick wall has a thin blackened and carbonaceous
outer layer and the asci are broadly clavate.

Family Cucurbitariaceae. In this family the main portion of the
stroma, the hypostroma, is well developed or consists of a limited mass of
mycelium intermingled with the remnants of the host tissue. From the
surface of this basal stroma there bulge out numerous, crowded, more or
less spherical stromatal projections or pseudoperithecia in which the
development of asci occurs without the production of true paraphyses.
These ascocarps have thinner walls than those of the foregoing family,
with a thinner blackened outer layer. Most members of the family are
saprophytic on wood and bark but possibly some are parasitic. In some
species conidia are produced in pycnidia. Cucurhitaria is the largest and
most frequently observed genus. It is placed by Theissen and Sydow
(1916) in this order.

The relationships and phylogeny of the orders considered in this
chapter are not at all certain. Theissen and Sydow, Gaumann (1928), and
Petrak (1923) would derive the Sphaeriales from the Pseudosphaeriales
which in turn they would derive from the Myriangiales and these from
near the Aspergillales. The author is inclined to consider that the simpler
forms of Sphaeriales and Hypocreales, with hyaline or light-colored, one-
celled ascospores and a definite hymenial layer are the more primitive and
that from these have arisen the other orders. This will be discussed more
fully in Chapter 17.

Key to the Orders of Ostiolate "Pyrenomycetes"

Ascocarps are true perithecia with or without marked stromatic outei- layer or
surrounding stromatic tissues. Periphyses and true ostioles usually present.
True paraphyses present or early disappearing, in some cases apparently
entirely lacking.
Perithecia with dark-colored, usually firm walls. Not lichen forming.

Order Sphaeriales
Perithecia dark-colored. Lichen forming. Order Pyrenulales

300 CLASS ASCOMYCETEAE

Perithecia and sustaining or enclosing stromata, when present, light-colored
or red or blue. Not lichen forming. Order Hypocreales

Ascocarps dothideal, i.e., consisting of cavities dissolved in a fairly massive in-
ternal or external stroma, each such perithecial cavity containing a cluster
of parallel or diverging asci separated by the paraphysis-like remains of the
stromatic tissue, or these disappearing very early. Ostioles formed by the
breaking or dissolving away of the apical portion of the stroma above the
perithecial cavity. Order Dothideales

Ascocarps pseudosphaeriaceous, i.e., consisting of perithecium-like stromata within
which the asci are formed in separate monascous cavities, or the intervening
tissues dissolve very early so that the asci form a parallel or diverging cluster
in a single large cavity. No true paraphyses. Ostioles formed by the breaking
or dissolving of the apical portions of the stromata.

Order Pseudosphaeriales

Ascocarps external, consisting of usually flattened stromata with a thin outer,
colored crust, the basal portion pseudoparenchymatous, with thin-walled
cells. The asci are formed in monascous cavities. Mycelium mostly external
or subcuticular, often very slightly developed and not easily detected.

Order Hemisphaeriales

Ascocarp obviously stromatic with rounded asci single in cavities which are in
one or'two layers or irregularly scattered.

Order Myriangiales (see Chap. 11)

Because of the large number of genera in the families the following
keys are made only for the families. A few of the more important genera
are merely mentioned under each family but not keyed out.

Key to the More Important Families of Order Sphaeriales

Perithecial walls thin,

Perithecia light brown, rarely colorless, superficial or sunk in a subiculum,
ostiole with papilla or well-developed neck. Paraphyses wanting.
Ascospores dark-colored, discharged at maturity.

Family Melanosporaceae
Melanospora, Neurospora, Gelasinospora.
Perithecia dark brown, superficial or sunk in a subiculum, rarely partially sunk
in the substratum or in a stroma, not conspicuously hairy. Ascospores
dark-colored, with slime coat or appendages, paraphyses present, asco-
spores discharged through the ostiole.

Family Fimetariaceae (Sordariaceae)
Fimetaria (Sordaria), Schizothecium (Pletirage), Hypocopra, Sporormia.
Perithecia dark, hairy all over and with a tuft of periostiolar bristles. Ascospores
dark-colored. Paraphyses wanting. Asci dissolving at maturity.

Family Chaetomiaceae
Chaetomium, Ascotricha.
Perithecial walls firm and dark.

Perithecia superficial or sitting in a subiculum, ostiole simple or in a low papilla.
Paraphyses mostly present. Ascospores hyaline or colored, one-celled
or several-celled. Asci not dissolving at maturity.

Family Sphaeriaceae
Zignoella, Melanomma, Rosellinia.

KEY TO THE FAMILIES OP ORDER HYPOCREALES 301

Perithecia superficial or slightly sunken, with a long neck. Ascospores one-celled
or several-celled, hyaline or colored, paraphyses usually present, asci
not dissolving at maturity.

Family Ceratostomataceae
Ceratostoma, Ceratosphaeria.
Much like the foregoing but the paraphyses wanting and the asci dissolving at
maturity and the hyaline one-celled ascospores exuding in a drop at
the tip of a long neck.

Family Ophiostomataceae
One genus Ophiostoma {Ceratostomella).
Perithecia in a cluster on a more or less well-developed subcortical stroma.

Family Cucurbitariaceae

(see Order Pseudosphaeriales)
Base of perithecium remaining sunk in the substratum at maturity.
Ostiole laterally compressed.

Family Lophiostomataceae^
Lophiostoma, Lophiotrema, Schizostoma.
Ostiole circular. Family Amphisphaeriaceae ^

Amphisphaeria, Caryospora, Winteria, Strickeria.
Perithecia remaining entirely enclosed in the substratum except the projecting
ostiole. No stroma present except in Glomerella.
Asci thickened at the top, with a distinct pore; ostiole mostly with a distinct
neck. Family Gnomoniaceae

Gnomonia, Glomerella.
Asci not apically thickened and without pore; ostiole simple or with a low
papilla. Families Mycosphaerellaceae and Pleosporaceae

(see Order Pseudosphaeriales)
Perithecia immersed in a stroma, mostly caulicolous.
Stroma in part consisting of remains of the host tissue.

Asci with long tapering stalks, not deliquescent, ascospores mostly
allantoid. Family Allantosphaeriaceae

Diatrype, Euttjpella, Anthostoma.
Stalks of the ascus dissolving early, ascospores ellipsoidal, fusoid, long
cylindrical, less often allantoid.

Family Diaporthaceae
Diaporthe, Valsa, Leucostoma, Endothia.
Stroma consisting entirely of fungous tissue, eventually external, asci cylin-
drical, paraphyses abundant, ascospores dark.

Family Xylariaceae
Hypoxylon, Daldinia, Poronia, Xylaria.
Perithecia in an endophyllous stroma, the perithecial walls well developed,
especially near the ostiole, where they pierce the clypeus-like portion
of the stroma. Leaf parasites. Largely tropical.

Family Phyllachoraceae
Phyllachora, Ophiodothella.

Key to the Families of Order Hypocreales

Perithecia without stroma or external to a stroma; ascospores ellipsoid to cylin-
drical, one- to many-celled. Family Nectriaceae
Nedria, Gibberella, Sphaerostilbe, Scoleconectria, Thyronedria.

2 Possibly belongs to the Order Pseudosphaeriales. \

302 CLASS ASCOMYCETEAE

Perithecia buried in a stroma, ascospores as above. Family Hypocreaceae

Hypocrea, Polystigma, Hypomyces.
Perithecia buried in a stroma with poorly developed perithecial walls, sometimes
only slightly developed near the ostiole. Ascospores filiform.

Family Clavicipitaceae'
Cordyceps, Claviceps, Balansia, Epichloe.

Key to Some Families of Order Pyreniilales*

Perithecia opening through round ostioles; not embedded in a stroma.

Perithecium more or less immersed in the crustose thallus. Paraphyses remain-
ing distinct. Algal host Trentepohlia. Mostly on trees, rarely on rocks.

Family Pyrenulaceae
Pyrenula.
Perithecia more or less immersed in the crustose thallus. Paraphyses mostly gela-
tinizing early. Algal host Pleurococcus. Mostly on rocks, rarely on trees.

Family Verrucariaceae
Verrucaria.
Perithecia immersed in the squamulose or foliose thallus. Paraphyses gelatiniz-
ing early. Algal host Pleurococcus. Mostly on soil or rocks.

Family Dermatocarpaceae
Dermatocarpon.
Perithecia embedded in a stroma. Thallus crustose. Algal host Trentepohlia.
Perithecia opening separately through round ostioles. Paraphyses remaining
distinct. On trees. Family Trypetheliaceae

Trypethelium.
Perithecia arranged radially in the stroma with the ostiolar necks converging
and opening separately or through a common ostiole. Paraphyses remain-
ing distinct. Mostly on trees. Family Astrotheliaceae
Astrotheliuni.
Perithecia in groups of two (rarely one) or more, the intervening walls sometimes
partially missing; the large, irregular ostiole sometimes serving two or more
perithecia. Paraphyses poorly developed or disappearing early or persist-
ing. Thallus crustose. Algal host Palmella or TrentepohHa. On trees.

Family Mycoporaceae
Mycoporum.

Key to the Families of Order Dothideales

Only recognized family.* Family Dothideaceae

Dothidea, Dothidella, Systremma.

Key to the Families of Order Pseudosphaeriales

Stromata small, internal or eventually external, perithecium-like. Asci more or

less parallel, formed in moiiascal cavities which long remain separate by

the persisting intervening stromatal tissues which connect from the base

to the top of the stroma. Family Pseudosphaeriaceae

Pyrenophora, Dothiora.

* Careful studies are needed to determine whether the ascocarps in this order are
true perithecia or pseudoperithecia. The distinction of families as usually made is
largely l)ased on the kinds of algal hosts as well as the habit of the thallus. Only part
of the 10 or more families are mentioned in this key.

*F()r Phyllachoraceae, see Order Sphaeriales.

LITERATURE CITED 303

Stromata small, perithecium-like, subepidermal, often eventually external. At
maturity only a single large cavity with a cluster of spreading asci, the
intervening stromatic tissues entirely destroyed or remaining as remnants
between the basal portions of the asci. Family Mycosphaerellaceae

Mycosphaerella, Dichjmellina, Guignardia.
Similar to the foregoing but the asci mostly parallel with more pronounced
remnants of stromatic tissues between them.

Family Pleosporaceae
Physalospora, Venturia, Pleospora, Leptosphaeria.
Stromata more pronounced, subcortical or subepidermal. Walls surrounding the
large central perithecial cavities thick.
Outer stromatal wall with thick blackened external layer. Asci broadly clavate.
The pseudoperithecia single or scattered. Family Botryosphaeriaceae

Botryosphaeria.
Outer wall with thin blackened layer. Asci elongated, nearly cylindrical. Pseudo-
perithecia densely clustered on the small or large basal portion of the
stroma. Family Cucurbitariaceae

Cucurbitaria.

Key to the Families of Order Hemisphaeriales

{This follows Theissen and Sydow, 1917, and probably needs radical rearrangement)

Ascocarps with more or less pronounced radial structure.

Ascocarps arising subcuticularly but emerging, at least at maturity. Vegetative
mycelium scanty or lacking. Family Stigmateaceae

Stigmatea.
Ascocarps produced externally on strands of hyphae from the internal my-
celium ("hypostroma"). Family Polystomellaceae
Polystomella, Parniularia (Schneepia).
Vegetative mycelium and ascocarps entirely superficial. Ascocarps round or
laterally compressed. Family Microthyriaceae
Microthyriuni, Asterina, Lembosia.
Mycelium conspicuous, external, radial or forming parallel ribbons of closely
united hyphae. Cover of the ascocarp arises as a thickening of the vege-
tative mycelium and is radial at least toward the margin.

Family Trichopeltaceae
Trichopeltis.
Ascocarps not showing radial structure. Mycelium reticulate and superficial or
almost lacking. Family Hemisphaeriaceae

Micropeltis.

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304 CLASS ASCOMYCETEAE

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1-10. 1931.
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Mycologia, 32(4):471-488. Figs. 1-6. 1940.
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Elliott, John A.: A cytological study of Ceratostomella fimbriata (E. & H.)

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Ellis, J. B., and B. M. Everhart: The North American Pyrenomycetes, iii +

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Figs. 1-5. 1947.
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Carroll WiUiam Dodge, xiv + 701 pp. 406 figs. 43 diagrams. New York,

McGraw-Hill Book Co., 192S.
Greathouse, Glenn A., and L. M. Ames: Fabric deterioration by thirteen

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: Tiber Discomyceten vortauschende Microthyriaceen, Ber. deut. botan.

Ges., 36(8):465-470. 1919.
Keitt, G. W., and M. H. Langford: Venturia inaequaHs (Cke.) Wint.: I. A

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and D. H. Palmeter: HeterothalUsm and variability in Venturia in-

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KiLLiAN, Karl: tJber die SexuaUtat von Venturia inaequalis (Cke.) Ad Z

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Klebahn, H.: Haupt- und Nebenfruchtformen der Ascomyceten, 395 pp. Il-
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Lieneman, Catherine : Host index of the North American species of the genus
Cercospora, A7in. Missouri Botan. Garden, 16(l):l-52. 1929.

LiNDAU, G.: Pyi'enomycetineae, in A. Engler und K. Prantl: Die Naturlichen
Pflanzenfamilien, Erster Teil, Abt. 1:321-491. Figs. 228-288. 1897.

Lindegren, Carl C: The genetics of Neurospora: I. The inheritance of response
to heat treatment. Bull. Torrey Botan. Club, 59(2):85-102. Figs. 1-4. 1932;
IL Segregation of sex factors in the asci of N. crassa, N. sitophila and N.'
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The inheritance of tan versus normal, Am. J. Botany, 21(2):55-65. PI. 1.
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32(2):243-256. PI. 14. Figs. 1-2. 1936a.

The structure of the sex-chromosome of Neurospora crassa, J. Heredity,

27{7) -.251-259. Figs. 1-4. 1936b
Lupo, Patsy: Stroma and formation of perithecia in Hypoxylon, Botan. Gaz

73(6) :486-495. PI. 18. Figs. 1-7. 1922.
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32(5) :652-666. Figs. 1-13. 1940.
Miller, Julian H.: Biologic Studies in the Sphaeriales, Mycologia 20(4) -187-

213, (6):305-339. Pis. 21-22, 35-38. Figs. 1-3. 1928.
: Studies in the development of two Myriangium species and the systematic

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306 CLASS ASCOMYCETEAE

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Figs. 1-5. 1918.
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509. 1 fig. 1943.

CLASS ASCOMYCETEAE: ERYSIPHALES,

ASPERGILLALES, MYRIANGIALES,

SACCHAROMYCETALES

AS WAS true for several other orders of the Class Ascomyceteae the
l\ investigations of the last three decades have necessitated numerous
changes in the limits of the orders to be taken up in this chapter. Even as
presented in this edition much must be recognized as only tentative, for a
great deal more very intensive study will be necessary before a satis-
factory classification can be attained.

Order Erysiphales (Perisporiales of most authors i). Lindau, in Engler
and Prantl (1897), placed in this order almost all of the plant-inhabiting
Ascomyceteae with external mycelium and with the perithecia also ex-
ternal. Ostioles are mostly lacking except in the Family Microthyriaceae.
The latter family has now been transferred to the Order Hemisphaeriales
(see Chapter 10) and the remaining two families have been increased to
five or six, mainly by the discovery of new forms and by the division
of the Perisporiaceae.

In general the Erysiphales carry on a parasitic, less often saprophytic,
existence on the surface of the host plant. Exceptions are found in the
species feeding on "honey dew" which occur wherever the latter accumu-
lates in sufficient quantity. As parasites the mycelium may not even
penetrate through the epidermis but in many cases the epidermal cells
of the host are penetrated by haustoria. The mycelium is septate and
branched, mostly with uninucleate cells. Rarely no mycelium is visible
except that making up the perithecium.

1 The Order Perisporiales and Family Perisporiaceae are based upon the genus
Perisporium. In view of the fact that the type species of this genus, P. gramineum Fr.,
has been shown not to belong to this order as customarily limited, it is necessary to
base the order upon a generally recognized genus whose connection with the order
is beyond doubt. Following the proposal of Gwynne-Vaughan the name Erysiphales
has been selected, based upon the genus Erysiphe.

307

308 CLASS ASCOMTCETEAE

Asexual reproduction is lacking or consists of the formation of single
conidia or the successive formation of conidia which may separate as fast
as formed or remain adherent in chains, the oldest conidium being the
terminal one. In the Capnodiaceae and Meliolaceae conidia may be
produced in pycnidia. The perithecia arise on the external mycelium or
may be partly surrounded by it except in a few cases where they are sub-
cuticular or subepidermal, becoming external through the rupture of the
cuticle or epidermis respectively. They do not possess an ostiole in the
Erysiphaceae but in the Capnodiaceae ostiolate and inostiolate species
occur, in some cases both in the same genus.

Following Theissen and Sydow (1917) this order may be divided into
five families as follows :

Erysiphaceae: external, mycelium white (cinnamon-yellow in Astomella), con-
idia hyaline, falling off singly or more often remaining attached in chains.
Perithecia without ostioles, external to the host, free or embedded in cottony
mycelium, external layer of peridium dark-colored, brittle at maturity, the
cells polygonal in outline, with various types of hyphal appendages. Asco-
spores hyaline (or yellow in one genus).

Meliolaceae: external mycelium colored, reticulately branched, the individual
cells cylindrical (not forming a moniliform hypha), sometimes with spines
and hyphopodia, not becoming slimy. Conidia mostly in ostiolate pycnidia.
Perithecia external (subepidermal in one genus), usually without true ostioles,
external cells polygonal, not becoming slimy. Perithecia without true ap-
pendages but often with bristles arising from an overlying layer of spiny
mycelium. Ascospores rarely one-celled, mostly two- to several-celled,
hyaline or more often brown.

Englerulaceae^: external mycelium mostly colored, parasitic on leaves or on
fungi on the latter. Chief distinction is the release of the asci from the peri-
thecia by the slimy histolysis of the latter.

Capnodiaceae: external mycelium of dark moniliform hyphae or of straight-
sided hyphae united laterally into dark sheets. In many cases saprophytic
on "honeydew." Conidia in elongated ostiolate pycnidia. Perithecia dark,
with or without ostioles, sometimes in the same genus. Walls of the often
stalked perithecia of rounded cells or of longitudinal hyphae united by slime.

Trichothyriaceae : mycelium dark-colored, creeping over the epiphyllous my-
celium of Meliola and other fungi. Perithecium wall of radial hyphae, origi-
nating at the tip of an upright hypha which then turns over so that the
morphological base of the perithecium is uppermost with the clusters of asci
at the upper end. Ascospores two- to several-celled, hyaline or colored.
Very doubtfully belonging in this order is the following:

Atichiaceae: forming small rounded or stellate cushions on leaves, horny when
dry, gelatinous when wet, with no free mycelium. Reproducing asexually by
clusters of cells (propagula). Perfect stage consists of asci scattered here
and there in one level in thickened areas of the thallus.

Arnaud (1925) does not recognize this order but distributes the
families here included among various other groups. Thus he places the

*See note on this family on p. 319.

OKDER ERYSIPHALES (PERISPORIALES OF MOST AUTHORS)

309

Erysiphaceae in what is called here the Order Hypocreales, Meliola and
Amazonia in the Dothideaceae, the Atichiaceae in the Myriangiales and
various Capnodiaceae in the Sphaeriales.

Family Erysiphaceae (The Powdery Mildea^s). These fungi are
parasitic upon Flowering Plants (Anthophyta) the world over, reaching
their greatest development in the temperate zones. They are usually con-
fined to leaves and young tissues of other portions of the plant, such as

Fig. 97. Erysiphales, Family Erysiphaceae. (A) Erysiphe graminis DC. Haustoria
in epidermal cells of host, seen from above. (B, C) Phyllactinia guttata (Wallr.) Lev.
Mycelium in substomatal chamber of host. (D) [Leveillula taurica (Lev.) Arnaud.
External and internal mycelium, and conidiophore. (After Arnaud: Ann. ^viphvt.,
7:1-115.) ^ '

310

CLASS ASCOMTCETEAE

the young shoots, the buds, fruits, etc. Sphaerotheca phytoptophila Kell. &
Swingle is found only on the lobed galls produced on the hackberry (Celtis
occidentalis L.) by a species of mite. Vncinula necator (Schw.) Burr,
attacks not only the leaves and young green shoots but also the immature
berries of the grape (Vitis) while Sphaerotheca mors-uvae (Schw.) B. & C,
on the gooseberry (Grossularia) is more often on the berries. The mycelium
is mainly superficial, obtaining its nourishment by haustoria penetrating
the epidermal cells or even to the cell layer immediately underneath. In
Phyllactinia part of the mycelium enters the stomata and sends its

Fig. 98. Erysiphales,
Family Erysiphaceae. Ery-
siphe graminis DC. Conidio-
phore and chain of conidia
arising from surface my-
celium. (After Salmon: Mem.
Torrey Botan. Club, 9:1-
292.)

haustoria into the mesophyll cells bordering the substomatal chambers.
In Leveillula the mycelium enters the leaf through the stomatal opening
and is confined to the mesophyll except that the conidiophores emerge
through the stomata. Foex (1912) pointed out that from this interior
mycehum there eventually creeps out through the stomata a thin myce-
lium growth which spreads out over the epidermis, held fast to it by
appressoria, but not producing any haustoria. This external mycelium
produces the perithecia and a few conidiophores bearing short chains of
small conidia which are smaller than those borne singly on the conidio-
phores emerging from the stomata. The mycelial cells are always uni-
nucleate. The conidia arise at the apex of short or elongated conidiophores.
In Leveillula and Phyllactinia the conidium falls off before the next suc-
ceeding conidium is formed, but in the remaining genera of the family the
conidia remain attached so that a chain of conidia is produced which in

OKDER ERYSIPHALES (PERISPORIALES OF MOST AUTHORS) 311

Erysiphe graminis DC. may consist of 20 or even more before the older
ones break off. Berlese (1898) reports for this species that after a cell is
cut off at the top of the conidiophore it divides into two conidia, the next
cell cut off from the conidiophore dividing similarly into two conidia. Thus
the terminal two conidia are of equal age and are the oldest pair, followed
by younger and younger pairs toward the base of the conidial chain. The
conidia are distributed by the wind and germinate on the epidermis of
the hosts, producing a short hypha which sends a haustorium into an
epidermal cell. Conidia are not described for the genus Astomella (Thiru-
malachar, 1947). (Figs. 97, 98.)

Brodie (1945) summarizes the experiments of himself and others upon
the germination of the conidia of Erysiphaceae at different degrees of
relative humidity. The conidia of Sphaerotheca pannosa (Wallr.) Lev. var.
rosae Wor. will not germinate at a relative humidity below 95 per cent. On
the contrary a considerable percentage of the conidia of Erysiphe graminis
DC, E. polygoni DC, and Microsphaera alni (DC.) Wint. germinate in
air entirely devoid of moisture, i.e., zero per cent relative humidity. This
accounts, at least in part, for the frequently observed phenomenon that
some powdery mildews appear abundantly in very dry weather. Possibly
also the dryness of the atmosphere keeps down the infection by Ciccin-
ohohts cesatii de By. and other fungi that are parasitic upon these mildews.
As long as the chains of conidia remain attached to their conidiophores
none of the conidia germinate but when detached the terminal conidia at
each end of the chain will germinate, but not those between.

After infection has taken place the mycelium grows rapidly, branching
in all directions, but gradually spreading radially. Conidial production
begins soon and may continue for some time but eventually gives way
to the production of perithecia. In some species of powdery mildews
conidia only are produced on some hosts, conidia and, later, perithecia
developing on other hosts. When perithecial development occurs it is
often on only one side of the leaf although conidia may be produced on
both sides. This is not universally true. Perithecial production is rare in
the tropics.

We owe our first clear knowledge of the sexual processes in this family
to Harper (1895 and later) although de Bary and others 20 to 30 years
earlier had described the external features. From neighboring hyphae in
contact there arise an antheridial and an oogonial branch, each at first
uninucleate. The latter is somewhat rounder and the former a little more
slender. They become appressed side by side or may even coil about each
other a little. The antheridial branch divides into a uninucleate stipe and
an apical uninucleate antherid which presses closely against the upper
portion of the oogone. The oogonial branch remains unicellular or may
divide into a smaller basal and a larger terminal cell (the oogone proper)

312 CLASS ASCOMYCETEAE

or the nucleus only may divide, producing a binucleate oogone one of
whose nuclei eventually disintegrates. An opening is formed from the
antherid into the oogone and through this the single male nucleus passes,
or the antherid nucleus divides and one nucleus passes into the oogone, the
other remaining in the antherid. According to Harper, Hein (1927), and
several other observers the male and female nuclei fuse in the oogone but
according to Dangeard (1907) and his students there is no passage of a
male nucleus into the oogone hence no nuclear fusion at this stage. Berg-
man (1941) reinvestigating the development in the species Sphaerotheca
castagnei Lev. confirms the passage of the antherid nucleus into the
oogone, where it passes by the larger oogone nucleus and takes up a basal
position. According to him there is no union of the nuclei at this stage. In
the meantime from the cell basal to the oogone there grow out branching
hyphae which push upward and around the oogone with the attached
antherid, thus forming a layer of 16 to 20 cells of which the antherid stalk
is one. An additional layer of hyphae arises in a similar manner from the
basal cell, within the first layer, and finally a third layer is produced. The
innermost layer consists of thin- walled cells, called by Hein "nurse cells,"
richly filled with food while the two outer layers form the cortex. This
eventually becomes dark and the cells become more or less polygonal in
outline. The zygote nucleus of the oogone divides rapidly according to
Harper and the oogone elongates, becoming an elongated plurinucleate
structure which is soon divided by septa into a row of from five to eight
cells, all uninucleate except the penultimate cell. Hein finds in his studies
that only three cells are produced, the middle one with two nuclei. Berg-
man describes the development in a different manner. The male and
female nuclei in the oogone do not unite immediately but each divides,
producing thus four nuclei in the somewhat elongated and curved oogone.
Two septa divide this into three cells of which the middle one is binu-
cleate. In Sphaerotheca and Podosphaera the two nuclei unite and the cell
enlarges and becomes the single ascus. The fusion nucleus divides suc-
cessively until eight nuclei are formed. Around each nucleus part of the
cytoplasm of the ascus is cut out by the formation of spore walls, thus
producing eight ascospores or, by degeneration of some of the nuclei or
partly formed spores, a smaller number. The ascospores are hyaline and
broadly ellipsoid. The ascus is obovoid. Miss Allen (1936) reports that in
Erysiphe polygoni DC. the formation of recognizable antherids is the
exception. Usually when any two suitable hyphae come in contact,
whether end to end or otherwise, the intervening walls dissolve and
nuclear transfer occurs. From the nearby hyphae the perithecium arises
containing a maze of interconnecting cells with varying numbers of
nuclei. Eventually the outer, colored cortex develops and several of the
contained cells enlarge to become the asci, two nuclei uniting in each.

ORDER ERYSIPHALES (PERISPORIALES OF MOST AUTHORS)

313

Fig. 99. Erysiphales, Family Erysiphaceae. Sphaerotheca castagnei Lev. Stages
in sexual reproduction. (A) Oogone and antheridial branch. (B) Antherid set off by a
septum. (C) Male nucleus has passed into oogone through an opening. (D) Row of
ascogonial cells, the penultimate one binucleate. (E) The nuclei of the penultimate
cell have united to form the primordium of the ascus. (F) Young perithecium showing
three layers of cells surrounding the young ascus whose nucleus has not yet divided.
(A-E, after Harper: Jahrb. wiss. Botan., 29(4):655-685. F, after Hem: Bull. Torrey
Botan. Club, 54(5):383-417.)

Already, therefore, it is apparent in this family as in other cases in the
Ascomyceteae that a substitution has occurred of uniting vegetative
hyphae for specially formed antherid and oogone. This tendency is one
that appears to become more pronounced in the higher groups of fungi
where (as in most Basidiomyceteae) no organs can be distinguished as
definite oogonial structures or in many cases as antherids. (Fig. 99.)

Harper's accounts indicate two chromosome reductions during the
nuclear divisions in the ascus, as would be necessary if there were nuclear
fusions both in the oogone and in the ascus. Dangeard, on the contrary,
who denies the occurrence of a nuclear fusion in the oogone, admits but
one reduction division in the ascus. In the other genera of the family the
binucleate penultimate cell of the row arising from the oogone undergoes
further division and produces a number of short ascogenous hyphae made
up of dicaryon cells. The terminal cell of each ascogenous hypha enlarges
and the nuclei fuse and undergo division, thus giving rise to an ascus with
ascospores. The latter develop as do those of Sphaerotheca. In a few genera
the ascospores are two- or four-celled (see Key to the genera of this family
on p. 353). The number of asci may vary from 5 to 8 in some species up
to 20 to 30 in others. As the asci enlarge the layer of nurse cells is gradually
destroyed, leaving only the usually two-layered cortex, (Fig. 100 A.)

314

CLASS ASCOMYCETEAE

Fig. 100. Erysiphales. (A) Family Erysiphaceae. Microsphaera quercina (Schw.)
Burr. Vertical section through an almost mature perithecium. At right and left are
the bases of two appendages. (B) Family Meliolaceae. Parodiopsis stevensii Arn.
Vertical section through almost mature perithecium. (Courtesy, Arnaud: Ann. sci.
nat. Botan., 7:643-723.)

From the outer cells of the cortex, mainly near the equatorial zone,
arise the characteristic appendages. These are simple and hypha-like in
Erysiphe, Leveillula, Sphaerotheca, Chilemyces, and Leucoconis, hooked or
spirally-coiled at the tip in Uncinula and Uncinulopsis, straight and once
or more dichotomously forked at the apex in Microsphaera, Podosphaera,
and Schistodes, or stiff and needle-like, with a bulb-like base, in Phyllac-
tinia. In some species the appendages are colorless, but often they are
colored basally. They do not seem to have the same function in all cases.
They may hold the perithecia fast to the mycelium or may push it up
above the surface of the mycelium, or may curve downward and pry the
perithecium loose, as in some species of Uncinula and in Phyllactinia.
The hooked or forked appendages would seem fitted for distribution by
insects, but that has not yet been demonstrated to be the normal means
of distribution. The upper half of the perithecium of Phyllactinia and of
Typhulochaeta bears short, penicillately branched mucilaginous cells.
These serve to fasten the perithecium, after its separation from the
mycelium, with its top side down, to objects with which it comes into
contact. Appendages are lacking in AstomeUa. (Fig. 101.)

When the ascospores are mature, which may not be until the following
spring, the asci and the inner cells of the perithecium absorb water and
swell until the perithecium is ruptured, at which time the asci also begin
to burst, discharging the enclosed ascospores with considerable force, or
when the perithecium bursts it may throw out the asci still containing
their spores.

Homma (1933) sowed a single conidium of Sphaerotheca fuliginea
(Schlecht.) Pollacci upon its host plant and on the resultant mycelium
were produced conidia and sexual organs. He therefore considered this

Fig. 101. Erysiphales, Family Erysiphaceae. Perithecia with various types of
appendages. (A-C) PhyllacHnia eleagni Linder. (A) Perithecial appendage, greatly
enlarged. (B) Perithecium with appendages. (C) Ascus with ascospores. (D) Sphaero-
theca humuli (DC.) Burr. Perithecia with appendages. (E) Microsphaera herheridis
(DC.) Lev. Perithecium with appendages. (F) Uncinula salicis (DC.) Wint. Peri-
thecium and appendages. (A-C, courtesy, Linder: Mycologia, 35(4):465-468. D-F,
after Tulasne, Selecta Fungorum Carpologia, Paris, Typographic Imp^riale.)

315

316 CLASS ASCOMYCETEAE

species to be "homothallic." Yarwood (1935) on the other hand found the
sunflower mildew to be "heterothalHc."

Salmon (1903, 1904a, b, c, 1905) in England, Reed (1916) in the United
States, and Hashioka (1938) in Japan, showed that some species are made
up of biological races which are confined to but a single host species or to
very closely related species. This is true of Erysiphe graminis, confined to
grasses (Poaceae) but in which the biological race on Poa will not infect
Bromus or Triticum and the strains on either of these genera will not
infect the other. Even for the genus Bromus there are some races of the
fungus that will attack certain species while other species are subject to
attack by other races. On the contrary Erysiphe ciclioracearum DC. is
very widespread in its host range and conidia from the fungus on one host
may infect many different hosts in families far apart systematically.

In his very excellent monograph of this family Salmon (1900) recog-
nized six genera with a total of 49 species and 11 varieties in the whole
world. His species limitation is much more conservative and broad than
that of most later mycologists. Blumer (1933) recognized 80 species
in Central Europe alone besides mentioning 50 extralimital species. It is
therefore perhaps safe to say that there are upward of 150 species in the
world when all regions have been carefully explored mycologically. It
must be noted that the known species of the family are mainly confined
to the extratropical parts of the earth. As noted before, in the tropics the
not too frequently noted specimens are almost without exception in the
conidial stage, perithecia rarely occurring.

In addition to the six genera recognized by Salmon in 1900 four more
have been added that like the others produce one-celled ascospores. The
author follows Theissen and Sydow (1917) in also including three more in
which the ascospores are two- or four-celled, the perithecia and mycelium
otherwise characteristic of the family. A key to these genera appears at
the close of this chapter.

Several species of Erysiphaceae are harmful parasites of cultivated
plants. Sphaerotheca mors-uvae (Schw.) B. & C, of relatively minor im-
portance as a parasite of the berries of the American species of gooseberry
(Grossularia) , has become very destructive when introduced into Europe
where the European species of gooseberry are exceedingly susceptible to
injury by it. The same thing is true of Uncinula necator (Schw.) Burr.,
which also is a minor enemy of the American species of grape (Vitis), but
when introduced into Europe proved very harmful to the susceptible
Vitis vinifera L. Podosphaera oxyacanthae (DC.) de Bary and P. leuco-
tricha (E. & E.) Salmon are sometimes harmful, especially to nursery
stock and young trees, to Prunus and Malus respectively, in America,
Europe, and other parts of the world. Sphaerotheca humuli (DC.) Burr, is
destructive to the hop {Humulus lupulus L.) wherever the host is grown

ORDER ERYSIPHALES (PERISPORIALES OF MOST AUTHORS) 317

in large plantings. Roses in Europe are frequently seriously injured by
S. pannosa (Wallr.) Lev., but according to Salmon the similar injury to
roses in America is more often due to another species. In America Micro-
sphaera alni (Wallr.) Salmon is exceedingly common as the cause of the
powdery mildew on the common lilac {Syringa vulgaris L.) but it almost
never occurs on that host in Europe. Erysiphe cichoracearum DC. is
probably the most widely distributed species of powdery mildews, attack-
ing hosts in the most varied families. It is not in general very destructive.
E. graminis DC. on the contrary is often very destructive to various
small grains in all parts of the world.

Family Meliolaceae (Perisporiaceae of most authors; see footnote on
p. 307). In this family, too, the mycelium is mostly superficial and spreads
in a network from the initial point of infection by conidium or ascospore.
The hyphae, in contrast to those of the Erysiphaceae, are usually dark in
color. They may form a dense crust or be more or less separate. They may
send haustoria into the epidermal cells or even into the next layer of cells
beneath or may, without producing haustoria, adhere closely to the epi-
dermis whose outer wall becomes more or less corroded with evidence of
some injury to the contents of the cell. In Meliola and some other genera
the creeping hyphae produce short opposite or alternate two-celled
branches, the terminal cell of which is enlarged, and rounded or angular.
These are the hyphopodia. They serve the double function of anchoring
the hypha in place and of producing a haustorium which penetrates the
cuticle to enter an epidermal cell of the host. The swollen cell of the pair
is homologous to the structure called the appressorium in many fungi. In
some genera of the order the hyphopodium may consist of only one cell.
In the genus Pampolysporium the mycelium and perithecia are sub-
epidermal, in Alina and Lasiobotrys subcuticular. In Stomatogene and
Piline the mycelium forms a sort of foot which enters through the stoma
into the substomatal chamber. Conidia produced singly on the superficial
mycelium are reported in one or two genera. In several genera ostiolate
pycnidia containing numerous conidia are the characteristic mode of
asexual reproduction. It has been shown that most of the reported cases
of conidia not in pycnidia are based upon fungi parasitic on the mycelium
of Meliolaceae, e.g., Arthrohotryum and Helminthosporium of each of
which Stevens (1918) has described several species.

The perithecia are typically without appendages, mostly black or
dark brown, without ostioles in the majority of genera. The asci arise in a
single layer or tuft in the base of the perithecial cavity. The ascospores
are usually four or eight in number. They are one-celled and hyaline in
one genus but in most cases are two- to many-celled or even muriform
and either hyaline or brown. In general the ascospores do not represent
types that would seem to be primitive. (Figs. 100 B, 102 A.)

CLASS ASCOMYCETEAE

Fig. 102. Erysiphales. (A, B) Family Meliolaceae. (A) Meliola corallina Mont.
Perithecium. (B) Irene echinata (Gaill.) Th. & Syd. Hypha with hyphopodia. (C)
Family Englerulaceae. Englerula effusa (Cke. & Mass.) Theiss. Perithecium with
wall dissolved into individual cells and a mass of shme. (D) Family Capnodiaceae.
Scorias spongiosa Schw. Perithecia and pycnidia. (A-B, after Engler and Prantl: Die
Natiirlichen Pfianzenfamilien, Leipzig, W. Engelmann. C, after Theissen and Sydow:
Ann. Mycolog., 15(6):389-491. D, after Ellis and Everhart: The North American
Pyrenomycetes.

The process of sexual reproduction has been worked out carefully in
Meliola circinans Earle by Graff (1932). Close to one another on nearby
hyphae there arise an ovoid uninucleate oogone with short stalk cell and
a slender somewhat spirally wound uninucleate antherid, also with a
short stalk. The two become appressed near their tips and an opening is
produced. The antherid nucleus disappears and what appears to be a
fusion nucleus is visible in the oogone, although the passage of the male
nucleus into the oogone and its fusion with the female nucleus was not
observed. Over the united antherid and oogone the surrounding vegeta-

ORDER ERYSIPHALES (PERISPORIALES OF MOST AUTHORS) 319

tive hyphae grow to produce a dark-colored, shield-like stroma. From the
stalk cells of the oogone and antherid hyphae grow out and form a
perithecium around these organs, under the stromatic shield. The
fertilized oogone elongates and divides into a number of uninucleate cells
of which two or three near the apex send out several branched ascogenous
hyphae which produce typical hooks which give rise to the asci. A few
paraphyses appear among the asci while from the upper part of the peri-
thecium arise periphyses which grow up together and pierce the stromatic
shield and spread apart to produce the ostiole. The eight nuclei produced
in each ascus in the usual manner are taken up by two's into the four-
developing ascospores. Two of these binucleate ascospores are destroyed
by the growth of the other two which eventually become five-celled, with
one nucleus in each cell except the middle cell which is binucleate. The
presence of a true ostiole lined by periphyses and the occurrence of
paraphyses would seem to suggest that perhaps this genus is more closely
related to the Sphaeriales.

In this family Theissen and Sydow (1917) distinguish 19 genera of
which Meliola and Irene with many hundreds of species each are very
abundant in the tropics. A few species occur even in temperate regions.
Stevens (1927, 1928) wrote a monograph of the genus Meliola which is
indispensable for the recognition of the species of this difficult genus. He
differs considerably from Theissen and Sydow in his interpretation of the
relationships of the genera centered about Meliola. In a subfamily
Meliolineae he includes Adinodothis, placed by those authors in Family
Polystomellaceae of the Hemisphaeriales, and Amazonia, assigned by
them to Family Microthyriaceae, of the same order. Both of these genera
have the same type of asci and ascospores. The perithecia of Amazonia
have been found to be complete, not incomplete below, and the spreading
mycelium has hyphopodia like those of Meliola and Irene. (Fig. 102 B.)

The fact that some genera of Family Capnodiaceae have much this
same type of superficial mycelium, with hyphopodia or bristles would also
indicate that the classification of the Meliolaceae and Capnodiaceae, as
well as of the Englerulaceae (see below) is rather artificial. Arnaud's
studies in this field (1925) suggest lines for a better arrangement when
more extensive investigations can be completed.

Family Englerulaceae. These are leaf parasites whose perithecial
cells dissolve into slime at maturity, exposing the enclosed asci. The 13 to
15 genera and 20 to 30 species are with few exceptions tropical. Petrak
(1928) after an extensive study o.'' most of the genera assigned by Theissen
to this family decided that it is a collection of heterogeneous forms placed
together on account of the one common character, the slimy dissolution
of the perithecia. Since this characteristic is known in other orders, e.g.,
Hemisphaeriales in some genera of Microthyriaceae and of Polystomel-

320 CLASS ASCOMYCETEAE

laceae; and in some Hypocreales, it does not seem to him to be of sufficient
importance to warrant the erection of a separate family. He distributes
the genera examined by him among the Microthyriaceae, Polystomel-
laceae, Molhsiaceae, Myriangiales, etc. The presence of hyphopodia in
some genera would suggest relationship to Meliola or related genera. Miss
Doidge (1942) included the genus Englerulaster in the genus Asterina in
the Microthyriaceae. (Fig. 102 C.)

Family Capnodiaceae. These are transferred by Arnaud (1925) to
the Sphaeriales but it seems best to follow Theissen and Sydow in retain-
ing them in the Erysiphales. It must be noted that the work of Miss
Fraser (1935) on the development of the ascocarp of Capnodium shows
that this starts as a stroma in which develops an archicarp with asco-
genous hyphae and asci. This would indicate that the relationship of this
genus lies with the Pseudosphaeriales. The investigations of Graff (1932)
on Meliola and of various authors on Erysiphaceae show that in those
fungi the structure is a true perithecium. Their relationship is probably
with the Sphaeriales. When further developmental studies are carried on
in other genera of the families tentatively placed in the Erysiphales a
better realization as to their true position may be obtained. The dark-
colored mycelium of the Capnodiaceae is usually superficial and in many
cases saprophytic on "honey dew," the sugary deposit forming on plant
parts from the droppings of aphids, scale insects, etc. This mycelium
sometimes forms a black papery layer that can be peeled off from the
underlying leaf. In a few genera hyphopodia are present. Miss Fraser
(1937) has made an extensive study of the physiology of the "sooty mold "
fungi, including their food requirements, and relations to temperature,
light, and to the presence of other fungi on the same leaf. The conidia are
borne in pycnidia of various shapes, sometimes elongated like a long
necked bottle. The external perithecial walls are formed of parallel, later-
ally adhering hyphae, not of polygonal cells as in the Erysiphaceae and
Meliolaceae. These component hyphae may be dematioid, i.e., monili-
form, or perisporioid, i.e., with parallel sides. In the same genus may be
found species with ostiolate perithecia and other species whose perithecia
lack ostioles. The perithecia may be sessile or more or less stalked, some-
times elongated like the pycnidia. The ascospores vary from colorless to
colored and from two- to many-celled. Theissen and Sydow recognize 25
or more genera and 50 or 60 species, mostly tropical or subtropical, but
some found in temperate regions. Capnodium salicinum Mont, occurs
upon willow {Salix) leaves and twigs in Europe. C. citri Berk. & Desm.
causes sooty mold on oranges, etc. wherever they are cultivated. A few
species of Limacinia occur on leaves of trees and shrubs in temperate re-
gions, even as far north as Germany and England. (Fig. 102 D.)

OKDER ERYSIPHALES (PERISPORIALES OF MOST AUTHORS)

321

Family Trichothyriaceae. This family of 5 or more genera and 20
or so species has been tossed about by mycologists since its members were
first studied. Mostly these forms have been assigned to the Erysiphales
or to the Hemisphaeriales, close to the Microthyriaceae. They are tropical,
parasitic on the epiphyllous mycelium of Meliola and on other fungi. It
was demonstrated by von Hohnel (1917) that the perithecia at maturity
lie inversely, with the original point of attachment outermost and the
asci radiating downward and outward from this point toward the onto-
genetically upper side. The perithecium arises terminally on a stout up-
right hypha which curves over as the development progresses. Eventually

Fig. 103. Erysiphales, Family Atichiaceae. Atichia millardeti Racib. (= Seuratia
coffeicola Pat.). (A) Thallus when wet, showing ascigerous cushions. (B) Section
through portion of an ascigerous cushion. (C) Propagula. (A-B, courtesy, Arnaud:
Ann. sci. nat. Botan., 7:643-723. C, courtesy, Mangin and Patouillard, Compt. rend.,
154(23) :1475-1481.)

322 CLASS ASCOMYCETEAE

on the apparent!}^ upper side the tissues break away forming an ostiolar
opening. The outer perithecial wall is radial in its structure, resembling
many of the Hemisphaeriales in that particular. The ascospores are two-
to several-celled and colorless or brown. Trichothyrium is the first de-
scribed and best known genus.

Family Atichiaceae. Because of their epiphyllous habit, apparently
entirely external to the leaf tissues and perhaps feeding saprophytically
on honeydew or parasitically on hyphae of other fungi this family may be
placed here, but with great doubt. There are no separate hyphae but a
gelatinous mass of anastomosing threads forming a cushion or a stellate
thallus. The cell walls are very much swollen so that when moist the
organism is gelatinous, when dry horny. The outer layer of cells has
dark-colored outer walls. In separate pockets (Phycopsis) or clustered in
basket-like structures (Atichia) certain cells divide and form a mulberry-
like or tetrahedral cluster of adherent cells (propagula) which eventually
are pushed out by the pressure of the underlying propagula or vegetative
tissues and which serve to establish new plants. These propagula are
dark-colored externally, hyaline within. In one species spermogonial
structures are known, but their function has not been determined. The
asci arise in more or less thickened cushion-like areas of the thallus. They
lie separate from one another in the tissue of the thallus, arising possibly
from ascogenous hyphae in among the other vegetative hyphae. As they
mature they elongate and push through the surface and discharge their
eight, two-celled, hyaline or slightly colored spores. The occurrence of asci
scattered in the tissues of the thallus led to the suggestion by Raciborski
(1909) that they are related to the Myriangiaceae, while on the contrary
von Hohnel (1910) placed them in the Saccharomj^cetaceae. Mangin and
Patouillard (1912), Cotton (1914) and Arnaud (1925) have also given
attention to these fungi. (Fig. 103.)

Order Aspergillales (Plectascales). This group is perhaps heterogene-
ous as regards certain of the included families. It shows similarities to the
Erysiphales and to some of the Sphaeriales and Hypocreales in its mode of
sexual reproduction as well as in the conidial formation. The chief differ-
ences lie in the internal structure of the perithecium. In the groups just
mentioned the ascogenous hyphae are of about the same length and arise
from one or more centers from which they radiate, resulting in the forma-
tion of a tuft or tufts of asci in the cavity of the perithecium or stroma, or
of a hymenium at the base and sides. The thin-walled cells making up the
interior portion of the perithecium or stroma give way before the out-
growing asci and eventually disappear completely or nearly so or the asci
and paraphyses grow out into the perithecial cavity. In the Aspergillales,
on the contrary, the ascogenous hyphae are of varying lengths so that
instead of arising in a tuft the asci are produced scattered throughout the

OEDER ASPERGILLALES (PLECTASCALES) 323

interior of the perithecium. The latter, as in the Erysiphales and Sphae-
riales, consists of a firmer exterior cortex (this is sometimes of loose
hyphae) and a thin-walled interior portion. Some of these thin-walled
cells are pushed aside or destroyed as the branching ascogenous hyphae
grow among them so that eventually the asci appear to be imbedded here
and there in the internal ''nucleus" of the perithecium. Finally most of
these interior cells as well as the ascus walls and the remains of the asco-
genous hyphae are dissolved, leaving the ascospores loose in the peri-
thecial cavity. True ostioles are lacking in most genera of this order. In
the Aspergillaceae they are present in Microascus and Emericella. If the
suggestion of Nannfeldt (1932) is followed and the Ophiostomataceae and
Chaetomiaceae are transferred from the Sphaeriales to this order these
families will add to the ostiolate forms in the order. The asci are formed
at the ends of the ascogenous hyphae, sometimes by the hook method, or
in chains by the transformation of successive dicaryon cells of the hypha
into asci.

Conidial formation is frequently catenulate, with the apical cell the
oldest, as in the Erysiphaceae. Many species occur mostly in the asexual
stage, only very rarely producing perithecia. This is especially true of the
very numerous species of Penicillium and Aspergillus which form some of
the commonest molds upon organic matter of every kind. The details of
sexual reproduction still remain to be studied for the great majority of
genera. Even in those cases that have been studied much still remains to
be learned, particularly regarding the behavior of the sexual nuclei. In
general a straight or coiled ascogonium and coiled antherid are produced.
The former may be several-celled, the terminal perhaps corresponding to
a trichogyne, or only one-celled. In a few cases the formation of an opening
has been observed between the antherid and the tip of the oogone (or of
the trichogyne), following which ascogenous hyphae grow out of one or
more of the ascogonial cells. The eminent French mycologist P. A. Dange-
ard (1907) has observed and figured the antherid and ascogonium in nu-
merous species of this order. He even figured the opening from the antherid
into the ascogonium or trichogyne in a few cases. He denied, however, that
this is a sexual process and considered that the antherid has entirely lost
its primary function as a male organ, perhaps functioning now as a nutri-
tive organ, which he therefore calls a trophogone. In the main the my-
cologists, apart from Dangeard's students and associates, do not agree
with him and look upon the antherid as a functional sexual organ, at least
in the majority of cases where it is present. The actual observation by
Schikorra (1909) and by Young (1931) of the passage of nuclei from
antherid to ascogonium in Monascus shows that this does occur. On the
other hand there are well authenticated cases where a coiled or straight
ascogonium is produced and no organ that can be in any way interpreted

324

CLASS ASCOMYCETEAE

Fig. 104. Aspergillales, Family Aspergillaceae. Monascus ruber van Tiegh. Sexual
reproduction. (A) Ascogone growing up sympodially at base of elongated antherid.
(B) Ascogone separated by septum from hypha. (C) Ascogone divided into a basally
located oogone and a trichogyne which has formed an opening to the antherid. (D)
Antherid has collapsed and the nuclei have passed through the trichogyne into the
oogone. (E) Two investing hyphae are growing up from just below the oogone. A
conidium is shown on the antheridial branch. (F) Perithecium wall completed; one
of the curved ascogenous hyphae is shown with three terminal asci, successively
younger from apex toward base. (G) Section of mature perithecium with ascus walls
mostly dissolved. (Courtesy, Young: Am. J. Botany, 18(7):499-517.)

as an antherid. From this ascogoniiim arise binucleate cells which give
rise to ascogenous hyphae. (Fig. 104.)

The formation of the ascus has been reported by Schikorra to take
place by the hook method in Monascus but this is denied by Young.
DeLamater (1937) has demonstrated the formation of croziers in Arachni-
otus aureus (Eidam) Schroet., of Family Gymnoascaceae. In general one
or more of the binucleate cells at the end of an ascogenous hypha enlarge,
the nuclei fuse and the young ascus is initiated. The perithecium is formed
by the growth of hyphae from near the point of attachment of the
ascogonium.

In perhaps the majority of species investigated cytologically the
mycelial cells and the conidia are plurinucleate. The young ascogonium
and young antherid may be plurinucleate or uninucleate. It is difficult to
determine which represents the more primitive condition.

Several families are recognized in this order, following Fischer (1896)
in the main. Those first to be considered show the closest affinity to the
Erysiphales and may well have arisen from or have given rise to that
order. The sexual organs and mode of origin of the perithecium are quite
similar in some of the Aspergillales to the corresponding structures of the
Erysiphaceae. If the short ascogenous hyphae of the latter should become
longer and of various lengths, pushing in among the tissues of the central

ORDER ASPERGILLALES (PLECTASCALES) 325

portion of the perithecium we would have the structure as it is found in
the Aspergillaceae. If the central and cortical tissues of the perithecium of
the latter instead of forming a continuous structure should remain more
or less loose and cottony we would have the Gymnoascaceae. If the peri-
thecium should become much enlarged (up to several centimeters) with a
firm cortex several layers of cells thick and a more or less permanent mass
of loose central tissue traversed by more or less well-developed sterile
"veins," in which the asci are scattered, the structure would be that of
the Elaphomycetaceae. In the Onygenaceae the ascocarp is differentiated
into a basal sterile portion and a somewhat larger head in which the asci
are scattered in a central mass of tissue which eventually breaks up into
a sort of capillitium. The Trichocomaceae are forms whose development
is not well understood and whose relationship to other families of the order
is more or less a matter of doubt. The Terfeziaceae, placed by Fischer
(1896) in this order were more recently (1938) placed by him in the
Tuberales. Myriangium and probably several other genera rather similar
in structure, forming the Family Myriangiaceae, possibly also belong in
this neighborhood but are better placed in a distinct order.

Family Aspergillaceae. The chief genera of this family are the
ubiquitous molds, PenicilUum and Aspergillus, of which the commonest
species are the blue and green molds found on all sorts of organic matter.
Most of their species are saprophytes but a few are animal parasites, caus-
ing cases of mycosis. The small orange or yellow perithecia of a species of
Aspergillus (perhaps more properly called Eurotium) are very frequently
found on jams and various conserves, as well as on imperfectly dried
herbarium specimens where it is called ''herbarium mold." Usually when
examined these are found to consist of nothing but a thin peridium sur-
rounding a cavity in which are found a large number of ascospores shaped
much like a pulley wheel. Rarely a few asci will be found but usually their
thin walls have completely dissolved away. In PenicilUum perithecia are
much more rarely formed and are more often sclerotium-like. The two
genera are best distinguished by their conidiophores. In Aspergillus the
conidiophore is swollen at its apex into a head from which radiate numer-
ous short sterigmata bearing at the apex of each a chain of spherical or
nearly spherical, smooth or roughened conidia, or sometimes producing
secondary sterigmata which bear in their turn the chains of conidia. The
species with the secondary sterigmata were formerly set apart in the
genus Sterigmatocystis but the modern usage is to include these in Asper-
gillus. In PenicilUum the conidiophore branches several times without
enlarging, the terminal portions lying more or less parallel, or at least not
widely divergent, and bearing one to three or four sterigmata with chains
of conidia. Thorn (1914) described in detail the method of formation of
the conidia of this genus. From the apex of the sterigma is produced a

326

GLASS ASCOMYCETEAE

Fig. 105. Aspergillales, Family Aspergillaceae. Aspergillus (Evrotium) sp. Peri-
thecia, ascus and ascospores. (After Ellis and Everhart : The North American Pyreno-
mycetes.)

narrow tubular extension near the tip of which a nucleus, formed by the
division of a nucleus in the sterigma, takes its position. A delicate cross
wall is then formed and the basal portion of the tube elongates and forms
a second cell in the same manner. This process continues until a chain of
cells is formed, the oldest at the tip. Within the segments of this tubular
extension the spores round up and each secretes a thicker wall within and
separate from the wall of the tube to which it may adhere or from which
it may be free. Thus the conidia are in reality formed internally and are
on that account called "endogenous" by Sartory and Sydow (1913).
The thin-walled portions of the original tube, lying between the conidia,
are often spoken of as "connectives." (Figs. 105, 194.)

In recent years the genus Penicillium, especially P. notaium Westling
and other species of this and related genera have acquired great impor-
tance because of the discovery that some of the products of the growth of
the mycelium are of great value as antibiotics. These are substances
which are able to bring about the destruction of various other organisms,
harmful or otherwise, even when introduced into the living bodies of man
and other animals infested by these organisms.

CitromyceH is much like Penicillium but the conidiophores or their
branches are somewhat thickened. The chief distinction is a biological
one, viz., the production of large quantities of citric acid when grown on
a medium containing sugar. Monascus is usually distinguished by its red
or pink mycelium. The conidia are on short chains. The mature peri-

ORDER ASPERGILLALES (PLECTASCALES) 327

thecium consists of one or two layers of cortical cells around the cavity-
enclosing numerous ellipsoid or globose ascospores, the interior perithecial
tissues and the walls of the eight-spored asci having dissolved rather early.
This was at first interpreted as a single many-spored ascus, hence the
name Monascus (meaning one ascus). One species is the "pink mold"
found in ensilage that was put into the silo in too dry condition. Other
species are used in the Orient for the fermentation of rice in the prepara-
tion of alcoholic beverages. (Fig. 104.)

The genus Lilliputia was established by Boudier and Patouillard
(1900) for a fungus shown much later by Dennis and Wakefield (1946) to
be a member of this family and in which the conidial stage belongs to the
form genus Gliodadium. This resembles closely Penicilliu in but the conidia
are surrounded by a slimy exudate so that the branched "brush" at the
upper end of the main stalk of the conidiophore is involved in a glijstening
drop filled with hundreds of spores which early lose connection with one
another so that they appear to be single or at most only in short chains.
The perithecia are relatively large for the family, from 0.5 to 1.0 mm. in
diameter, with a thick firm cortical region and round to ovoid asci, each
containing eight large spherical, yellowish to brown ascospores which are
rough or prickly. The most common species is L. insignis (Wint.) Dennis
and Wakefield (1946) based on Eurotium insigne Wint. This occurs in
Europe and in the United States on various types of organic substrata,
such as dung of geese and kangaroos, dead seaweeds along the shore, and
old stable manure. Brefeld (1908) gave the name Lysipenicillnim to this
genus but it is antedated by Lilliputia. It must be noted that the asci and
ascospores are very similar to those of Terfezia to which Boudier and
Patouillard considered them closely related. This casts doubt on the cor-
rectness of transferring the Terfeziaceae to Order Tuberales. Magnusia
produces depressed globose -oblong, dark-colored perithecia with elon-
gated, apically circinate appendages arising from near the base. (Fig.
106 A.)

Sexual reproduction has been studied in several members of this
family. Both Schikorra (1909) and Miss Young (1931) have studied this
process in Monascus. A slender plurinucleate antherid is produced at the
end of a hypha. Sympodially from the cell below arises the ascogonial
branch which bends so as to lie parallel to or to coil somewhat around the
antherid. A basal cell cuts this off from the main hypha and soon an
apical trichogyne cell is set off by another septum. The trichogyne and
oogone both contain several nuclei. An opening appears between the
trichogyne and antherid through which the nuclei of the latter pass, the
trichogyne nuclei having previously disappeared. The septum between
trichogyne and oogone dissolves out and the male nuclei pass into the
latter, the septum then being regenerated. The nuclei pair by twos in the

328 CLASS ASCOMYCETEAE

oogone but fusion of these pairs has not been observed. In the meantime
hyphal branches arising from just below the oogone form the beginning
of the perithecial wall which is one or two layers of cells in thickness. From
the base of the fertilized and enlarged oogone grow several ascogenous
hyphae, each of several dicaryon cells. The apical and second and third
cells may develop into asci, each with eight ascospores. Ascus walls and
the remainder of the ascogenous hyphae and of the oogone and antherid
dissolve and leave the ascospores free in the perithecial cavity. De Bary
(1870) and many years later Miss Dale (1909) reported that in Aspergillus
glaucus Lk. (= Eurotium herhariorum Lk.) a tightly coiled ascogonium
is formed, at first one-celled but soon dividing into several multinuclear
cells. An antherid rising from lower down or from another branch creeps
up the side of the coiled ascogonium. A fusion of antherid and ascogonium
was not observed and possibly does not occur. It is not certain that this
is really an antherid. Possibly it is one of the investing hyphae that grow
up from below the ascogonium to form the perithecial wall. Soon the
middle cells of the ascogonium divide into binucleate cells from which
arise the branched ascogenous hyphae. From the cells supporting the asco-
gonium arise the hyphae which form the perithecium with a cortex of one
layer of polygonal cells the interior being filled with thin-walled cells. In
the species called by Brefeld (1874) Penicillium crustaceum Ft., this
mycologist reports the formation of short coiled antherid and ascogonium
from adjacent cells of the same hypha. These fuse at the apex and give
rise to a several-celled structure from which branch out the ascogenous
hyphae while around them a dense sclerotium-like perithecium develops.
The nuclear behavior has not been followed. Dangeard (1907) reports in
P. vermiculatum Dang, that there is formed a long straight multinucleate
ascogonium around which coils a slender antheridial hypha with an en-
larged uninucleate antherid at the apex. This fuses with the ascogonium
and the male nucleus may pass into the latter or may remain in the
antherid. The ascogonium then divides into binucleate cells from which
arise the ascogenous hyphae. Dangeard reports that the male nucleus is
not functional. Derx (1925) reported that the ascospores in P, luteum
Zukal produced mycelia of two sexual phases that are mutually compat-
ible and self-sterile.

Family Gymnoascaceae. This family is characterized by the lack of
a firm-walled perithecium, this })eing represented by a more or less loosely
tangled mass of mostly branched hyphae among the bases of which the
asci are clustered. When mature they are usually visible through this
tangle of hyphae. The chief generic distinctions are based upon the
character of these interwoven filaments and the color of the ascospores.

The ascocarps are mostly not over 1-2 mm. in diameter, often smaller,
and sessile, globose or depressed globose, almost colorless or dark-colored.

ORDER ASPERGILLALES (PLECTASCALES)

329

Fig. 106. Aspergillales. (A) Family Aspergillaceae. Magnusia nitida Sacc. Peri-
thecium. (B-D) Family Gymnoascaceae. Arachniotus trisporus Hotson. (B) Asco-
gonial branch coiled around the antherid. (C) Ascogenous branch with asci. (D)
Mature perithecium with cortex of loosely woven hyphae surrounding the asci.
(A, courtesy, Ames: Mycologia, 29(2):222-225. B-D, courtesy, Hotson: Mycologia,
28(6):497-502.)

In some cases the color of the ascospores gives the color to the ascocarp.
Conidia are formed in some species, singly on short conidiophores or in
chains. Chlamydospores also are known in some species. These fungi are
often saprophytic on animal matter such as feathers, dead animal bodies,
excrement, etc. Arachniotus trachyspermus Shear (1902) was found asso-
ciated with diseased cranberries and A. trisporus Hotson (1936) was iso-
lated from contaminated milk. A number of skin parasites of man and
other animals have been credited to this family, but the evidence is not
too convincing.

The details of sexual reproduction have been described by several
authors. In Ctenomyces, Eidam (1880) described a multinucleate asco-
gonium coiled around the straight multinucleate antherid. Passage of
nuclei from the latter to the former was observed. The ascogonium soon
divides into numerous short binucleate cells each of which produces a
more or less coiled ascogenous hypha. During this time the near-by hy-
phae have formed a loosely woven perithecial wall around the developing
ascogenous hyphae. In Gymnoascus reessii Baran. the union of antherid
to the tip of the coiled ascogonium was observed by Baranetzky in 1872.

330 CLASS ASCOMYCETEAE

In 1903 Miss Dale observed the passage into the ascogonium of several
male nuclei. The ascogonium divides into a number of cells each of which
produces ascogenous hyphae. The perithecial wall in this genus is a mass
of loosely interwoven hyphae with spines and prongs. Hotson (1936) ob-
served similar reproductive structures in Arachniotus trisporus Hots., as
did DeLamater (1937) in a species close to A. aureus (Eidam) Schrot. The
latter demonstrated the passage of a nucleus from the antherid into the
ascogonium coiled about it. The nuclei then multiply by conjugate divi-
sion and the ascogonium divides into binucleate cells from each of which
a crozier buds out to give rise to an ascus. The uninucleate tip and basal
cells of the crozier often unite and produce another crozier. Nuclear fusion
occurs in the young asci and eight ascospores are formed. Fifty single-
spore cultures were made and all were fertile showing that this species is
self-compatible. (Fig. 106 B-D.)

Family Elaphomycetaceae. These fungi are subterranean and prob-
ably saprophytic although possibly they may produce mycorrhiza. The
ascocarps are large, up to 2 or 3 cm. in diameter with a very thick perid-
ium, usually hard and roughened externally, the central portion consisting
of the ascogenous hyphae and asci and the thin-walled central cells of the
ascocarp, traversed radially by the "veins" which probably are conduc-
tors of foodstuffs. The cell walls of the central portion dissolve leaving the
numerous ascospores free in the center of the ascocarp. Clemencet (1932)
reported that in Elaphomyces no sexual organs are to be found but that in
Ascoscleroderma (a segregate of that genus) a stout ascogonial filament
coils around a straight antherid which however is sexually functionless.
From the ascogonium arise branching ascogenous hyphae which produce
at their tips rectascous asci, i.e., the cell elongates and the nuclei fuse
without any curving or hook formation. In Elaphomyces on the contrary
the asci are subterminal and, judging by the illustrations, are formed by
means of croziers. C. W. Dodge (1929) recognized 24 species of Elapho-
myces, mainly from Central and Southern Europe and the United States.
One species is reported from Italy, France, and Australia.

Family Onygenaceae. This family contains a few species in the
single genus Onygena, found mostly in temperate Europe and North
America. They grow on old feathers, hair, hoofs, horn, felt, and other
animal matter. The ascocarps are a few millimeters up to 1 or 2 cm. tall,
consisting of a stalk and a somewhat enlarged head within which the asci
are scattered in the manner characteristic of this order. The tissues break
up into a sort of capillitium. (Fig. 107.)

Family Trichocomaceae. In this family consisting of the one genus
Trichocoma, and only a few species, the ascocarp is sessile. When young
it is closed with a firm peridium which is thinner above. The ascogenous
portion develops centrally and consists of irregularly scattered asci with

ORDER MYRIANGIALES

331

vertical sterile plates. The latter grow at the base so that at maturity a col-
umn of vertical honeycomb structure bursts the upper peridium and pushes
upward. In the meshes of the honeycomb there are developing asci at the
base while near the top the ascus walls are dissolved and the ascospores
lie free. As the upper ends of the column weather away the ascospores are
released. Tropical and warm temperate species. Developmental studies
of the earlier stages are lacking, in default of which the exact relationship
of this family is uncertain.

Fig. 107. Aspergillales, Family Onygenaceae. (A-
C) Onygena equina Pers. ex Fr. (A) Habit sketch. (B)
Vertical section. (C) Asci. (D) Ascospores of Onygena
caprina Fckl. (After Engler and Prantl: Die Natiir-
lichen Pflanzenfamilien, Leipzig, W. Engelmann.)

Family Terfeziaceae. Although the fungi of this family with their
large subterranean ascocarps resemble in structure in many ways some of
the genera of the order Aspergillales, and especially in ascus and ascospore
structure those organs in Lilliputia, the author follows Fischer (1938) and
Miss Gilkey (1939) in placing them in the Tuberales, but with consider-
able doubt as to the correctness of this transfer.

Order Myriangiales. This order, in the first edition of this textbook
considered to be closely related to the Pseudosphaeriales, appears, from
the studies of Julian H. Miller (1938) discussed below, to be more nearly
related to the Aspergillales. Some of the earlier mycologists suggested a

332 CLASS ASCOMYCETEAE

close relationship to the Pezizales. Whether the order should be con-
sidered to consist of but one family or of several depends upon the weight
given to the arrangement of the asci in the spore fruit, i.e., in a single layer
or scattered at various depths below the outer surface, and to the septa-
tion and color of the ascospores. Until life-history studies like those by
Miller have been completed on the other genera of this order it seems
advisable for the present to include all in one family.

Family Myriangiaceae. Julian Miller (1938) described in detail the
development of the ascocarps of two species of Myriangium and con-
cluded that this genus, and the other genera usually associated with
it in this family, are not properly ''Pyrenomycetes" but on the con-
trary should be associated with the Aspergillales. The species of this
genus are parasitic upon scale insects which in their turn are feeding upon
many kinds of trees and shrubs in the tropical and warm temperate re-
gions of the world. Growing out from the body of the host the mycelium
forms a firm cushion-like stroma which is rather strongly anchored to the
bark of the host by hyphae that may penetrate the lenticels. That there
may be some direct parasitism upon the woody host is suggested by the
fact that the bark cells underneath the center of the stroma die. Further-
more a stroma may remain alive and produce ascocarps for several years,
long after the scale insect has been killed by the fungus. At the central
thicker portion of the stroma arise the archicarps just under the upper
surface. These are upright, more or less coiled, hyphae of uninucleate
cells (as are all of the cells of the stroma), the upper cell being attenuated
and one or more of the middle or lower cells enlarged and soon multi-
nucleate. Accompanying the archicarp is a more or less loosely coiled
hypha of slender uninucleate to plurinucleate cells, apparently the an-
theridial hypha. However, no connection between antherid and archicarp
was observed. No spermatia were produced anywhere in the stroma. The
enlarged ascogonial cells divide repeatedly producing a layer of large
multinucleate cells. From the apex of each such cell arises an ascogenous
hypha of binucleate cells. Thus is produced a flat or concave disk-like
thickening on the stroma consisting entirely of parallel, closely packed
ascogenous hyphae. These are more or less branched. Terminal and inter-
calary cells of these hyphae divide longitudinally or diagonally. The two
nuclei of one of these daughter cells divide again and two septa are formed,
producing a row of three cells, the central one binucleate. These two nuclei
unite and the cell enlarges to become the ascus. Since many cells in each
ascogenous hypha may produce asci and these hyphae are packed side by
side to form the disk of the ascocarp the resultant asci are scattered at
various depths below the upper surface but are separated by the tissues
consisting almost entirely of ascogenous hyphae. There is no dissolution
to form monascous cavities such as is found in the Pseudosphaeriales.

ORDER MYRIANGIALES

333

In ilf . duriaei Mont. & Berk, only one archicarp is formed in the young
ascocarp, the whole disk being the product of the divisions of its asco-
gonial cells and of the branching ascogenous hyphae arising from them.
In M. curtisii Mont. & Berk. 30 to 50 archicarps are produced and the
resultant ascogonia and ascogenous hyphae do not form as compact and
extensive a structure as in the other species. (Fig. 108 A-C.)

The asci when mature have a two-layered wall. When the ascocarp is
wet by rain it becomes soft and the mature asci expand, pushing their

Fig. 108. Myriangiales, Family Myriangiaceae.
(A-C) Myriangium duriaei Mont. & Berk. (A)
Stroma, top view; the cuplike bodies are asco-
genous. (B) Ascus. (C) Dehiscing ascus, the outer
wall ruptured and the inner wall elongated but
still intact. (D, E) Elsinoe veneta (Speg.) Jenkins.
(D) Vertical section of ascocarp. (E) Acervulus
(Sphaceloma) stage. (A-C, after Fetch: Brit.
Mycol. Soc. Trans., 9:45-80. D-E, after Burk-
holder: Cornell Agr. Expt. Sta. Bull., 395:155-183.)

tips through to the surface. The outer ascus wall splits and contracts
around the base of the ascus while the thin-walled inner layer pushes out
further and discharges its eight greenish yellow muriform ascospores a
distance of several centimeters. The lower asci push up through the spaces
left by the disappearance of the upper asci and repeat the process. There
is no weathering away of the tissues as previously believed. The ascocarps
of the other genera usually placed in the same family with Myriangium
have not been given sufficiently intensive ontogenetic study to make it
certain that they have the same type of development although that prob-
ably is so. Apparently the asci in these other genera are of the same type
as those o" Myriangium.

334 CLASS ASCOMYCETEAE

Elsinoe consists of a good many species of plant parasites which pro-
duce their stromata under or within the epidermis. By the rupture of the
latter the stroma containing the scattered asci becomes exposed. Unlike
Myriangium, in which no conidia have been observed, there are produced
acervuli or sporodochia bearing unicellular, hyaline conidia, of the form
genus Sphaceloma. E. veneta (Speg.) Jenkins is the cause of anthracnose
of Rubus, E. ampelina (de Bary) Shear of anthracnose of Vitis, E. piri
(Wor.) Jenkins of anthracnose of pear (Pirus) and apple {Malus). Other
species of Elsinoe cause serious diseases of species of Citrus, Canavalia,
Phaseolus, etc. The ascospores are in most cases septate transversely but
in a few cases longitudinal septa also are present. Including those so far
known only in the characteristic Sphaceloma stage about 175 species of
Elsinoe are recognized in the whole world (Jenkins, 1947) of which above
40 occur in the continental United States and the island possessions,
Puerto Rico, Hawaii, and Guam. The diseases produced are in the later
literature called "spot anthracnoses." Other genera are reported mainly
from the tropics, some parasitic in plants, some upon insects. (Fig. 108
D, E.)

The relationship of this family to other Aspergillales or its justification
as a separate but closely related order is uncertain. In some particulars it
reminds one of the Trichocomaceae, and more distantly of the Elapho-
mycetaceae. Raciborski (1909) has suggested the possible relationship of
the Atichiaceae to the Myriangiaceae, but the structure and development
of the ascocarp appears to be too fundamentally different in the two
families in so far as the meager information is available on these points in
the former family. The possible relationship of the Myriangiaceae to the
Pseudosphaeriales is discussed in Chapter 17.

Order Saccharomycetales. This order represents, in the opinion of the
author, the ultimate degree of simplification in the Class Ascomyceteae.
On the other hand it must be noted that by many eminent mycologists,
such as de Bary (1884), Brefeld (1889), Gaumann (1926), Dangeard
(1907), Atkinson (1915), and others, the members of this order are looked
upon as primitively simple, representing almost the first steps in the evo-
lution of the Ascomyceteae from the Phycomyceteae. To agree with this
theory the arrangement of the orders of this class should be the reverse of
that used in this book, placing the Saccharomycetales first and the
Pezizales and their allies last.

The fungi making up this order are filamentous or unicellular (Yeasts).
In the latter case the cells often remain attached until several cell divi-
sions have occurred forming an irregular mass but usually not a cylin-
drical hypha. The fundamental common character of all the members of
the order is the production of single asci, usually as the result of a sexual
process but sometimes parthenogenetically, instead of a cluster of asci

ORDER SACCHAROMYCETALES 335

from branched ascogenous hyphae. Furthermore there is lacking the
formation of a protective structure, the perithecium or apothecium.

The mycelial forms of this order mostly take on the unicellular (i.e.,
yeast) habit when growing in nutritive media of considerable concentra-
tion. On the other hand many of the yeasts can be made to develop more
or less typical hyphae when grown on special media, usually those not too
rich in soluble organic substances. In the main the species that have the
yeast habit possess uninuclear cells. That is also true of many of the
filamentous forms, although some of the latter may have several nuclei to
the cell.

The majority of Saccharomycetales are saprophytes on organic mat-
ter, mostly of vegetable origin. However, some are parasites in plants
(e.g., species of Nematospora) or in Man or other animals. When growing
as saprophytes in organic media of plant origin they frequently cause fer-
mentation, the products usually being CO2 and various alcohols or organic
acids. Some species of Saccharomyces are of very great importance, espe-
cially in the commercial production of alcohol for industrial or beverage
purposes, or of CO2 for leavening bread. Many of these fungi can grow
anaerobically if sufficient soluble organic foods are present in the medium.

As in most of the Ascomyceteae in which such studies have been made
the cell walls are mostly lacking in cellulose but it is not agreed whether
or not chitin is present. In Schizosaccharomyces octosporus Beij. and Sch.
vcrsatilis Wick. & Duprat (1945) the ascus walls give a distinct blue reac-
tion with iodine-containing media, indicating the presence of carbo-
hydrates closely related to dextrin or starch. This should be compared
with the similar staining of the apical portions of the asci of some Pezi-
zales and of the b^-sidiospore walls of some Agaricales.

In the filamentous forms asexual reproduction may take place by ter-
minal or lateral budding from the hyphal cells or by the breaking up of
the hyphae into short unicellular pieces, often called "oidia." Short
conidiophores may be produced in some species which bear conidia singly
or in short chains. The unicellular forms may reproduce by fission into
equal cells which remain adherent at first and eventually separate from
one another. These are the Fission Yeasts. The majority of the yeasts
however reproduce by the formation of small buds which enlarge until
they equal the mother cell in size. In the meantime other buds may arise
on the mother cell and on the buds already in process of development so
that a colony results consisting of many cells of various sizes and ages. In
the process of budding a small bud is produced and then into it projects
an extension of the nucleus of the mother cell. According to Lindegren
(1945, 1949) this extension separates by mitosis from the mother nucleus
and then the opening between the two cells is closed except for a small
central pore which always remains through the separating septum in all

336 CLASS ASCOMYCETEAE

the Higher Fungi. This is closed before the cells separate from one another.
Fabian and McCullough (1934) reported that by growing some yeasts
under certain cultural conditions or by the addition of lithium salts to the
culture media the vegetative cells may be induced to break up into
minute cells which differ, not only in size and shape but in their ability to
cause fermentation, withstand changed environment, etc. On inoculation
upon the customary culture media they resume the normal size and char-
acteristics of the original cells. This is of interest as it shows a parallelism
in behavior to that demonstrated by Hadley (1927) for some of the true
bacteria.

Sexual reproduction consists of the union of two equal or unequal cells
and the almost immediate fusion of the two gamete nuclei. This diploid
cell or some of its products by division enlarges to become a single ascus
within which are produced usually eight or four or sometimes fewer asco-
spores (even as low as one in the genus Monospora). In Ascoidea and
Dipodascus many ascospores are formed. Unlike the case in many of the
Ascomyceteae the ascospores in this order are mostly not expelled vio-
lently by the bursting of the asci.

In the older classifications of the families within this order the forms
normally with mycelium are placed in two or three families and the yeast
forms in about as many. The fact that the same type of ascospores may
occur in genera assigned to several of these families throws doubt on the
validity of these classifications (Zender, 1926). Thus the hat-shaped asco-
spores occur in the unicellular genus Hansenula (Willia), in some species
of the filamentous genus Endomyces, and in the filamentous Ascoidea
ruhescens Bref. These three genera are usually placed in three separate
families. This spore type is apparently closely related to that found in
those species of the genus Aspergillus in which there are two parallel
ridges with a furrow between them.

It seems to the author that the yeast forms are to be considered as
derived more or less independently from the Endomycetaceae and perhaps
other families. In that case the customary yeast family ox families are not
true phylogenetic units but must eventually be merged with the families
from which they are derived. Pending further research by students of
these groups the more customary family distinctions are followed here.
The most recent extensive work on these organisms with the modern sys-
tem of classification is that by Miss Stelling-Dekker (1931), Miss Lodder
(1934), and Diddens and Lodder (1942).

AspoROGENOus Yeasts. Closely resembling the yeasts and the fila-
mentous genera of this order are many forms sometimes called the Asporo-
genous Yeasts. These do not produce asci nor are there any evidences of
sexual reproduction. Possibly some of them may be true yeasts that have
permanently lost the power of producing asci. Others, however, may rep-

OEDER SACCHAROMYCETALES 337

resent derivatives of entirely different orders or even classes of fungi which
have evolved into yeast-like forms. Under certain conditions of culture
yeast-like cells are formed by some Mucorales, Ustilaginales, and many
families of Ascomyceteae. Although their true relationships are doubtful
they are given consideration here. Apart from these are two families that
possess sexual reproduction and a simplicity of structure that implies
either great reduction or, on the contrary, a great degree of primitiveness:
the Spermophthoraceae and Pericystaceae. Their relationships are very
doubtful. Their consideration in this chapter is due to a lack of conviction
on the part of the author as to where else they might more properly find
their relationship, rather than because of any evident kinship with the
Saccharomycetales.

The first two families to be discussed are those that are normally
filamentous and ascus-producing.

Family Endomycetaceae (Including Eremascaceae of Some
Authors). Typically forming a branched mycelium with the cells uninu-
cleate or containing several nuclei. Asexual reproduction by the breaking
up of branches of the mycelium into " oidia" or by the formation of bud-
ding branches. Some species are capable of alcoholic fermentation. Sexual
reproduction takes place by the union of projections of usually (but not
always) adjacent cells to form an ascus perched upon the tips of the united
processes. In some cases the gametangia are distinctly a small antherid
terminal to a slender antheridial branch and a large oogone. In some
species the two processes do not unite or even only one is produced so that
the ascus is formed parthenogenetically. Ascospores are eight or four or
less, frequently fewer. They are ellipsoidal or hat-shaped.

Eremascus fertilis Stoppel and E. albus Eidam produce eight asco-
spores after the union of the tips of adjacent cells which may be tightly
coiled around one another in the latter species. Frequently union fails to
occur and the parthenogenetically produced asci contain four or fewer
ascospores. The mycelium produces no conidia (either oidia or budding
cells). Endomyces is distinguished from Eremascus by the production of
four or fewer ascospores per ascus and by asexual reproduction by means
of oidia. Endomycopsis differs from both by the production of budding
cells as well as occasionally of oidia. In both genera ascus formation may
be parthenogenetic or may be the result of the union of two equal or
unequal gametangia. Some species of both genera are able to ferment
various sugars but some lack this power. Endomycopsis albicans (Vuill.)
Dekker has been found in cases of the disease of the mouth known as
thrush although this mostly seems to be due to an asporogenous filamen-
tous yeast. The various other species of the two genera are found in soil
and on various vegetable products. It must be noted that these two
genera are very close to the true yeasts and possibly the distinction be-

338

CLASS ASCOMYCETEAE

Fig. 109. Saccharomycetales, Family Endomycetaceae. (A-D) Eremascus fertilis
Stoppel, stages in isogamic sexual reproduction. (E-H) Endomyces magnusii Ludwig,
stages in anisogamic sexual reproduction. (After Guilliermond: Ann. fermentations,
2:129-151, 257-277.)

tween this family and the two famihes of yeasts will have to be abolished.
(Fig. 109.)

Attention should be drawn to two filamentous fungi which agree with
this order in simplicity of structure and the formation of asci singly and
not in a hymenium. Both are parasitic in the hymenium of species of
Corticium. In the authors' opinions these are highly derived, simplified
forms possibly derived from apothecial genera. They are Trichomonascus
(Jackson, 1947) and Myriogonum (Cain, 1948). Their relationship to the
filamentous Saccharomycetales is doubtful, but they may be placed here
until they have been studied in culture and their full ontogeny observed.

Family Ascoideaceae. In many respects this family shows relation-
ship to the Endomycetaceae, but the differences seem large enough to
warrant their being kept distinct. The two genera Dipodascus and
Ascoidea form well-developed mycelium. In the former asexual reproduc-
tion is by the breaking up of the terminal portion of the hypha from the
apex downwards to form oidia in one species and is lacking in the other.
In Ascoidea the conidia are produced singly at the tips of the hyphae
which by sympodial development grow by the conidium and produce
another one until the hypha appears to have one terminal and many

ORDER SACCHAROMYCETALES

339

lateral conidia. The asci are elongated and many-spored. They open at
the apex and the ascospores escape, sometimes in a worm-like mass.

Dipodascus alhidus Lag. was found growing in the gummy exudate
of the cut surface of a tropical species of Bromeliaceae. Its septate
mycelium consists of multinucleate cells. From two adjacent cells in a
hypha arise multinucleate branches which come into contact and fuse
at their tips. These branches are then separated from the main hypha by
septa. At the tips of each of the gametangia a single "privileged" nucleus
enlarges and these two unite in or near the passageway formed by the

Fig. 110. Saccharomyce-
tales, Family Ascoideaceae.
Dipodascus uninucleatus
Biggs. Stages in sexual re-
production. (A) Early stage
in conjugation of adjacent
cells. (B) Zygote cell en-
larged and nuclei united,
thus forming primordium of
ascus. (C) Young ascus in
4-nucleate stage. (D) Young
ascus with 16 nuclei. (E)
Mature ascus. (After Biggs:
Mycologia, 29(l):34-44.)

dissolution of the intervening walls. From this region grows upward the
somewhat tapering ascus into which pass not only the cytoplasm of the
gametangia but also their many nuclei and the larger zygote nucleus.
The latter divides, apparently meiotically, and probably the four nuclei
thus produced continue to divide. It is assumed that it is the nuclei thus
formed that serve as the centers about which the ascospores are produced.
There are many degenerating nuclei in the ascus which may be the original
haploid nuclei introduced from the gametangia. Eventually the apex of
the ascus dissolves and the ellipsoidal ascospores escape in a gummy
mass. The vegetative hyphae reproduce asexually by the formation of
oidia. In D. uninucleatus Biggs it was shown by Miss Biggs (1937) that
the cells of the mycelium and the gametangia are uninucleate. As a result

340

CLASS ASCOMTCETEAE

there are no supernumerary nuclei in the ascus, the many nuclei there
produced being the product of the division of the zygote nucleus. Other-
wise the ascus resembles that of D. alhidus. No asexual reproduction has
been observed in this species. (Fig. 110.)

Ascoidea, with the single species A. rubescens Bref., also grows in
plant exudates (such as the beech, Fagus silvatica L.) and the mycelium
consists of multinucleate cells. No fusion of separate gametangia occurs
but the oval asci are produced terminally on upright hyphae. They are
from the beginning multinucleate and produce very numerous hat-shaped
ascospores like those of Hansemda and some species of Endomyces.

*•«££■

'is •■.'115 ^ W-

Fig. 111. Saccharomycetales, Family Ascoideaceae. Ascoidea rubescens Bref.
(A) Sympodial formation of conidia. (B) Proliferated asci. (C) Young multinucleate
ascus proliferating in discharged ascus. (D) Ascus approaching spore formation, nuclear
divisions simultaneous. (E) Portion of ascus with spores. (F) Mature spore. (G-I)
Stages in conjugation and germination of spores. (A-B, After Engler and Prantl: Die
Natiirlichen Pflanzenfamilien, Leipzig, W. Engelmann. C-I, after Walker: Mycologia,
27(2):102-127.)

ORDER SACCHAROMYCETALES 341

According to Varitchak (1931) two of the many nuclei in the young ascus
enlarge and then unite. From the division of this zygote nucleus it is
supposed that the nuclei of the ascospores are formed. After the ascospores
are set free by the dissolution of the apex of the ascus the basal septum
arches upwards and a new ascus is produced by proliferation. This may
occur many times as in the formation of sporangia in Saprolegnia. Miss
Walker (1935) was unable to confirm Varitchak's report of the union of
two privileged nuclei and believes that the ascus develops partheno-
genetically. She noted earlier (1931) that the ascospores frequently fuse
by twos after their escape and produce mycelium originating from the
conjugation tube connecting the two ascospores. It should be noted that
in Dipodascus uninucleatus frequently the ascus may be produced par-
thenogenetically. If that should occur also in Ascoidea it may be that
both Varitchak and Miss Walker were correct in their reports. (Fig. 111.)

The true yeasts, i.e., those forms of this order that are normally not
hyphal, may be divided into the families Schizosaccharomycetaceae in
which the cells divide by fission and Saccharomycetaceae in which they
divide by budding. Many of them are of great industrial value because
of their power of fermenting various sugars and producing alcohol and
CO2, but many are unable to ferment the commoner sugars. Some of the
alcohol-producing yeasts grow at the top of the solution to be fermented,
the so-called "top yeasts," while others are more abundant where the
oxygen supply is less, in the lower portion, the "bottom yeasts."

Sexual reproduction is quite varied. Perhaps the more common mode
is the union of two cells, more often equal in size but sometimes of un-
equal size, by means of short tubes. The intervening walls are dissolved
and the two nuclei unite, usually in the conjugation tube. The number of
ascospores is more often four but in many cases is eight. Probably by
failure of some of the nuclei to function in the production of ascospores
the number may be less than four, in a few cases only one. The mature
ascospores may be distributed evenly or unevenly between the two con-
jugating cells or may be found in only one. They are more often ellipsoidal
but may be hat-shaped or spherical or needle-shaped. A modification of
the foregoing type is the conjugation of a mother cell with a bud produced
from it. Both cells may produce conjugation tubes or the partition be-
tween the larger cell and the bud may dissolve away. This process has
been called pedogamy. It is clear that this is a modification of the pre-
ceding type. In both types the vegetative cells may be assumed to have
haploid nuclei with immediate meiotic division of the zygote (diploid)
nucleus to bring about the ascospore production.

In some yeasts the four ascospores unite by twos within the ascus.
From the conjugation tube a process pushes through the ascus wall and
produces a daughter cell outside. This cell multiplies by budding or fission

342 CLASS ASCOMYCETEAE

depending upon the type of asexual reproduction characteristic of the
species. Eventually some of these cells enlarge a little and the nucleus
divides (probably meiotically) into four and the four ascospores are
produced. These in their turn conjugate by twos. Yeasts of this type are
apparently diploid during their whole vegetative life, the ascospores ^lone
being haploid. Winge (1935) showed that sometimes a pair of ascospores
may be so situated in the ascus that they do not unite. In that case each
buds vegetatively. The colonies thus formed consist of haploid cells
which are smaller and more rounded than the normal diploid cells of the
species. Soon one or more pairs of these haploid cells, of varying degrees of
relationship, fuse and the resulting diploid cell multiplies by budding to
produce the larger more elongated cells characteristic of the normal vege-
tative development of the species.

A third category of yeasts has been assumed to produce its asci par-
thenogenetically. Saccharomyces cerevisiae Meyen, the common yeast of
bread and beer, does not show conjugation of the ascospores within the
ascus nor is such conjugation observed immediately prior to ascus
formation. However the Lindegrens (1943, 1949) have shown that the four
ascospores of a normal four-spored ascus when germinated separately in
the proper culture media produce colonies of haploid cells resembling
those mentioned above in the case of failure of a pair of ascospores to
conjugate within the ascus. Mostly these remain of the haploid type
although occasionally conjugation between two cells may occur to form an
"illegitimate" diploid colony. This may, under conditions favorable for
ascosporogenesis, produce asci but these usually contain only two asco-
spores which are mostly incapable of germination. These illegitimate
diploid colonies when perpetuated in conditions favorable for asexual
multiplication may be of industrial value since these strains are frequently
very constant and free from mutations. When the haploid colonies from
the four ascospores are mated vigorous diploid colonies develop in certain
matings and these under conditions favorable for ascospore formation
produce four-spored asci whose ascospores are capable of germination.
These matings show that two of the four ascospores in the normal ascus
are of one mating type and two of another. The haploid strains from the
original four ascospores may develop into lines that under no mating
combinations are able to produce ascospores. In other words they may by
mutation become asporogenous yeasts to which the name Torulopsis
(Toriila) has been given in the past (see below). Saccharomyces paradoxus
Batsch, as shown by Guilliermond (1936), is variable as to the point
where sexual reproduction occurs. Under some conditions the diploid
phase vegetative cells become asci and produce ascospores which con-
jugate in the ascus. From the zygotes thus formed arise by budding the
usual vegetative cells of the yeast. Sometimes in the same species under

ORDER SACCHAROMYCETALES

343

Fig. 112. Saccharomycetales, Family
Saccharomycetaceae. Saccharomyces cere-
visiae Meyen. (A) Cell before beginning
of formation of bud. (B) Bud formed but
still without nucleus. (C) Nucleus push-
ing out into bud. (D) Nucleus divided,
with one daughter nucleus in each cell.
(After Guilliermond : Ann. Mycolog.,
2(2):184-189.)

Fig. 113. Saccharomycetales, Family Saccharomycetaceae. Saccharomyces para-
doxus Batsch. (A) Two ascospores (a and b) have, without conjugation, formed systems
of budding cells (1-4) with haploid nuclei. (B) Cells 2 and 3 from ascospore a have
conjugated and formed bud (a) with diploid nucleus, and there has occurred further
formation of haploid buds in the system from ascospore b. (C) Cells 2 and 5 from
ascospore b have conjugated and produced the diploid buds (a), (b), and (c), and
further diploid buds (b) and (c) have developed from the previously formed zygote.
(Courtesy, Guilliermond: Ann. fermentations, 2:129-151, 257-277.)

344 CLASS ASCOMYCETEAE

CO (TO

A B

Fig. 114. Saccharomycetales, Family Schizosaccharomycetaceae. Schizosaccharo-
myces odosporus Beijer. (A-E) Stages in the conjugation of two equal haploid cells
and the formation of the ascus and ascospores. (Courtesy, Guilliermond: Arm. fermen-
tations, 2:129-151, 257-277.)

other conditions the ascospores do not conjugate but bud and form small
colonies of adhering haploid yeast cells. The cells of a single colony arising
from a single ascospore conjugate by pairs, thus giving rise to the diploid
phase again. This species does not have the two mating types which are
possessed by S. cerevisiae. A very full discussion of the various types of
sexuality in the yeasts is given by Guilliermond (1936). (Figs. 112, 113.)

Family Schizosaccharomycetaceae. The only genus is Schizo-
saccharomyces. Only two species are definitely recognized by Stelling-
Dekker (1931), Sch. pom^e Lindn., with mostly four (or fewer) ascospores,
and capable of fermenting saccharose, and Sch. odosporus Beijer., with
eight ascospores and incapable of fermenting saccharose. In both species
conjugation of two vegetative cells initiates ascus formation. Sch.
versaiilis Wick. & Duprat (1945), by its production of hyphae under
anaerobic conditions, shows relationship to Endomyces. (Fig. 114.)

Family Saccharomycetaceae. In this family are found most of the
industrial strains of yeasts. The genus Saccharomyces is the most impor-
tant. Many species have been described, but the differential characters
have been largely based upon their fermentation abilities with different
kinds of sugar. In the light of the recent genetic studies of S. cerevisiae it
seems likely that many of these represent merely segregations or muta-
tions of genetic characters which have been perpetuated nonsexually.
Saccharomycodes differs from Saccharomyces mainly in the fact that the
ascospores conjugate within the ascus so that the whole life history is
diploid. The genus Hansemda (Willia) resembles Saccharomyces vege-
tatively but the ascospores are spherical or ellipsoidal with an equatorial
ridge or spherical and flattened on one side with a rim at the edge, giving
the appearance of a hat. In Deharyomyces the ascus arises from isogamous
or heterogamous conjugation and the spherical ascospores are roughened
with fine warts. They are usually one (rarely two to three) per ascus. The
genus Nemato.spora contains two or three species of yeasts that are para-
sitic in plant tissues but easily cultivated in various culture media.
Asexual reproduction is by budding. In some species short mycelia are
produced. The single yeast-like cells or the cells of the mycelium may be-
come transformed into asci within which are produced mostly eight (rarely
four or fewer) ascospores. These are usually in two bundles of four spores

ORDER SACCHAROMYCETALES 345

each, at the two ends of the ascus. The spores are needle-shaped, tapering
at one end to a slender flagellum-like thread which is not motile. Sexual
union of two cells to produce the ascus has been claimed for N. lycopersici
Schneid. by Schneider (1916), by Wingard (1925) in A'^. phaseoli Wing.,
and in N. coryli Peglion by Miss Manuel (1938), but is denied by Guillier-
mond (1928). Probably closely related to this genus is Ashbya gossypii
(Ashby & Now.) Guill. which causes injury to the bolls of cotton {Gos-
sypium). It differs in having a very limited yeast stage, mainly developing
a coenocytic, variously branching hypha, one end of which, without a
separating septum, functions as the ascus. Guilliermond claims that the
nuclei functioning in the ascospore development do not arise from the
division of one original ascus nucleus but by the division of several nuclei
already present in that portion of the
hypha. Both Nematospora and Ashbya
are apparently spread by the punc-
tures of various sucking insects. The
genus Eremothecium is very similar to
Ashbya. E. ashbyii Guill. (1936a) was,
like it, isolated from diseased cotton
bolls. It differs chiefly in the slightly
different form of the ascospores. In . Fig. 115. Saccharomycetales, Fam-
, , i 1 . 1 ily Saccharomycetaceae. Saccharo-

several respects these three genera ,„^^^^^^ ;^^^,„-^.. Hansen. (A) Ascus

suggest the gametophytic generation with four ascospores. (B) Conjugation

of Spermophthora gossypii Ashby & of ascospores by twos. (C) Zygotes
--. ,, ,.,.,. Ill germinating to form new diploid cells.

Nowell, which IS discussed below (Courtesy, Guilliermond: Ann. fer-
(under Family Sperm ophthoraceae). mentations, 2:129-151, 257-277.)
(Fig. 115.)

Monosporella bicuspidata (Metschn.) Keilin, parasitic in Daphnia (a
Crustacean), and Coccidiascus legeri Chatton, parasitic in a species of the
fly Drosophila, have similar ascospores and possess a budding yeast-like
stage. They probably are closely related to the foregoing genera.

The intensive studies by Miss Stelling-Dekker (1931) upon the large
collection of yeasts in the Centraalbureau voor Schimmelcultures in
Baarn, Netherlands, have led her to the conclusion, already expressed by
Van Tieghem in 1906, that the distinction between the three families En-
domycetaceae, Saccharomycetaceae, and Schizosaccharomycetaceae can-
not be maintained and that they must be united under the older name
Endomycetaceae with the four subfamilies Eremascoideae, with the
single genus Eremascus; Endomycoideae, containing Endomyces and
Schizosaccharomyces; and Saccharomycoideae, with Endomycopsis and
most of the sporogenous genera of yeast habit; and Nematosporoideae.
This scheme of classification was adopted by Guilliermond (1936b) in a
contribution in which the various genera are discussed and illustrated.

346 CLASS ASCOMYCETEAE

It seems likely that this tentative scheme may in the future form the best
basis for the classification of the sporogenous yeasts and near relatives.
Further cytological studies are desirable before this can be considered as
entirely settled.

The Asporogenous Yeasts are those in which no formation of asci is
known. Since it has been demonstrated by Lindegren and Lindegren
(1943) and suggested by others, also, that races of species of the genus
Saccharomyces may lose their power of ascus production, probably as a
result of some gene mutation, the distinction between the two groups is
very doubtful as regards some genera.

Family Nectaromycetaceae. This very doubtful family contains the
single genus Nectaromyces named by the Sydows (1918). This fungus
grows in the nectar of flowers and is easily cultivated. It produces yeast-
like budding cells in abundance, but under certain conditions produces a
cluster of four cells cruciately arranged or "airplane-like." Also long
branched hyphae may occur, which bear conidia at their tips.

Family Torulopsidaceae. The members of this family are in the
main quite similar to the yeasts of the Saccharomycetaceae but differ
from them in the complete absence of ascus formation. As mentioned
above this may be due in some cases to gene mutations in sporogenous
yeasts. On the other hand it must be borne in mind that yeast-like cells
are produced under the proper conditions of environment and nutrition
in such widely distinct fungus orders as the Mucorales, Ustilaginales, and
some of the Heterobasidiae and Eubasidiae, besides various species of
different orders of the Ascomyceteae. Hence it is conceivable that this
family may be of heterogeneous origin. Many of the species are capable of
fermenting various sugars with production of CO2 and alcohol or related
substances; others can oxidize the sugars in the presence of oxygen but do
not cause fermentation. Many of the latter and a few of the former are
pathogenic in Man and other animals. Following Lodder (1934) and
Diddens and Lodder (1942) the family is divided into two subfamilies:
Torulopsoideae, in which no mycelial stages occur and Mycotoruloideae,
in which mycelia or pseudomycelia occur, from which at the apex or along
the hypha budding takes place. Besides this the fungus may grow as
simple budding cells.

In the Torulopsoideae Lodder recognizes seven genera of which Toru-
lopsis and Pityrosporum may be mentioned. Torulopsis is known in medi-
cal literature more commonly as Torula or as Cryptococcus and sometimes
as Blastomyces. The name Torula properly belongs to a genus of filamen-
tous fungi of the Family Dematiaceae of the Fungi Imperfect!. On that
account Berlese proposed, according to Lodder, the name Torulopsis in
1894 for the yeast-like, asporogenous organisms to which the other names
are so often given. The species of this genus resemble those of Saccharo-

ORDEB SACCHAROMYCETALES 347

myces except for the failure to produce asci. The cells are mostly short or
long ellipsoidal, propagated by budding and never producing true hyphae.
The genus is divided into two subgenera : the species of the first cause the
fermentation of various sugars and are almost without exception non-
pathogenic to animals, the species of the second subgenus produce no
fermentation in sugars although they may oxidize them as a source of
energy. A number of these are capable of parasitic life in animals. The
genus Pityrosporum contains a few species that grow in the skin of Man or
other animals, causing scaliness and falling of hairs (pityriasis capitis).
Besides the short, oval, budding cells there occur some cells that are
flask-shaped, with a larger basal portion and a smaller rounded apical
portion. The genus Mycoderma of Persoon as amended by Lederle (see
Lodder, 1934) is much like Torulopsis but under some conditions the
budding cells remain attached to form a short branching pseudohypha.
The species grow in wine, beer, etc., forming a surface film. They feed
upon the sugars and alcohols but produce no true fermentation. The genus
Kloeckera resembles Torulopsis but the cells are in part apiculate, resem-
bling a lemon. Some of the species cause fermentation of various sugars.
In the Mycotoruloideae (established under the name Mycotoruleae
by Ciferri and Redaelli in 1929) the cells form definite septate or non-
septate hyphae or pseudohyphae from which sprout crowns or tree-like
growths of buds. All the species bud freely and in certain media do not
produce mycelium. Many are pathogenic for animals. In medical litera-
ture a great many are reported under the generic name Monilia. This
name is usually reserved by mycologists for saprophytes or parasites on
plant tissues for many of which the perfect stage is known to belong to
the Discomycetous family Sclerotiniaceae. Blastodendrion produces pseu-
domycelia from the cells of which sprout crown-like or branching clusters
of ellipsoidal cells. A number of species growing more or less pathogen-
ically occur in Man in the alimentary canal or on moist skin, nails, etc.
Candida (synonyms: Monilia, Geotrichoides, etc.) produces regularly
septate hyphae at whose apices chains of yeast-like cells may be produced
as well as single such cells or clusters at the joints of the hypha. Many
species are pathogenic. The species reported as C. albicans (Robin)
Berkh. is found as a cause of the disease of the mouth known as "thrush."
It is placed by C. W. Dodge (1935) in a closely related genus Syringospora.
This fungus is often confused with Endomycopsis albicans (Vuill.) Dekker
which occurs in cases of thrush and is similar to the foregoing in many
respects, but produces asci and ascospores. The classification by C. W.
Dodge (1935) of the sporogenous and asporogenous yeasts and nearly
related filamentous fungi that are pathogenic to animals is very different
from that given above which is based mainly upon Ciferri and Redaelli
(1929) and Dekker (1931) and Lodder (1934). This monumental work of

348

CLASS ASCOMYCETEAE

Dodge should be consulted by all mycologists interested in medical
mycology.

Family Rhodotorulaceae (The Red Yeasts). This family was set
apart by Lodder from the Torulopsidaceae because of the production in
the yeast-like cells of carotinoid pigments, thus giving the colonies a red
to orange color. Rhodotorula is the one genus recognized. The cells are
round to elongated and bud freely. Occasionally the resulting cells remain
attached in a row to form a short moniliform pseudohypha. No true fer-
mentation of sugars occurs but dextrose and various other sugars are
oxidized in aerobic respiration. In some species growth occurs in ethyl
alcohol. Some species are soil inhabitants, probably saprophytes, but
some are found in human sputum and others apparently attack hair. The
relationship of this genus is uncertain. It may be a true yeast that has
lost its power of producing asci or it may be a much reduced, yeast-like
fungus of entirely different origin.

Family Sporobolomycetaceae. In 1930, Derx recognized two genera,
with seven species in Sporobolomyces and two in Bullera, and another
genus, Tilletiopsis, which he suggested showed affinities to these. In 1948,
the same author described another genus, Itersonilia, which together with
the other three he now recognizes as belonging to the family. They are all
characterized by the production of aerial "ballistospores," a name pro-
posed by Derx for spores usually asymmetrically perched at the apex of

Fig. 116. Saccharomycetales (?), Family Sporobolomycetaceae. Sporobolomyces
roseus Kl. & van N. (A) Vegetative cell. (B-D) Stages in formation of bud. (E-H)
Stages in formation of aerial spore. (I) Mother cell after discharge of spore. (Courtesy,
Buller: Researches on Fungi, vol. 5, London, Longmans, Green & Co.)

ORDEB SACCHAROMYCETALES 349

the sterigmata and discharged violently, often with the production of a
droplet of water at the base of the spore just before its discharge. As in the
Subclasses Teliosporeae and Heterobasideae these spores are capable of
germination by repetition, i.e., by forming sterigmata upon which similar,
but somewhat smaller ballistospores are produced. In the first two genera
colonies are formed in which the usual mode of reproduction is by bud-
ding, as in yeasts, producing considerable mucilage. In the other two
genera true hyphae are produced and no budding occurs. Nyland (1949)
has described still another genus clearly belonging to this family, to
which he has given the name Sporodiobolus. Like Itersonilia the mycelium
has an abundance of clamp connections. The cells are binucleate. An
abundance of chlamydospores is produced in Sporidiobolus. These are
binucleate but soon the nuclei unite and the spore becomes golden-brown.
In the younger stages of growth the cells bud, as in yeasts, forming a
yeast-like colony similar to that of Sporobolomyces. No asci or ascospores
occur in any of the four genera. These fungi occur on leaves covered
with sooty mold or injured by insects or by fungus parasites, and fre-
quently may be isolated from the soil. The aerial ballistospores are
distributed by air currents. (Fig. 116.)

In Sporobolomyces a carotinoid pigment is present so that the colonies
are red to salmon colored, but in Bullera the pigment is lacking and the
colonies are pallid to yellowish. Neither of these two genera normally
produces hyphae, but it should be noted that sometimes in Spor^obolo-
myces, in old cultures, a few short, branching hyphae may arise and on
them develop laterally and terminally the characteristic spore-bearing
sterigmata. In normal development the first extensive growth is by
budding. Later some of the surface cells send out one (or even two or
three) sterigma each, which in some cases may become forked. Perched
obliquely at the tip of the sterigma a usually asymmetrical spore is
formed, like a basidiospore on its sterigma.

The genera Tilletiopsis and Itersonilia form definite hyphae but not
yeast-like colonies of budding cells. From these hyphae arise sterigmata
from which the spores are discharged as described above. In Itersonilia a
clamp connection is formed at every septum while these are lacking in
Tilletiopsis.

Kluyver and van Niel (1924) suggested that this mode of spore dis-
charge of the aerial spores warrants the idea that these fungi may be very
much reduced Basidiomycetes. Buller (1933) investigated this process
more fully and accepted the idea. Lohwag (1926) and Guilliermond (1927)
did not agree with this suggestion because the cells of the then recognized
species are uninucleate and there is no fusion of nuclei prior to spore
formation, so that these cells can not be considered as much reduced
basidia. Buller, on the other hand, pointed out that in a number of the

350 CLASS ASCOMYCETEAE

Basidiomyceteae the cells are uninucleate, including the basidia, and
there is no nuclear fusion in the latter. This can be explained, according
to him, by loss of sexuality, as has happened in many of the Saccharo-
mycetales and other fungi, or by the occurrence of + and - strains which
in the absence of the reciprocal strain proceed to produce their basidio-
spores parthenogenetically. With the discovery of Itersonilia, with its
mycelium made up of dicaryon cells (as evinced by the clamp connec-
tions), and of Sporidioholus it seems that probably these are really much
reduced forms of Basidiomyceteae from either the Subclass Teliosporeae
or Subclass Heterosporeae, in both of which the teliospores often germi-
nate by repetition.

The two families whose discussions follow are of very uncertain rela-
tionship. It is possible that the Spermophthoraceae are not far from the
Endomycetaceae. The Pericystaceae are of still more doubtful kinship.
Both need further investigation in order to confirm the validity of the
reported life histories and the conclusions drawn therefrom as to their
relationship.

Family Spermophthoraceae. The only known species, Spermophthora
gossypii Ashby & Nowell, was determined to be the cause of the disease
called stigmatomycosis affecting the seeds of cotton (Gossypium) and
fruits of tomato {Ly coper sicon) in the West Indies. This was studied first
by Ashby and Nowell (1926) and further by GuilHermond (1928). The
germinating ascospores give rise to a nonseptate, coenocytic, dichoto-
mously branching mycelium with apical growth. The apices of the
branches may continue to grow while the older parts die and are cut off
by callose plugs as in some Phycomyceteae. No cellulose reaction is
shown upon treatment with chloriodide of zinc. Sometimes small cells
bud off from this mycelium and these may bud in turn, but it has not been
ascertained whether these are functional asexual spores. A short distance
back from the hyphal tips the mycelium forms spindle-shaped swellings,
the gametangia, which become separated from the usually short tip cells
by a cross wall, another septum setting this swelling off from the main
portion of the mycelium. The gametangia contain at first from four to ten
nuclei. These divide twice simultaneously and the resulting nuclei and
most of the cytoplasm form a dense axial strand surrounded by a vacuo-
late epiplasm. The axial portion divides into fusiform, uninucleate cells
which enlarge at the expense of the epiplasm. The rupture of the game-
tangium wall permits the nonmotile gametes to escape. Whenever two lie
in contact they unite by a conjugation tube within which, usually, the two
nuclei unite. From this conjugation tube there grows out a rather Hmited
branched septate mycelium of uninucleate cells. The ends of the hyphae
enlarge and are cut off by a septum and become spherical asci. The
nucleus divides three times and around each nucleus is formed a more or

ORDER SACCHAROMYCETALES

351

Fig. 117. Saccharomycetales (?), Family Spermophthoraceae. Spermophthora
gossypii Ashby & Nowell. (A) Gametophytic, coenocytic mycelium. (B) Developing
gametangia. (C) Mature gametangia containing gametes. (D-G) Stages in union of
gametes. (H-J) Sporophytic mycelium bearing asci. (After Guilliermond : Rev. gin.
botan., 40:328-704.)

less fusiform or lemon-shaped ascospore. As the eight ascospores enlarge
they use up the epiplasm. They usually remain in a rather compact
bundle even after the ascus wall disappears. Rarely a gamete germinates
parthenogenetically to form a septate mycelium bearing apparently
normal asci. Although the cytological details could not be made out
Guilliermond believed that the first two divisions of the ascus are meiotic
so that the fungus shows an alternation of a gametophyte with haploid
nuclei and a sporophyte with diploid nuclei. In 1936 he reported that in
cultures maintained for ten years the fungus had lost its power to produce
the ascogenous mycelium. The gametes from the gametangia no longer
underwent conjugation but germinated directly to repeat the gameto-
phytic generation. He suggested that Ashhya and Eremothecium may
represent a similar development in which sexuality has been lost and only
an asexual generation is present. (Fig. 117.)

Guilliermond (1928) believed that Spermophthora occupies a position
intermediate between the Phycomycetes and Ascomycetes. Comparing
it with higher Ascomyceteae (e.g., Pyronema) he considered its sporo-
phytic stage with uninucleate cells and diploid nuclei to be homologous
to the system of ascogenous hyphae with cells containing pairs of haploid
nuclei. In Spermophthora the single diploid nucleus in the ascus divides to
form the nuclei of the ascospores ; in Pyronema the two haploid nuclei of
the ascus unite and then divide to form the nuclei of the ascospores. In
both the ascospores with their haploid nuclei give rise to a gametophyte.
In Spermophthora the gametangia produce individual gametes which are
set free and unite ; in the higher Ascomyceteae two gametangia (antherid

352

CLASS ASCOMYCETEAE

and oogone) unite but the contained nuclei do not unite now but wait
until the ascus is formed. On the basis of this homology Guilliermond
considered the Saccharomycetales to represent a lateral offshoot in which
the union of gametangia has become established but in which the sporo-
phytic phase has undergone reduction to a single ascus. Thus Dipodascus,
Ascoidea, Endomyces, etc., are, according to him, not in the line of evolu-
tion to the higher Ascomyceteae which arose more directly from forms
intermediate between them and Spermophthora.

Family Pericystaceae. The only genus recognized is Pericystis, the
cause of "chalk-brood" and other troubles in beehives. Two species have
been described, apparently both feeding upon the stored pollen in the
cells of the comb. The first species described was P. alvei by Miss Betts
(1912) and the second was P. apis described by Maassen in 1916. The
latter besides feeding upon the pollen appears to attack the larvae in the
cells of the comb. It was given careful study by Claussen (1921). The
systematic position of this genus is in doubt. Fitzpatrick (1930) placed it
among the doubtful Phycomyceteae. Varitchak (1933) who studied the
life history and nuclear behavior of P. apis, concluded that it is a primitive
Ascomycete related to Dipodascus and Ascoidea but far nearer the sup-
posed phycomycetous ancestors. The mycelium, as in Ascoidea, contains
chitin and not cellulose. As in that fungus it is septate with multinucleate
segments, the septa being centrally perforated, thus permitting the flow
of cytoplasm from segment to segment. The mycelia are of two sexes, the
fungus being heterothallic. When they come into contact multinuclear
gametangia are produced, each separated by its septum from the main
hypha. They resemble at first the early stages of conjugation in Mucor.

Fig. 118. Fungus of uncertain relationship. Family Pericystaceae. Pericystis apis
Maassen. (A) Conjugating gametangia. (B) The same, in optical section. (C) Oogone
with zygotes. (D) Oogone with zygotes which have produced spores and thereby have
become spore balls. (After Claussen: Arh. biol. Reichsanstalt Land- u. Forstw., 10:467-
521.)

KEY TO THE GENERA OF FAMILY ERYSIPHACEAE 353

The female gametangium enlarges and the male gametangium sends a
conjugation tube into it. Numerous male nuclei and some cytoplasm
enter the oogone, whose nuclei divide several times as do the introduced
male nuclei. Many nuclei of both sexes degenerate but many pairs of
uniting nuclei are found. Around each such zygote nucleus is organized a
mass of cytoplasm, called by the investigator an "egg." There is no cell
wall between this egg and the rest of the protoplasm. The number of such
zygote nuclei and eggs varies, usually being large but being reduced to
one in rare cases. Each zygote nucleus divides several times and the egg
cytoplasm undergoes cleavage until as many spores are formed as there
were nuclei produced. Spore walls are formed and within the gametangium
are now found as many spore balls as there were uniting pairs of nuclei.
Varitchak called each ball of spores an ascus and the whole structure a
"synascus." By reduction of the number of uniting nuclei and resultant
spore balls a condition would be attained, according to him, similar to
that in Dipodascus whose ascus he calls a "hemiascus." By reduction of
the number of nuclei in the uniting gametangia to one in each as in
Endomyces a true ascus would be formed. (Fig. 118.)

Key to Families of Order Erysiphales

Aerial mycelium hyaline. Outer colored layer of perithecial wall one cell in thick-
ness, of polygonal cells, brittle. Family Erysiphaceae
Aerial mycelium (when present) dark. Outer peridium layer not brittle.

Peridium parenchymatous, not slimy; mycelial hyphae cylindrical, not slimy.

Family Meliolaceae
Peridium dissolving into slime; mycelial hyphae moniliform or cylindrical.

Family Englerulaceae
Perithecium walls built of moniliform or cylindrical hyphae, more or less slimy.
Mycelium moniliform or cylindrical, united in strands.

Family Capnodiaceae
MyceUum dark-colored, parasitic on epiphyllous Meliolaceae and similar fungi.
Perithecium inverted at maturity, with the morphological base upward.

Family Trichothyriaceae
Forming rounded or stellate cushions on leaves, gelatinous when wet, horny
when dry, with no free mycelium. Asci scattered in thickened area of the
thallus. Family Atichiaceae

Key to the Genera of Family Erysiphaceae

Ascospores one-celled.

Perithecia normally containing only one ascus.

Appendages hyplia-like, usually unbranched. Sphaerotheca

Appendages stiff, dichotomously forked at tip. Podosphaera

Perithecia normally containing several to many asci.

Appendages hypha-like, mycelium external, conidia in chains.

Erysiphe
Appendages hypha-like, mycelium internal as well as somewhat external,
conidia falling off singly, sometimes producing short chains.

Leveillula

354 CLASS ASCOMYCETEAE

Appendages more or less stiff, dichotomously forked at tip.

Microsphaera

Appendages hooked or coiled at tip.

Mycelium entirely external, conidiophores straight. Uncinula

Mycelium internal as well as external, conidiophores with spirally twisted
base. Uncinulopsis

Appendages with swollen base, straight and pointed, mycelium external and

in substomatal chambers. Phyllactinia

Appendages wanting, surface of perithecium without gelatinous cells.

Brasiliomyces
(Viegas, 1944)
True equatorial appendages wanting, on upper surface of perithecium numer-
ous gelatinous penicillate cells. Typhulochaete
Ascospores two-celled.

Appendages hypha-like. Chileniyces

Appendages dichotomously forked at tip. Schistodes

Appendages lacking, mycelium and spores cinnamon-yellow. Astomella
Ascospores four-celled, appendages hypha-like. Leucoconis

Key to the Commoner Genera of Family Meliolaceae

Perithecia or mycelium intramatrical.

Perithecia subepidermal, asci many-spored, spores hyaline, two-celled.

Pampolysporium

Perithecia extramatrical, hyaline, mycelium in the epidermis but breaking out

as a superficial dark mass. Spores two-celled, brown, perithecia scattered.

Alina
Perithecia and dark mycelium external but emerging in strands through the
stomata.
Perithecia on peg-like strands from the stomata. Stomatogene

Mycelium superficial but penetrating the stomata in narrow bundles. Perithecia
on short mycelial branches. Piline

Perithecia and mycelium entirely external.

Mycelium with hyphopodia but no bristles. Irene^

Mycelium with hyphopodia and bristles. Meliola

Mycelium without hyphopodia.

With bristles, spores brown, two-celled. Phaeodimeriella

With bristles, spores brown, four- to five-celled. Meliolina

Without bristles, spores hyaline, two-celled Dinierina

Without bristles, spores brown, two-celled, perithecium rust-colored.

Parodiopsis
Without bristles, spores brown, two-celled, perithecium black.

Dimerium

Key to the Commoner Genera of Family Englerulaceae

{Based on Theissen and Sydow, 1917, but see Petrak, 1928)

Perithecial walls of rounded, soft cells, deliquescing into slime in which these cells
are scattered.

» Stevens (1927) segregates Irenopsis and Irenina from the genus Irene on some
minor characters, and also includes in this family (Meliolaceae) Adinodothis and
Amazonia which are placed by Theissen and Sydow in Order Hemisphaeriales.

KEY TO THE FAMILIES OF ORDER ASPERGILLALES 355

Perithecia sessile, with many asci.
Mycelium with hyphopodia.

Spores two-celled, colorless. Schiffnerula

Spores two-celled, brown. Phaeoschiffnerula

Mycelium without hyphopodia, spores two-celled, brown

Englerula
Perithecia stalked, with one ascus, spores two-celled, brown

Thrauste
Perithecial walls of parallel hyphae which separate at maturity. Mycelium exten-
sive, with hyphopodia, ascospores two-celled, brown.

Parenglerula

Key to the Commoner Genera of Family Capnodiaceae

Perithecia stalked or elongated vertically, wall of parallel hyphae.

Mycelium with parallel walls, forming a thick spongy mass, perithecium long-
stalked, round, spores four-celled. Scorias
MyceUum of moniliform hyphae, perithecia vertically elongated, sessile or
stalked, spores muriform. Capnodiwn
Perithecia sessile or on a very short stalk, spherical, walls mostly of monilioid
hyphae.
Bristles on mycelium or perithecia.

Perithecia without bristles, spores two-celled, hyaline. Tangled bristles on

the mycelium which has no hyphopodia. Dimerosporina

Perithecia without bristles, spores two-celled, brown, ascus single; short,

dark bristles on the myceUum which has hyphopodia. Balladyna

Perithecia with several asci, spores four- to more-celled, hyaline, bristles on
perithecia or myceUum or both. Chaetothyrium

Bristles lacking.

Spores several-celled, colorless. Limacinia

Various other genera based on form and color of spores, etc.

Key to Some Genera of Family Trichothyriaceae

Forming ribbon-Uke strands of hyphae covering the myceUum of the host fungus.
Spores colorless, two-celled. Trichothrjrium

Spores colored, two-celled. Trichothyriella

Spores colorless, three- to four-celled. Trichothyriopsis

Mycelium disappearing early, on stromata of fungi, spores colorless, two-celled.

Loranthomyces

Key to the Genera of Family Atichiaceae

Propagula clustered in basket-like structures. AticMa

Propagula in separate pockets. Phycopsis

Key to the Families of Order Aspergillales

Ascocarps subterranean, 1 cm. or more in diameter, cortex of many cell layers,
surrounding a "gleba" of irregularly arranged asci, with a central sterile por-
tion. Conidia not known. Family Elaphomycetaceae

Ascocarps not subterranean, external or in some cases buried in the substratum,
mostly not over 2-4 mm. in diameter.

356 CLASS ASCOMYCETEAE

Ascocarps with peridium consisting of loosely interwoven hyphae, conidia

formed in chains. Family Gymnoascaceae

Ascocarps at maturity with thin cortex or more stroma-like with a thick cortex

surrounding one or more glebal masses of asci. Conidia mostly catenulate.

Family Aspergillaceae
Ascocarps stromatic at base, the mass of asci and ascospores pushing out of the

top as a columnar structure. Conidia not known.

Family Trichocomaceae
Ascocarps stalked, with a spherical head which dehisces variously at maturity

to release the spores. Conidia not known. Family Onygenaceae

(If Nannfeldt (1932) is followed the two families below must be added to this key.)
Perithecia thin-walled, with long slender ostiolate necks, from whose apex the

hyaline ascospores are exuded in a slimy drop. Family Ophiostomataceae
Perithecia thin-walled, without neck, covered with numerous long, dark-colored,

variously branched or coiled hairs. Ascospores dark and extruded in a slimy

drop. Family Chaetomiaceae

Key to the Commoner Genera of the Family Aspergillaceae

Perithecia without neck or ostiole.

Perithecia without distinct hair coating or appendages, at least at maturity.

Perithecia small, usually bright-colored, at maturity reduced to a thin cortex,

containing the ascospores freed by the dissolution of the internal tissues

and the eight-spored asci. Ascospores biconvex, with a grooved edge, often

with two ridges. Conidial stage Aspergillus. Eurotium or

Aspergillus

Perithecia small, bright- or dark-colored, of two types: sclerotial with thick,

firm cortex with the central portion occupied by the eight-spored asci and

with ascospores as in the preceding, or with loose, hyphal cortex and the

ascospores without marginal groove. Conidial stage Penicilliurn.

Penicillium
Perithecia small, at maturity with thin cortex, at first containing many eight-
spored asci, but these and the surrounding tissues dissolve so that the
spherical or ellipsoidal hyaline or red ascospores lie free. Mycelium red or
purple in mass. Conidia single or in short chains. Monascus

Perithecia 0.5-1 mm. in diameter, with a thick, firm cortex containing numer-
ous ovoid asci, each with eight spherical, roughened, yellow to brown asco-
spores. Conidial stage Gliocladvum. Lillipuiia
Perithecia stalked, 2-3 mm. in diameter, with firm cortex, and containing
several irregular, gleba-like masses of sporogenous tissue. Asci two- to
eight-spored, ascospores with several ridges running in different directions.
Conidial stage resembling Penicilliurn or Sterigmatocystis. Tropical.

Penicilliopsis
Perithecia with distinct hair coating or coiled appendages.

Spherical or depressed, with simple or club-shaped hairs. Cortex firm and
thick. Ascospores small, ellipsoidal or lenticular, colored.

Cephalotheca
Depressed, often bluntly angled, with long appendages at each corner, coiled
at the tip. Magnusia

Perithecia with neck or ostiole. Doubtfully belonging to this order. Here could
be placed Microascus and Emericella if they are transferred from the Order
Sphaeriales.

KEY TO THE GENERA OF FAMILY ELAPHOMYCETACEAB 357

Key to Some Genera of Family Myriangiaceae*

Parasitic upon scale insects, but possibly eventually entering and parasitic upon
the tissues of the plant host. Basal stroma well developed, bearing one or
more disk-like ascigerous portions (ascomata) throughout whose tissues
the asci are scattered in no definite layers. On stems and leaves of various
plants, largely tropical.
Ascomata usually several; ascospores strongly muriform, hyaline or only pale-
colored. Myriangium
Ascoma usually single, with a narrower base; ascospores several times trans-
versely septate, occasionally one or two cells longitudinally divided,
hyaline or only pale-colored. Kusanoa
Parasitic within the tissues of leaves, stems, and fruits, or on their hairs, or in some
cases possibly parasitic upon fungi within the plant hosts.
Attacking only the hairs of the host plants, forming little ascomata, within
which only a few asci are produced ; ascospores mostly muriform,
Ascoma globular, with homogeneous tissue. Molleriella
Ascoma globular, strongly gelatinized except a few traversing brown hyphae.

Nostocotheca
Ascoma flattened. Saccardinula

Attacking the synnemata of Helostroma, and forming a disk-like or pulvinate
ascoma, attached by a narrow base. Ascospores strongly muriform and
dark-colored when mature. Cookella

Parasitic within the tissues of the host plant.

Ascigerous portion strongly developed outside the host tissue, spreading from

a narrower base. Uleomyces^

Ascigerous portion not strongly di&tinct from the internal stromatic tissue,
sometimes barely rupturing the epidermis, or forming a well-developed ex-
ternal cushion. Asexual reproduction by acervuli characteristic of the form-
genus Sphaceloma. Elsinoe (Plectodiscella)

Key to the Genera of Family Gymnoascaceae

Hyphae of the ascocarp without appendages.

Ascospores hyaline, yellowish or red. Arachniotus

Ascospores brown to brown violet. Amaurascus

Hyphae of the ascocarp with appendages.

Appendages consisting of spines or prongs. Gymnoascus

Appendages circinate. Myxotrichurn

Appendages comb-like. Ctenomyces

Key to the Genera of Family Elaphomycetaceae

Ascocarps with sterile base. Ascoscleroderma

Ascocarps without sterile base. Elaphomyces

* This tentative key is modified from Theissen and Sydow (1917) and Arnaud
(1925). Further studies are needed to reveal whether the many genera ascribed to
this family are valid.

5 The genera Myriangina, Myriaruiinella, and Kusanoopsis are probably closely
related to Uleomyces and Elsinoe and perhaps all five should be united in the one
genus Uleomyces

358 CLASS ASCOMYCETEAE

Key to the Families of Order Saccharomycetales and Appended Fungi

Asci formed, at least under special conditions.

Under normal conditions producing a mycelium of uninucleate or multi-
nucleate cells.
Asci containing from one to eight ascospores.

Family Endomycetaceae
Asci containing very numerous ascospores.

Family Ascoideaceae
An alternation of haploid and diploid generations.

Family Spermophthoraceae
Under normal conditions producing a yeast type of growth.

Cells dividing by fission. Family Schizosaccharomycetaceae

Cells multiplying by budding. Family Saccharomycetaceae

(The following key is based upon Lodder, 1934.)
Ascus formation unknown (Asporogenous Yeasts).

No violently expelled aerial spores formed (baUistospores)
Without carotinoid pigments

Cells reproducing by budding, but also producing aerial conidia on short

conidiophores or sterigmata Family Nectaromycetaceae

Cells reproducing by budding, but aerial conidia lacking. In some cases
short chains of buds formed or even hyphae. Many parasitic in Man and
other animals. Family Torulopsidaceae

Cells with carotinoid pigments, reproduction mostly by budding, no true
conidia or hyphae Family Rhodotorulaceae

Aerial conidia formed which are violently expelled, usually also budding
present Family Sporobolomycetaceae

Keys to the Commoner Genera of Saccharomycetales and Appended

Fungi

Key to the Commoner Genera of Sporogenous Yeasts Included in Families
Endomycetaceae, Schizosaccharomycetaceae, and Saccharomycetaceae

(Based upon Stelling-Dekker, 1931)

Spores spindle-shaped.

Only one spore per ascus. Parasitic in Arthropods. Monosporella

At least four spores per ascus.

Spores with a nonmotile, flagellum-like extension at one end. Parasitic in
plant tissues.
Mostly growing in yeast-like colonies. Nematospora

Mostly growing as hyphae ; ascospores formed in the terminal portion of a
hyphal branch, not separated by a septum from the remainder of the
hyphae. Ashbya

Spores without flagellum-like extension.

Parasitic in plant tissues, forming hyphae. Eremothecium

Parasitic in insects, hyphal formation lacking. Coccidiascus

Spores not spindle-shaped.
Producing true hyphae.

No production of oidia or budding cells. Asci mostly eight-spored.

Eremascus
Oidia produced, asci mostly with four or fewer spores. Endomyces
Asexual reproduction by budding, rarely by oidia. Endomycopsis

KEYS TO THE COMMONER GENERA OF SACCHAROMTCETALES 359

Normally not producing hyphae or only meagerly.

Asexual reproduction by fission. Schizosaccharomyces

Asexual reproduction by budding.

Spores hat-shaped (see also species of Endomyces) .

Buds produced from all sides of the cells. Hansenula

Budding bipolar, i.e., from opposite ends of the cells.

Hanseniaspora
Spores round with weakly or strongly verrucose walls, one, rarely two per
ascus.
Buds produced from all sides of the cells. Debaryomyces

Budding bipolar (mostly), buds with a narrow base. Spores with an
equatorial ridge. No completed conjugation, though rudimentary
conjugation tubes are produced. Schwanniomyces

Budding bipolar, but with a broad base. Spores without equatorial ridge.
Zygotes produced by union of a mother and a daughter cell.

Nadsonia
Spores round to oval, smooth, one to four per ascus. Conjugation taking
place between pairs of ascospores in the ascus. Vegetative cells large,
with bipolar buds with broad base. Saccharomycodes

Spores round to oval, smooth, one to four per ascus. Conjugation occurring
immediately before ascus formation or between vegetative cells in
colonies arising from germination of the spores. Not forming a thick
surface layer on beer wort. Fermentation of glucose marked.

Saccharomyces
Spores round to angular, one to four per ascus. Conjugation before ascus
formation or unknown. Forming a thick surface layer almost imme-
diately on beerwort. Fermentation feeble or lacking.

Pichia

Key to the Commoner Genera of Family Torulopsidaceae

(Based upon Lodder, 1934; Diddens and Lodder, 1942)

Not forming true mycelium. Cells mostly single or in irregular groups.

Subfamily Torulopsoideae
Cells prevailingly limoniform, with bipolar budding. Kloeckera

Cells prevailingly triangular, budding at the three angles.

Trigonopsis
Cells prevailingly flask-shaped, budding mainly at the broader basal part.

Pityrosporum
Cells round, ovoid or cylindrical.

Producing a thick surface growth on beerwort. No fermentation of glucose.
Buds often separating from mother cells by fission. Schizoblastosporion
Buds separating from mother cells in the usual manner.

Mycoderma
Not producing a thick surface growth on beerwort. Fermentation of glucose
in some species, no fermentation in others. Torulopsis

Forming pseudomycelium or true mycelium. Many species pathogenic to
animals.

Subfamily Mycotoruloideae
Pseudomycelium or true mycelium, reproduction by budding.

Cells often abruptly pointed, under aerobic conditions strong acid produc-
tion. Brettanomyces

360 CLASS ASCOMYCETEAE

Cells not pointed, under aerobic conditions not strongly acid producing.
Chains of yeast-like buds produced at the ends of the hyphae and single
cells or clusters at the joints of the hyphae. Many species are pathogenic
to animals. Candida

Crown-like or branching clusters of yeast-like cells produced from the cells
of the pseudomyceUum. Mostly pathogenic to animals.

Blastodendrion
In addition to budding cells, arthrospores (conidia) are also produced. Mostly
pathogenic to animals. Trichosporon

Keys to Other Families Possibly Related to Yeasts

Family Nectaromycetaceae

Only genus. Nectaromyces

Family Rhodotorulaceae

Only genus. Rhodotorula

Family Spermophthoraceae

Only genus. Spermophthora

Family Pericystaceae

Only genus. Pericystis

Family Sporobolomycetaceae

Forming yeast-like colonies reproducing mainly by budding, rarely few short
hyphae.

Cells with red or salmon-colored pigment. Sporobolomyces

Cells pallid to yellowish. B idler a

Forming hyphae, yeast-Uke colonies, reproduction by budding wanting.

Mycelium without clamp connections, spores falcate. Tilletiopsis

Mycelium with clamp connections, spores not falcate, reniform or ellipsoid or
ovoid. Itersonilia

Forming hyphae with clamp connections, yeast-like Ijudding present in the
younger colonies. Sporidiobolus

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LITERATURE CITED

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362 CLASS ASCOMTCETEAE

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LITEKATUEE CITED

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VON Hohnel, Franz: Atichia treubii v. Hohnel (Saccharomycetes), Ann. Jardin
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364 CLASS ASCOMYCETEAE

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CLASS BASIDIOMYCETEAE: SUBCLASS
TELIOSPOREAE

Introduction

IN CONTRAST to the Condition in the CUass Ascomyceteae the ultimate
reproductive spores in the Basidiomyceteae are produced externally
upon usually terminal cells within which caryogamy and subsequent
meiosis produce successively a diploid and then four haploid nuclei. In the
Ascomyceteae the haploid nuclei formed as a result of the meiotic divi-
sions of the diploid nucleus become the centers of ascospores formed
within the ascus. The difference is apparently that in the one class the
spores are produced internally but in the other externally. In reahty even
in the latter case distinct spore walls are sometimes produced around the
spores within the pocket-like outgrowths from the main cell so that the
basidiospore is an internally produced spore surrounded by and usually
adhering to the wall of the projecting pocket from the basidium.

(Fig. 119.)

The basidiospores or sporidia throughout the class are universally
unicellular when first formed and usually uninucleate (except in the more
unusual cases where two nuclei enter the spore from the basidium). In
many species the nucleus divides within the spore before the latter is set
free. In Dacrymyces and some other genera the basidiospore becomes
transversely septate into several cells at or shortly after maturity. In
color the spores in this class vary from hyaline (the majority of species)
to pink or to yellow-brown, brown, purple-brown, and black.

Typically in the Ascomyceteae there are various means by which the
ascospores are expelled from the ascus, thus providing for their distribu-
tion. Similarly in most of the Basidiomyceteae where the spores are
produced in the open air there is a special provision for the discharge of
these spores from their points of attachment. The spores are usually dis-
charged with considerable violence, usually all four spores being dis-
charged successively at intervals of several seconds to several minutes,

366

INTRODUCTION

367

\o No

r>,

°tz)

Fig. 119. Class Basidiomyceteae. Diagrammatic sketches comparing the nuclear
behavior in the formation of the ascus and the basidium and the further steps leading
to the production of the ascospores and basidiospores, respectively. (A) Development
of the ascus and ascospores. (B) Development of the basidium and basidiospores. (a)
Dicaryon terminal cell of a hypha. (b) union of the two nuclei to form a diploid nucleus;
(c, d) as the cell enlarges the first and second meiotic divisions occur, the daughter
nuclei in each division being indicated by a connecting line; (e) ascospores are formed
around each nucleus in the ascus; (f) external pockets are formed near the apex of the
basidium; (g) a wall is formed surrounding the cytoplasm and nucleus and separating
them from the sterigma. Spores are now mature.

according to the observations of Buller (1909). The distance to which the
spores are discharged may be ten to twenty times the length of the spore.
By affixing a clean glass coverslip a short distance above the surface of the
mature hymenium, e.g., a gill of a mushroom, the basidiospores will
sometimes be found sticking to the glass in groups of four. In this manner
it is possible to obtain and make cultures from the four spores arising
from a single basidium. Buller has made very extensive studies of this

368 CLASS BASIDIOMYCETEAE

discharge of basidiospores and its relation to the distribution of the spores
by air currents. (Figs. 120, 121.)

In all cases where the basidiospores are expelled violently Buller has
pointed out that they seem to be attached somewhat obliquely to the
sterigmata. Just before the spore discharge a drop of liquid begins to
appear at one side of the point of attachment, attaining full size in 5 to
20 seconds. It is worthy of note that in Sporoholomyces the spores are
discharged in the same way. This mode of attachment and release is
entirely different from that of true conidia (except conidia of Tilletiaceae) .

In the Ascomyceteae the vegetative mycelium is mainly of the mono-
caryon type, with the haploid nuclei one to a cell or at least not closely
associated by twos. The ascogenous hyphae on the contrary have two
nuclei to each cell, i.e., they are dicaryotic. These alternating types of
mycelium may be called primary and secondary respectively. Mainly the
latter is short-lived and limited in growth. However, in the Taphrinales
the main vegetative mycelium is of the secondary type, the primary
mycelial stage being very much abbreviated. In the Basidiomyceteae the
secondary "dicaryon" type of mycelium is more often the preponderating
stage, the primary mycelium usually being of much shorter life than the
secondary mycelium.

In very many of the Ascomyceteae there are definite male and female
reproductive structures (antherids and oogones, respectively). Yet in
some groups sexuality is on the decline and then these special reproductive
organs may be substituted for by the union of vegetative cells. This.latter
condition is the more frequent in the Basidiomyceteae where only in the
Rusts (Uredinales) are special female receptive hyphae produced. Even
in this group contact of vegetative hyphae of compatible types is suffi-
cient for sexual reproduction. In the Rusts special male gametes (sperm
cells or spermatia) are formed in definite organs called spermogonia,
but only on monocaryon mycelium. In many other members of the
Basidiomyceteae there are produced small hyaline, one-celled and uni-
nucleate spores called oidia. They arise on oidiophores and as they are
set free they cling together in a drop of sticky liquid, forming little shining
balls. The oidiophores may be simple or branched. The branches or the
apical portions of the unbranched oidiophores break up into oidia succes-
sively from the tip toward the base, leaving eventually only a short stub.
Brodie (1931), Vandendries (1924), and others have shown that mostly
the oidia arise only from primary mycelium. Long ago Dangeard (1894-
1895) described the formation of uninucleate oidium-like cells from the
dicaryon mycelium of Dacrymyces and more recently Vandendries and
Martens (1932) have demonstrated their occurrence on the dicaryon
mycelium of Pholiota aurivella (Batsch ex Fr.) Quel. In this species from
the same secondary mycelium may be produced thick-walled binucleate

INTRODUCTION

369

I
A

Fig. 120. CUass Basidiomyceteae. Steps in the discharge of a basidiospore or of a
sporidium from its sterigma. (A-I) Subclass Heterobasidiae, Order Dacrymycetales,
Family Dacrymycetaceae, Calocera cornea Fr. (J-M) Subclass Teliosporeae, Order
Uredinales, Family Pucciniaceae, Endophyllum euphorbiae-sylvaticae (DC.) Wint.
(Courtesy, Buller: Researches on Fiuigi, London, Longmans, Green and Co. A-I,
vol. 2, p. 7; J-M, vol. 3, p. 54.)

5 _4

Fig. 12L Class Basidiomyceteae. The successive discharge of four basidiospores
from one basidium of Agaricus campestris Fr. (Courtesy, Buller: Researches on Fungi,
London, Longmans, Green and Co., vol. 1, p. 52.)

370

CLASS BASIDIOMYCETEAE

Fig. 122. Class Basidiomyceteae, Coprinus lagopus Fr. (A) Monocaryon mycelium
bearing heads of oidia in mucilaginous drops. (B) Detail of oidiophore. (C) Oidia
(a, c) of one sexual phase fusing with monocaryon hyphae (6, d) of the opposite
(compatible) sexual phase. (Courtesy, Brodie: Ann. Botany, 45(178) :3 15-344.)

"gemmae" and thin-walled binucleate "oidia," both of which give rise to
dicaryon mycelium, and also uninucleate oidia. The latter begin as
binucleate, spindle-shaped cells which become divided by a septum into
two uninucleate cells which fall apart to form the oidia. These latter upon
germination give rise to monocaryon mycelium. The sticky droplets con-
taining the oidia attract insects. Brodie (1931) has demonstrated that
small flies carry these oidia from monocaryon mycelium to monocaryon
mycelium in Coprinus while Craigie (1927) demonstrated the same thing
for Rusts. (Fig. 122.)

In many of the Basidiomycetae the fungus occurs in two sexual
phases. It was shown by Vandendries (1924) and by Brodie (1931)
that the oidia from one phase are able to fertilize the mycelium of the
opposite phase as Craigie (1931) proved for the Rusts. The germinating
oidium unites with a cell of the monocaryon mycelium and thus initiates
the dicaryon stage of growth. This dicaryon cell may grow out by elonga-
tion and division into a dicaryon hypha or the nucleus introduced into this
cell by the oidium divides and one of the daughter nuclei passes through
the wall into the next cell and so on until the whole hypha is " diploidized "
as demonstrated by Lehfeldt (1923) and by Buller (1930). The latter made

INTRODUCTION

371

-P

E Ce»iQ»i»e"T

Fig. 123. Basidiomyceteae. Diploidization of monocaryon mycelium. (A) Two
monocaryon mycelia, mutually compatible, have united and diploidization has been
initiated in one cell. (B) Diagrammatic representation of the steps (1-6) in the
diploidization of a monocaryon mycelium by a dicaryon one. (Courtesy, Buller:
Nature, 126(3183) :686-689.)

experiments which showed that the rate of progression of the diploidizing
nuclei through the mycehum was 1.2 to 1.5 mm. per hour in Coprinus
lagopus Fr. When the introduced nucleus or one of its descendants arrives
in the terminal cell of the hyphal branch the two nuclei divide by con-
jugate division and thenceforth a typical dicaryon mycelium is produced
from this hypha. The initiation of the dicaryon phase does not depend
alone upon the oidia. Two monocaryon hyphae of opposite sexual phase
may unite and the result is the same as when an oidium unites with a
hyphal cell. Brodie demonstrated that the oidia of Coprinus lagopus are
capable of germinating and forming a mycelium made of very slender
monocaryon hyphae. From these arise very numerous oidiophores and
heads of oidia. When two such oidial mycelia of opposite sexual phase
meet diploidization occurs. Also the oidia from one mycelium may
fertilize the oidial mycelium of the opposite sexual phase. Possibly the
oidia and the oidial mycelium may represent a residual male sexual
structure while any cell of the normal monocaryon vegetative mycelium
possesses a female tendency, this not being restricted merely to an oogone
as in many of the Ascomyceteae. Vandendries and Brodie (1933) find
however that in some Basidiomyceteae the oidia are also capable of func-
tioning as conidia, producing a typical monocaryon mycelium which is
capable of diploidizing and being diploidized by a monocaryon mycelium
of appropriate sexual phase. (Fig. 123.)

The disappearance of definite sexual organs has not done away with
the three fundamental phenomena of sexual reproduction, cytogamy,
caryogamy, and meiosis, nor with the modification into various types of
compatibility and incompatibility of sexual strains such as occur in many
Ascomyceteae. Indeed, it seems possible that this development of
sexually compatible and incompatible strains has become greatly in-
creased in the Basidiomyceteae.

372

CLASS BASIDIOMYCETEAE

Very frequently, but by no means universally, associated with the
secondary type of mycelium is the production of clamp connections.
These are produced more often at every septum in the more slender
hyphae but may be absent on a portion of the same hypha where the cells
are broader. This has been frequently noted by the author in the tissues
of the pilei of various Agaricaceae, where slender hyphae bearing clamp
connections may as they elongate produce a series of broader cells without
these structures and perhaps still further on becoming slender again
bearing clamp connections. On the other hand even these broader hyphae
may show clamp connections, as illustrated in some of the figures by
Ktihner (1926) though other figures show the broader hyphae without

+ y* o

Fig. 124. Class Basidiomyceteae. Diagram-
matic representation of the steps in the for-
mation of clamp connections. (Courtesy, Ben-
saude: Recherches sur le cycle evolutif et la
sexualite chez les Basidiomycetes, Nemours,
published by author.)

such structures. Whenever clamp connections are present the mycelium
is thereby indicated as being dicaryotic, i.e., of the secondary type, but
from the foregoing it is evident that the absence of these structures does
not in all cases indicate the primary monocaryon nature of the hyphae.

The mode of development of clamp connections was reported inde-
pendently by Kniep (1915) and Mile. Bensaude (1918). The details of the
formation are as follows : The two nuclei of the terminal cell of a dicaryon
hypha lie a short distance apart in the longitudinal axis of the cell. Be-
tween them a lateral pocket is formed in the wall. The two nuclei now
divide simultaneously (conjugate division) and the lower daughter nu-
cleus of the upper pair passes into the pocket. This is now cut off from
the main cell by the formation of a septum. At the same time the upper
daughter nucleus of the lower pair, lying at about the level of the upper
end of the pocket becomes separated from its sister nucleus by a trans-

INTRODUCTION 373

verse wall. At this stage the terminal cell now has two nuclei, the cell
below it has one nucleus, while the fourth nucleus lies in the lateral pocket.
The latter curves around until it is in contact with the lateral wall of the
upper end of the penultimate cell and the intervening walls are dissolved
and the nucleus passes into the latter cell. The lateral pocket has acted
as a by-pass through which a nucleus has been transferred from the
terminal to the penultimate cell in such a manner as to provide each cell
with a daughter nucleus of each of the two nuclei originally in the terminal
cell. This by-pass is known as a clamp connection. Buller (1933) has
followed the formation of these clamp connections in living mycelium and
found that the process requires only a short time. In Coprinus lagopus Fr.
the time elapsed from the first appearance of the projecting lateral pocket
until the passage of the nucleus out of the pocket into the penultimate
cell was 23 minutes, while in C. sterquilinus Fr. it was 40 to 45 minutes.
The conjugate division of the nuclei was completed in the first species in
from 12 to 14 minutes. (Fig. 124.)

The phenomena involved in the formation of the clamp connections
are generally considered (e.g., by Kniep, Bensaude, and most students
since then) as being homologous to those occurring in an ascogenous
hypha when an ascus is forming by the hook or crozier method. The
Moreaus (1928) have shown that the formation of the hook does not
necessarily lead immediately to the formation of an ascus, for the terminal
binucleate cell may elongate and again form a hook while the tip of the
original hook unites with the cell below. This may continue several times
until a series of dicaryon cells is produced, each connected to the cell
below by a clamp connection. De Ferry de la Bellone (1886) and Mat-
tirolo (1887) described and figured typical clamp connections on the
mycelium of Tuber layideum Matt., and other species. Unless they mis-
took some intermingled strands of Basidiomycetous mycelium for that of
the fungi they were studying the occurrence of this structure in the
Ascomyceteae as well as in the Basidiomyceteae must be considered
substantiated.

Buller (1933) did not beheve that the formation of clamp connections
is at all homologous to the processes occurring in the ascogenous hyphae.
The clamp connections play, he beheved, an important part in the trans-
fer of food through the mycelium. An actual flow of protoplasm was
observed by him through the clamp connection and its centrally perfor-
ated upper septum. Possibly the occurrence of whorls of clamp connec-
tions in some fungi would support Buller's view as to their function in
nutrition. Kemper (1937) studied the development and cytology of the
clamp connections which may occur in whorls instead of singly at the
septa of Coniophora cerebella Pers. (now called C. puteana (Schum. ex Fr.)
Karst.). In this species the uninucleate young basidiospore becomes bi-

374

CLASS BASIDIOMYCETEAE

nucleate by the division of the nucleus before discharge. It gives rise to a
coenocytic mycelium with cross walls at rather long, intervals. The nuclei
divide apparently independently of one another and no conjugate divi-
sions can be noted. In the younger mycelium clamp connections arise
singly but as the mycelium becomes older they may arise by twos and
in still older mycelium in whorls. Their formation has no apparent con-
nection with nuclear divisions. From one to several nuclei may pass into
each pocket which fuses with the cell below the septum and discharges all
the contained nuclei into that cell, or a lateral branch may be formed into
which the nuclei pass. Sometimes the terminal cell of the hj^pha is left
without any nuclei. The author believes that these whorls of clamp con-
nections serve to distribute various types
of nuclei to the new branches. (Fig. 125.)
The greater prevalence of clamp con-
nections in the Basidiomyceteae is probably
due in the first place to the fact that in
most members of this class the dicaryon
mycelium represents a much greater por-
tion of the life history of the plant than
do the rather transitory or entirely wanting
ascogenous hyphae of the Ascomyceteae.
Furthermore, in the latter group these
hyphae are broader in general, so that at
conjugate division the two dividing nuclei
may lie side by side instead of some dis-
tance apart in the longitudinal axis of a nar-
row hypha. Only in the latter case is a by-pass really necessary. Some
whole genera of the Basidiomyceteae lack clamp connections entirely.

The prevalence of clamp connections varies greatly in different parts
of the same mycelium and is, furthermore, modified greatly by the en-
vironment. Thus mycelium submerged in liquid media may have but few
or even no clamp connections while the aerial portions may produce them
in abundance. They may be present on the slenderer hyphae m the pileus
of the mushroom and absent in the broader extensions or branches of
these same hyphae. In some species they are found only at great intervals
while in others they occur at every septum. In some species they are only
found in the subhymenial tissues of the spore fruit but not elsewhere, even
when all the tissues consist of dicaryon mycelium. Hirmer (1920) ob-
served conjugate divisions in the mycelium of Agaricus campestris Fr.,
although clamp connections were completely absent. Within the genus
Coprinus Brunswik (1924) has found some species which lack the clamp
connections entirely and other closely related species in whose mycelium
they are abundant. He interprets this as the gradual loss of a structure

Fig. 125. Class Basidio-
myceteae. Whorls of clamp con-
nections in Coniophora puteana
(Schum. ex Fr.) Karst. (After
Kemper: Zentr. Bakt., Para-
sitenk. und Infektionskr. , Zweite
Abt., 97(4-8) :100-124.)

INTRODUCTION

375

inherited from the Ascomycetous ancestors but whose function is no
longer indispensable.

Mile. Bensaude and Kniep were among the first to show the existence
of different sexual phases (or as they called it "heterothallism") in the
Basidiomyceteae. Miss Mounce (1922) and Miss Newton (1926) and
various other investigators have shown that many of this class are " homo-
thallic," i.e., will produce the dicaryon mycelium in culture from a single
basidiospore while other species are always "heterothallic." Rarely a
monocaryon mycelium of a heterothallic species after a considerable time
begins to produce dicaryon hyphae in a manner not yet satisfactorily
explained. In the homothallic species in which the basidiospore is uni-
nucleate it must be assumed that the genetic factors for incompatibility
are absent or are both present and mutually cancelling in the same
chromosome. Sass (1929) studied the behavior of the nuclei in certain
homothallic forms in which the nuclei are two in number in the basidio-
spore. Thus in Coprinus ephemerus Fr. there exist forms in which four
uninucleate basidiospores are produced on each basidium ; these forms are
heterothallic. In C. ephemerus forma hisporus only two basidiospores are
produced each with two nuclei. Mostly these give rise to homothallic
mycelia but sometimes they show heterothallism. Sass found that from
ninety per cent of such binucleate spores there is produced a coenocytic
mycelium which as it grows begins to become septate until the apical
portions of the hyphae consist of uninucleate cells. Some of these hyphal
branches on coming into contact with other uninucleate hyphae from the
same mycelium fuse with them to form typical dicaryon mycelium (with
clamp connections) and from this mycelium arise the normal fruiting
bodies. Evidently the two nuclei of the basidiospore in this case repre-
sented opposite sexual phases. In about ten per cent of the basidia the
two nuclei of the spore are clearly of the same sexual phase. Such spores
produce a mycelium consisting from the first of uninucleate cells. Only
when two such mycelia of opposite sexual phases come into contact is the
secondary, fruiting mycehum produced. The situations described by Sass
are very similar to those described by Ames (1932) in Schizothecium (see
Chapter 10).

The four basidiospores may represent two or sometimes four sexual
phases which are mutually fertile by twos. Because of the presence of male
sexual cells (oidia) on the mycelium of both uniting mycelia and of the
fact that each mycelium or its oidia can diploidize the other mycelium it is
manifest that we can not look upon these as representing opposite sexes.
As in homologous cases in the Ascomyceteae it is a question of self-
incompatibility, comparable in a way to that occuring within a given
horticultural variety of pear which may be sterile to its own pollen but
fertile to pollen of another (but not of every other) variety. Since the term

376 CLASS BASIDIOMYCETEAE

heterothallism as originally applied refers to mycelia representing differ-
ent sexes it seems inadvisable to use the terms heterothallic and homo-
thallic in these higher fungi, at least without qualification.

Apparently the factors governing the compatibility are two allelo-
morphic factors borne on different chromosome pairs. Kniep (1928) and
others have pointed out that only those unions of cells lead to diploidiza-
tion that bring about a combination that is heterozygous for both these
sets of factors. This need not involve a union of the nuclei into a diploid
nucleus, a process that occurs only in the basidium, but concerns merely
the bringing together, without union, of two haploid nuclei in the same
cell, so that so far as the cell is concerned diploidization has taken place
even though the two haploid nuclei remain separate for the present. Thus
a monocaryon mycelium with the compatibility factors A and B (these
being on separate chromosomes) could unite with another monocaryon
mycelium of the formula ab; similarly a mycelium with the formula aB
could unite with one of the formula Ab. In either case the formula for the
resultant cells would be AaBb, a condition heterozygous for both sets of
characters and fulfilling the requirements for the mating of the mycelia.
In the basidium the diploid nucleus undergoes two meiotic divisions to
consummate the reduction process. If both pairs of chromosomes bearing
the factors for incompatibility or compatibility undergo disjunction in
the first division it will be a matter of chance whether the resultant
daughter nuclei will be Ab and aB or AB and ab. Since each nucleus di-
vides again, this time by splitting the chromosome, there will be four
nuclei (one for each spore), two each of the formula Ab and aB or AB and
ab respectively. If the disjunction does not occur in either chromosome
pair until the second meiotic division the first division may represent
merely the splitting of the chromosome so that the two daughter nuclei
will be like the parent nucleus with the formula AaBb. When these divide
by the disjunction division it will give this time Ab and aB or AB and ab,
for each of the two nuclei. If the separation of the chromosomes or the
chromosome pairs occurs in the same direction in both nuclei there will
result four nuclei, alike two by two, but this separation may be in opposite
directions in the two dividing nuclei for one of the chromosome pairs so
that the four nuclei may all be different. If disjunction of one chromosome
pair occurs at the first division and of the other pair at the second division
the four resultant nuclei will also be all different, viz., Ab, aB, AB, ab.
Miss Newton (1926) showed that in Coprinus lagopus all of the foregoing
arrangements may be found in the various basidia of the same hymenium.
Apparently disjunction occurs more often at the first division for both
sets of chromosomes, for in the majority of cases two of the basidiospores
of a given basidium will be of one sexual phase and the other two of the
opposite phase. In some species only two sexual phases appear to occur.

INTRODUCTION

377

These are spoken of as possessing bipolar sexuality in contrast to those
species with quadripolar sexuality such as described above. In a bipolar
species only one allelomorphic pair of incompatibility factors need be
assumed.

Not infrequently a monocaryon mycelium may produce spore fruits
but in this case the basidia are less numerous and are either sterile or
give rise to but two, uninucleate basidiospores, both of the same sexual
phase as the parent mycelium. In Coprimis fimetarms Fr., Oort (1930) has
shown that two monocaryon mycelia may intermingle and together build
spore fruits when they represent phases alike genetically for one factor
and heterozygous for the other, e.g., Ab and AB. Such fruits are not
normal but may produce two kinds of two-spored basidia which are about
equally divided between the component phases, in this case Ab and AB.
It is apparent that no true sexual union has occured in such a case. That
this is more than a simple intermingling of separate monocaryon mycelia
is demonstrated by the fact that clamp connections may be formed, vary-
ing from incomplete or abnormal structures to those of perfectly normal
appearance. They vary from only occasional to frequent. The external
conditions appear to have considerable effect upon the number of clamp
connections produced. These unions do not arise in all incompatible com-
binations. Vandendries and Brodie (1933) showed that in Hypholoma
candolleanum (Fr.) Quelet, a quadripolar species, the mycelia to which
they ascribe the formula ab' when mated with a'b', or ab when mated
with a'b cause mutual partial inhibition of growth so that when growing
close together they are much smaller than when grown apart or in the
combination a'b X a'b'. It should be noted that the formulae ab, a'b, ab',
and a'b' correspond to AB, aB, Ab, and ab in the preceding portion of the
paragraph. The combination studied by Oort which produced a spore fruit
corresponds to the formula a'b X a'b'. Oort also found that such spore
fruits were not produced in certain other combinations. Quintanilha
(1935) made genetic and cytological studies of normal and "illegitimate"
crosses in Coprinus fimetarius and found that nuclear fusions may occur
in the basidia even in the latter type of cross.

Vandendries and Brodie (1933) and Vandendries (1934) and Brodie
(1934, 1936) described what they termed "barrage sexuel." This had been
noted before by Oort (1930) and by Brunswik (1924) but not studied
intensively. This amounts to a mutual repulsion of the hyphae of some of
the incompatible matings. When two such cultures are established in the
surface of an agar medium the mycelia as they grow leave a gap between
the two colonies. This is especially marked in the aerial mycelium whose
hyphae show abrupt curvatures away when they approach the other
mycelium at a distance of 3 to 5 mm. These authors showed further that
the interposition of thin low plates of glass did not prevent this repulsion

378 CLASS BASIDIOMYCETEAE

nor did sheets of mica, very thin sheets of silver, lead, and other sub-
stances. Apparently the repulsion is due to some sort of emanation from
the mycelium, perhaps gaseous. Experiments showed that certain nuclear
combinations give these mycelial repulsions but not others. Thus the
combinations ab X ab' and a'b X a'b' show "barrage" but not ab X a'b
or ab' X a'b'. Clearly the repulsion is between the b and b' mycelia.
The repulsion was also demonstrated between dicaryon mycelium and
monocaryon mycelium and between dicaryon mycelia where the two
sets of nuclei were different, e.g., (ab + a'b') X (a'b + ab'). This phe-
nomenon of barrage as well as the inhibition of growth does not occur in
all species. Brodie (1935) showed that barrage or "aversion" occurs also
in some strains of fungi growing naturally in wood.

A peculiar phenomenon in connection with the occurrence of bipolar
and quadripolar sexual phases is that of "geographic races." This has
been studied in Coprinus by Hanna (1925) and by Vandendries (1924)
and extensively by Brunswik (1924) and in Ustilago by Bauch (1930,
1931). In some species of Coprinus the two sets of incompatibility factors
may be alike in fungi growing in the same locality but one or both sets
may be different in fungi (of the same species) growing in different local-
ities. In the latter case all sexual phases of one fungus would be compatible
with all sexual phases of the other fungus, while if only one pair is different
in the two fungi certain combinations will be incompatible. It is evident
that these are explained best on the hypothesis of multiple allelomorphs,
familiar to geneticists in both animals and plants. When either of the
factors produces an allelomorph by mutation a legitimate cross becomes
possible. Since these mutated factors are frequently discovered in fungi
from different geographic locations the term "geographic races" is often
used (see Kniep, 1928). In Ustilago longissima (Sow.) TuL, Bauch rec-
ognized over 15 allelomorphic strains of the A-a factors and about 8 of the
B-b factors. The resulting interfertility of all sexual phases of one collec-
tion of species with all sexual phases of another collection has been found
to be rather general. Thus Mounce and Macrae (1936) found complete
compatibility between collections made in different localities or on differ-
ent hosts in Gloeophyllum saepiarium (Wulf.) Karst. {Lenzites saepiaria
(Wulf.) Fr.) and in Trametes americana Overh., and Vandendries (1936)
showed its occurrence in Leptoporus adustus (Willd. ex Fr.) Quelet.
Barnett (1937) made similar observations for some of the Heterobasidiae.

The Class Basidiomyceteae may be divided into three subclasses:
Teliosporeae (as a class in the earlier work of the author), Hetero-
basidiae, and Eubasidiae. They may be distinguished as follows:

Teliosporeae : parasitic in the leaves, stems, f luits, and sometimes in the roots,
of Pteridophyta, Strobilophyta (Coniferae), and Anthophyta (Angio-
spermae). The septate parasitic mycelium is mostly intercellular in the host

SUBCLASS TELIOSPOREAE 379

and consists of primary (monocaryon) and secondary (dicaryon) phases. In
the latter certain cells enlarge and their two nuclei unite to form a diploid
nucleus. This cell is a teliospore. It usually has a thick, colored wall and
serves as a resting spore. With the proper environmental conditions this
teliospore germinates by sending out a thin-walled straight or curved, four-
celled or one-celled hypha, the promycelium, within which the zygote nucleus
divides meiotically to form four (or, by further mitotic division, more)
nuclei. Usually four sporidia are formed on the promycelium and into each
of these a single nucleus enters. These sporidia are shot off with violence
or (in the Family Ustilaginaceae) fall off, and give rise to the primary type
of mycelium or by union of two sporidia of suitable sexual compatibility the
secondary type of mycelium arises. The teliospores are formed within or
upon the host tissues in so-called sori. Various secondary types of (asexual)
reproduction are produced.

Heterobasidiae: mostly saprophytic, but in many cases parasitic upon various
types of Bryophyta or Vascular Plants. Mycelium septate, mostly falling
into two phases, the monocaryotic and dicaryotic. From the latter are pro-
duced the spore fruits characteristic of the subclass. The basidia are usually
produced in a hymenium, i.e., they stand side by side like a palisade, but
may be more scattered. They arise as enlarged binucleate cells whose nuclei
fuse. Only rarely does the basidium become dark-colored and thick-walled.
Usually the formation of the basidiospores takes place without much delay.
The basidia may be one-celled and forking (Dacrymycetales), transversely
four-celled (Auriculariales), longitudinally four-celled (Tremellales), or one-
celled with four enlarged epibasidia bearing the basidiospores (Tulasnellales) .
Spore fruits are more often gelatinous when wet, or waxy. The basidiospores
often germinate by budding at many points in water. Possibly Sporoboloniyces
was derived from this subclass.

Eubasidiae: Mostly saprophytic but some species parasitic. Very often living
in the wood of living or dead trees. Basidia one-celled, without epibasidia
but bearing usually four long or short sterigmata, each bearing a single
basidiospore which usually germinates by a germ tube, even when fallen
into water. Basidia produced with some exceptions in definite hymenial
layers which may be exposed to the air before the spores become mature
(the "Hymenomycetes"), or enclosed in the spore fruit until after the spores
are completely mature (the "Gasteromycetes"). In this subclass are found
the largest spore fruits of the whole Phylum Carpomyceteae (Higher Fungi).

Subclass Teliosporeae

The subclass Teliosporeae corresponds to Dietel's limits of subclass
Hemibasidii in the second edition of Engler and Prantl's Die Natiirlichen
Pflanzenfamilien (1928). These fungi have been sometimes called the
Brand Fungi. Usually the two orders Uredinales and Ustilaginales have
been considered to be more or less closely related. They were placed close
together by Fries (1832) as Ordo IV* Hypodermii,i and by Plowright
(1889) and by most of the later botanists. Bennett and Murray (1899)

1 It should be noted that Fries did not believe that these were true fungi but that
the spores arose by the transformation of the tissues of the host plant and were not
used for the propagation of the rust or smut.

380 CLASS BASIDIOMYCETEAE

separated the orders, placing the Ustilaginales in their Class Zygomycetes
while the Class Uredineae was placed between the two Classes Asco-
mycetes and Basidiomycetes, the affinities being considered to lie with
the former. Because of the fancied resemblance of the teliospores of some
Rusts to asci, Charles E. Bessey (1894) was inclined to include the
Uredinales and Ustilaginales in the Class Ascomyceteae, a position from
which he receded when the cytological phenomena of these groups became
better known. That they are rather closely related to the other Basidio-
myceteae the studies of Brefeld (1881 and later) and the cytological in-
vestigations of Sappin-Trouffy (1896), Harper (1898, 1902), and others
leave little doubt. That they stand apart from the majority of families of
that class is equally certain. The author beheves that the differences are
sufficiently great to warrant placing them in a separate subclass, a posi-
tion that does not deny their relationship within the Basidiomyceteae but
leaves each subclass much more homogeneous.

The fungi of this subclass are parasitic. In the Order Ustilaginales
some of the species are capable of saprophytic growth in media rich in
food; in the Uredinales growth is strictly parasitic and the fungi have
never been cultured except on the living tissues of the host. The mycelium
is long, slender, and branching, growing intercellularly within the host. In
the majority of cases studied occasional or frequent haustoria are pro-
duced. The cells of the mycelium are mostly uninucleate in one stage of
development (monocaryon stage) and binucleate in the remainder of the
life cycle (dicaryon stage). As the cells of the latter type of mycelium
divide the two nuclei divide simultaneously so that one daughter nucleus
of each of the original nuclei is found in each of the two daughter cells.
This is the type of nuclear division to which the term "conjugate divi-
sion" was given. Clamp connections occur in many of the Ustilaginales
but have been rarely reported in the Uredinales. Finally, on the dicaryon
mycelium are produced certain larger cells, usually terminal to a hypha
or its branches, which become thicker walled. Within these cells the two
nuclei unite to form the only diploid nuclei in the life cycle of the fungus.
These cells are the teliospores. In the Ustilaginales (Smuts) these are often
spoken of as "chlamydospores," a misuse of this name which is rightly
applied only to vegetative cells which become filled with food and develop
thick walls to permit survival over winter or through other unfavorable
environments. In true chlamydospores there is no nuclear fusion and their
germination is in the manner usual for asexual spores.

Plowright (1889) recognized tlfe essential homology of the teliospores
in the two orders. They have a typical manner of development. The
diploid nucleus divides by meiotic divisions into four nuclei, in the
teliospore, or more often the exospore ruptures and a thin-walled hypha
(promycelium) grows out into which the diploid nucleus passes. The

SUBCLASS TELIOSPOREAE 381

meiotic divisions then take place in the promycehum instead of the body
of the teliospore. In some cases the nuclei divide once or twice more
resulting in the production of 8 or 16 or even more nuclei. The promyce-
lium may remain nonseptate but more often becomes transversely septate
into four cells. From each of these cells is produced a sessile or stalked
sporidium or several such sporidia. From the nonseptate promycelium 4
to 16 or more sporidia bud out at the apex. If the teliospore is deeply
buried in the tissues of the host or among other teliospores the pro-
mycelium at first is a slender hypha, its emergent portion developing the
typical structure.

The sporidia are borne in the Uredinales and in Family Tilletiaceae
in the Ustilaginales at the tips of pointed sterigmata from which they are
flung violently at maturity. Their position is slightly asymmetrical at
the tip as is the case in the Heterobasidiae and in the Hymenomyceteae.

Many writers (e.g., Gaumann, 1926, Arthur, 1929, etc.) apply
the names basidium and basidiospore, respectively, to the struc-
tures here called by their older names promycelium and sporidium. The
latter is doubtless homologous to the basidiospore but the term basidium
must include both the teliospore, in which the nuclear fusion occurs, and
the outgrowth from it, the promycelium, in which the meiotic nuclear
division usually occurs and upon which the sporidia are borne. More
correctly, as will be seen further on in the next chapter, the teliospore is
probably homologous to the hypobasidium and the promycelium to the
epibasidium of some of the Auriculariales.

Asexual reproduction is known in both the Uredinales and the
Ustilaginales. In the latter it occurs by means of colorless, often spindle-
shaped or sickle-shaped conidia which are uninucleate or binucleate. The
latter arise only from dicaryon mycelium but the uninucleate conidia
may arise from either monocaryon or dicaryon mycelium. They arise on
very short conidiophores, or rather sterigmata, from the sides of the
mycelial cells. They are usually in the true sense of the words "repeating
spores" for they produce the same type of mycelium as that from which
they arose, except in the case of uninucleate conidia from dicaryon
mycelium. These give rise to monocaryon mycelium. These conidia are
mostly produced on the saprophytic mycelium, but in some species are
developed on sterigmata which emerge through the epidermis of the living
host. In the Uredinales the urediospores (uredospores) are dicaryon
repeating spores and are, therefore, true conidia. There are no mono-
caryon repeating spores in the Rusts. The aeciospores (aecidiospores) are
the result of a sexual fusion involving cells but not the nuclei. They bridge
over the step from the monocaryon phase to the dicaryon phase. They
are not strictly homologous to ordinary conidia but yet show great
resemblance to the urediospores.

382 CLASS BASIDIOMYCETEAE

Sexual reproduction occurs in two usually separated steps : the union
of two monocaryon cells to initiate the dicaryon mycelial phase and
eventually the union of the nuclei in the teliospores to form diploid nuclei.
The points at which the dicaryon phase may be initiated are quite variable
in the Smuts but much more definite in the Rusts. The details will be
taken up in the discussion of the respective orders.

Order Uredinales (The Rusts). These form a group which manifests
a very high degree of evolutionary development in many directions. The
7,000 or more species are all strict parasites of Ferns (Pteridophyta),
Conifers (Strobilophyta), and Flowering Plants (Anthophy ta) . Not only
are the Rusts strict parasites but in many cases they are highly specialized
into biologic races which are confined to certain species of a host genus or
even to special agricultural varieties of a host species much as in the case
of Erysiphe graminis. In Europe Eriksson (1894, 1902) was the first to
make extensive studies into physiologic races confined to related species.
He was followed by numerous other investigators on that continent. In
the United States Carleton (1899, 1904) was the first to follow up Eriks-
son's work, followed by Freeman and Johnson (1911), and by Stakman
(1914), and others. The study of the physiologic forms on special agri-
cultural varieties of these species has been carried out in this country
very extensively by Stakman and various collaborators (1922). For
example in Puccinia graminis tritici Erikss. & Henn., the physiologic race
that attacks common wheat (Triticum aestivum L.) and durum wheat
{T. durum Desf.), over 150 physiologic forms have been distinguished in
North America and many additional forms, mostly distinct from these, in
Europe. These distinctions were made by means of a study of their power
of infecting 10 or more differential varieties of wheat. Similar physiologic
forms have been found in leaf rust of wheat (P. ruhigo-vera iritici (Erikss.
& Henn.) Carl.) by Mains and Jackson (1926) in the United States,
Scheibe (1929) in Germany, and Sibelia (1936) in Italy. Stakman and
others (1928) have also found them in maize rust (P. sorghi Schw.).
Physiologic forms have also been distinguished in flax rust (Melampsora
lini (Pers.) Lev.) by Flor (1935), in Puccinia helianthi Schw. b}^ Brown
(1936), in P. iridis (DC.) Wallr., by Mains (1938), in Uromyces phaseoli
typica Arthur, by Harter and Zaumeyer (1941), etc.

With one or two exceptions the teliospores are produced within the
tissues of the host and remain internal or break out through the epidermis
separate and free or united together in a waxy mass or attached by stalks.
The sporidia are capable of wind distribution but are short-lived and
delicate so that they can be carried only short distances in a living condi-
tion. Two other spore forms are usually produced, the aeciospores and
the urediospores, both of which are relati\ ely thick-walled and capable
of remaining alive while being carried many miles by the wind (hundreds

ORDER UREDINALES (tHE RUSTS)

383

Fig. 126. Monocaryon and dicaryon mycelium and haustoria in Subclass Telio-
sporeae, Order Uredinales. (A) Cronartium ribicola Fischer, monocaryon phase in
tissues of Pinus strobus L. ; intercellular hyphal cells and haustoria uninucleate. (B)
Phragmidium rubi (Pers.) Wint., dicaryon phase in leaf of Rubus sp.; intercellular
hyphal cells and haustoria binucleate. (A, courtesy, Colley: /. Agr. Research, 15(12):
619-660. B, after Sappin-Trouffy : Le Botaniste, 5:59-244.)

of miles in some cases). In general in this order the teliospores remain
attached in the sorus until after the formation and discharge of the
sporidia. However, Pady (1948) has shown that in Puccinia tumidipes
Peck the very much swollen pedicel will absorb water and rupture, throw-
ing the teliospore some distance (2 to 4 mm.). It is probable that the same
phenomenon occurs in other species with enlarged pedicels, such as some
species of Phragmidium and Gyinnospor-angium.

The mycelium occurs in two well-marked alternating phases, the
monocaryon and the dicaryon phase. Clamp connections have been
reported by Voss (1903) on the mycelium of the aecial phase but are
apparently rare, or at least not readily demonstrated in thin sections.
Their presence was confirmed by Wang and Martens (1939) in close
proximity to the aecium of Puccinia coronata Cda. on Rhamnus frangula
L. They were demonstrated by maceration of the tissues, not by sections.
Haustoria are frequent, sometimes small and spherical, sometimes large
and branched, and often partially surrounding the nucleus of the host
cell. They are uninucleate or binucleate according to the type of mycelium
producing them. Colley (1918) showed for the White Pine Blister Rust
{Cronartium ribicola Fischer) that the haustorium does not truly pene-
trate the protoplast of the host cell but causes invagination. Miss Rice
(1934) demonstrated this also for Uromyces caladii (Schw.) Farl. Appar-
ently the host and parasite stand in a very perfect balance for a very
considerable time, especially in the case of rusts that are well adapted
to the host. The host cells are not killed outright in such cases. In some
of the rusts that are not well adapted to their host Stakman (1914)
demonstrated that the host cells are killed immediately around the site of

384 CLASS BASIDIOMYCETEAE

infection, thus isolating the parasite from the host tissues and causing the
early death of the fungus before it does much damage to the host. The
death of the affected parts of the host plant is gradual in most cases. In
some cases, e.g., the rusts of small grains, the splitting of the epidermis
by the numerous elongated sori of the rust seems to increase the water
loss of the host plant to a very detrimental degree. (Fig. 126.)

The promycelium of the Rusts is normally four-celled, each cell
having but one nucleus. It emerges through a thin spot in the tehospore
wall, the germ pore. In shape the promycelium is variable depending
upon the species. It may be long and slender and nearly straight, each
cell with a long sterigma, or short and thick and curved, constricted more
or less at the septa. On such a curved promycelium the sporidia always
arise on the convex side, often on rather short sterigmata. In a number of
genera the promycelium does not entirely emerge from the teliospore.
Thus in Zaghouania the swelling of the teliospore bursts its thick wall
allowing the emergence of a thin-walled, four-celled promycelium whose
basal portion still remains enclosed within the old cell wall. In Coleo-
sporium and Gallowaya the teliospore divides by cross walls into four cells
without emerging from the cell wall, thus producing a promycelium that
is entirely internal. From each cell a long slender sterigma grows up
through the gelatinous stratum that covers the tops of the layer of later-
ally adhering teliospores. Weir (1912) reported that occasionally in this
genus the four cells produced by the division of the teliospore are cru-
ciately arranged, resembling the condition in Tremella. In Chrysopsora
each teliospore of the two forming the stalked compound teliospore
divides transversely as in Coleosporium. In Gopla7ia dioscoreae (B. & Br.)
Cummins (1935b) the teliospores arise from the base of a gelatinous
matrix and push up into it but not through to the exterior. Each telio-
spore, as in Coleosporium, divides by transverse septa into four cells from
each of which a slender sterigma pushes out to the surface where the
sporidia are formed. This is very similar to the case in the gelatinous
species of Auriculariales.

The uninucleate sporidia may be long, ellipsoid, pointed or rounded
at one or both ends, or may approach a spherical shape. The cell contents
are usually somewhat yellow. The sporidium is shot off with more or less
violence from the tip of the sterigma. Usually the whole contents of the
promycelial cell pass into the single sporidium arising from that cell but
occasionally a case is met with where the cell nucleus divides and one
nucleus and part of the cytoplasm pass into the sporidium, so that a
second sporidium may be produced by the same cell, as occurs more
frequently in the Ustilaginales. A sporidium that fails to fall upon a
suitable host is capable, under proper conditions of moisture, of pro-
ducing a secondary sporidium at the tip of the sterigma and this in turn

ORDER UREDINALES (tHE RUSTS)

385

a tertiary or even quaternary sporidiiim, each being shot off from its
sterigma. The successive sporidia are smaller and smaller. Under condi-
tions of extreme humidity the author has observed the primary sporidia
and the succeeding ones of Kunkelia nitens (Schw.) Arthur remaining
attached in a short chain of four or five successively smaller cells.

When a sporidium falls upon the epidermis of a suitable host it germi-
nates in a drop of rain or dew or film of water, forming a slender germ tube
which, Dr. Ruth F. Allen (1930) showed, penetrates the cuticle and cell
wall into the epidermal cell. The actual pore of entry is very small, the
hypha on either side being several times as thick. Within the epidermal
cell the hypha elongates and becomes
divided into several uninucleate cells.
From each of these a branch grows
through the interior wall of the host
cell either into an underlying cell,
where it acts just as did the infection
hypha in the epidermal cell, or into
an intercellular space where growth
becomes much more rapid and the
hypha becomes larger and more vigor-
ous, sending haustoria into the cells
between which it passes. A few cases
have been reported where entry took
place through a stoma but these seem
to be not usual. Pady (1935) reports
that in the uninucleate race of Kun-
kelia nitens the sporidia upon germi-
nation infect intracellularly the epi-
dermal cells and from these continue

to progress as intracellular mycelium for ten days or so, producing then
in the phloem tissues of the host the intercellular mycelium which forms
haustoria that are coiled more or less. More often after the first two or
three cells are formed the production of septa leads to the formation of
uninucleate (monocaryon) cells. In Melanipsora lint (Pers). Lev. the cells
remain one to three or four nucleate up to the regions where the definite
reproductive cells are formed when monocaryon cells predominate (Allen,
1934a). The infected area of the leaf often becomes thickened, in part at
least through rapid formation of large amounts of mycelium which forces
the host cells apart and in some cases crushes them. The presence of the
rust may cause marked changes in the manner of growth of the host. In
normally prostrate species of Chamaesyce {Euphorbia) the infected shoots
become upright, a phenomenon also observed in other families of host
plants. Abnormal growth of axillary buds in infected shoots gives rise to

Fig. 127. Subclass Teliosporeae,
Order Uredinales. Infection of leaf of
Berberis vulgaris L. by sporidium of
Puccinia graminis Pers. Infection
hyphae are within epidermal cell and
the attached sporidium is empty.
(Courtesy, Allen: J. Agr. Research,
40(7):585-614.)

386 CLASS BASIDIOMYCETEAE

the witches' brooms so characteristic of some rust infections on Conifers.
Dodge (1923) showed that in leaves of Rubus infected by the orange
rusts {Kunkelia nitens and Gyntnoconia peckiana (Howe) Trotter) the
production of stomata which is usually confined to the lower epidermis is
also brought about in the upper epidermis. (Fig. 127.)

The monocaryon mycelium is of rather short duration in the leaves
and stems of herbaceous plants. In woody plants it may persist for years,
e.g., Cronartium rihicola Fischer, in the tissues of the white pine {Pinus
strohus L.) or Gymnoconia peckiana and Kunkelia nitens in Ruhus where
the mycelium penetrates to the roots and infects the new shoots next year.
From this mycelium arise the spermogonia (pycnia). These may be sub-
cuticular or subepidermal on leaves, green stems or fruits, or even sub-
cortical on woody stems. They consist of a basal pseudoparenchymatous
mass of uninucleate cells from which arise numerous parallel slender
uninucleate sporophores. In these the nucleus divides and the upper
daughter nucleus passes out into a terminal sperm cell (pycniospore) con-
taining very little cytoplasm and surrounded by a thin cell wall. This
spore is pushed loose by the formation of a second sperm cell below it, etc.
At the same time a sugary slime is secreted which partially or completelj^
fills the cavity of the spermogonium. The subepidermal spermogonia may
be more or less spherical structures with a marginal series of paraphyses
around the sporiferous portion. These push up through the epidermis,
rupturing it and producing an ostiole through which the sugary slime
containing the sperm cells exudes as a shining drop which is sweet in
taste. Sometimes the mass of spermogonia is fragrant. The more diffuse
subcuticular and subcortical spermogonia also rupture the overlying
cuticle or cortex respectively, exposing the sugary exudate. Various in-
sects, particularly flies, attracted by the sweet liquid and accompanying
fragrance visit the spermogonia and feed on the exudate and go from leaf
to leaf and plant to plant. In this way the sperm cells adhering to their
feet or mouth parts become scattered widely over various plants or other
parts of the same plant. Rain also doubtless helps in the dissemination of
the sperms. (Fig. 128.)

From the same mycelium that has given rise to the spermogonia and
often near to these or on the opposite side of the leaf the hyphae begin to
mass themselves, frequently at first in a substomatal chamber or other
large intercellular space. Eventually this becomes a pseudoparenchy-
matous mass of cells, those nearer the surface being larger and less filled
with food, the underlying cells being smaller and better supplied with
food, the whole mass being more or less surrounded by several layers of
hyphae. All the cells are normally uninucleate. This structure is called an
aecial primordium. Buller (1938) suggested for this structure the term
proto-aecidium. Massee (1888) reported the presence of an oogone with

Fig. 128. Subclass Teliosporeae, Order Uredinales. Various types of spermogonia.
(A) Subcuticular spermogonium in Phragmidiuin violaceum (Schulz.) Wint. (B)
Subepidermal spermogonium in Gynuwsporangium clavariaefonne (Jacq. ex Pers.)
DC. (C, D) Subcortical spermogonia. (C) In Cronartium comptoniae Arthur, section
of twig of Pinus sp. (D) Portion of spermogonium of Cronartium ribicola Fischer,
showing development of sperm cells. (A-B, courtesy, Blackman: Ann. Botany,
18(71) :323-373. C, after Adams: Penn. State Coll. Agr. Exp. Sta. Bull., 160:31-77.
D, courtesy, Colley: J. Agr. Research, 15(12) :619-660.)

387

388 CLASS BASIDIOMYCETEAE

attached antherid in this primordium. The more recent investigations
seem to show that this report is incorrect. In the dense portion of the
pseudoparenchymatous tissue there begin to appear cells with two or
more nuclei, whose origin will be discussed in the succeeding paragraphs.
These cells elongate and may branch. At the apex binucleate aeciospores
are budded off. After each spore is formed there is produced a smaller
"disjunctor" cell, likewise binucleate, then another aeciospore, etc., until
a chain of alternate spores and disjunctor cells results. Between the
original binucleate or plurinucleate cells others push their way in from
the basal side of the aecial primordium and these also give rise to spore
chains. In the meantime the large almost empty cells begin to collapse
and digest, forming a space into which the chains of spores are pushed.
Finally all of this tissue is crushed or destroyed. Around the sporogenous
area a peridium may be produced or it may be wanting. The mass of
spores eventually ruptures the overlying host tissues and the mass of
spore chains and loose spores is exposed to the air. The spores are usually
light to dark yellow or orange and somewhat roughened.

Craigie demonstrated in 1927 that for Puccinia graminis Pers., grow-
ing on barberry (^Berheris vulgaris L.) and P. helianthi Schw., growing on
sunflower (Helianthus) an infection with a single sporidium produces a
sorus within which are produced numerous spermogonia and aecial
primordia. If this is protected from the visits of insects the primordia
never develop to spore production. However, if insects are allowed free
access to this sorus and other sori as well, the aecia will develop within a
few days. Furthermore, by transferring the sperm cells from one sorus to
another in about half of the cases aecia will develop. Craigie finally
demonstrated that the four sporidia of a promycelium were of two sexual
phases, two sporidia of each phase, and that sperm cells from the sorus of
one sexual phase, would fertilize the sorus of the other sexual phase, and
vice versa. In later papers (1928, 1931, 1933) he has added P. coronata
Corda, P. pringsheimiana Klebh. (P. caricis grossulariata Arth.) and
Gymnosporangium sp. to the list. Miss Allen (1932a) showed this to be
true for P. triticina Erikss. (P. ruhigo-vera triiici (Erikss. & Henn.) Carl.)
and several other species of rusts. Indeed it seems probable that almost
all rusts producing spermogonia have these two sexual phases. If two sori
from sporidia of opposite sexual phases lie close together aecia will develop
where their myeelia come into contact, without the help of the sperm cells.
Brown (1932) showed that a dicaryon mycelium (e.g., uredial sorus) can
also induce aecial formation in a sorus from a sporidial infection. Hanna
(1929a) found that 48 hours after a mixture of sperm cells of opposite
sexual phases was applied to a sorus of Puccinia graminis Pers., on the
leaf of barberry, the cells at the base of the aecial primordium became
binucleate. Using mixtures of sperm cells from different physiologic races

ORDER UREDINALES (tHE RUSTS)

389

oiPuccinia graminis on wheat, Waterhouse (1929), in Australia, obtained
two hitherto unknown forms which are probably to be interpreted as
crosses. Similar experiments by Miss Newton (1930) and others in Canada
and by Stakman (1930) and collaborators in Minnesota resulted not only
in interracial crosses but in crosses between P. graminis tritici and P.
graminis secalis and between the former and P. graminis agrostidis. In
the course of the various crosses several new physiological races have been
produced. The study of the results reveals that many of these races must
be heterozygous. This was further demonstrated by the production of
selfed strains. Thus physiologic race #17 when selfed (i.e., fertilized by
sperm cells of the same race but of opposite sexual phase) produced
aeciospores which gave rise to #17 and seven others besides, while race #53
gave besides itself seventeen other races. On the other hand race #9
proved to be homozygous.

Fig. 129. Subclass Teliosporeae, Order Uredinales.
Receptive hypha with attached sperm, just below
mouth of stoma. (Courtesy, Andrus: /. Wash. Acad.
Sci., 23(12) :544-557.)

Andrus (1931, 1933) demonstrated for the rusts Uromyces phaseoli
typica Arth. and U. phaseoli vignae (Barcl.) Arth, that there are certain
elongated filaments, which he called "trichogynes," whose tips emerge
from the stomata or from between epidermal cells and to which the sperm
cells adhere. The nucleus of the sperm enters the trichogyne and passes
down through it. Only then do the multinucleate cells appear in the aecial
primordium. Miss Allen (1932a) demonstrated the occurrence of similar
receptive hyphae in Puccinia triticina and other species of rusts and J. L.
Forsberg (in an unpublished thesis, 1932) has shown their presence in
Kunkelia nitens (Schw.) Arth. and Gymnoconia peckiana (Howe) Trotter,
but not in the monocaryon race of the former in which functional spermo-
gonia are not produced. Miss Rice (1933) showed that in a number of
other rusts similar structures are formed. According to Miss Allen (1932a)
the receptive cells, into which the one or more (up to ten or so) sperm
nuclei pass, elongate and branch and form a mycelium which penetrates

390 CLASS BASIDIOMYCETEAE

into the basal portion of the aecial primordium where dicaryon or pluri-
nucleate cells give rise to the chains of spores. In a later paper Miss Allen
(1933c) showed that in Puccinia sorghi Schw., whose aecial stage is pro-
duced in species of Oxalis, the paraphyses extending from the ostiole of
the spermogonium are receptive hyphae as was demonstrated for P.
helianthi Schw. by Craigie. The union of sperms to these paraphyses and
the actual passage of the sperm nuclei into them w^as illustrated. In her
work on P. iriticma and more particularly on P. graminis she reported
that receptive hyphae may emerge from stomatal openings or from be-
tween epidermal cells in addition to the spermogonial receptive hyphae.
Pierson (1933) and Buller (1938) demonstrated the presence of slender
flexuous nonseptate receptive hyphae growing up from the base of the
spermogonium and out into the drop of spermogonial nectar in Cronartium
ribicola Fischer and Puccinia graminis Pers., respectively. When a drop
of mixed nectar is added sperm cells were found attached to these hyphae
within a few hours. In the latter of the foregoing species there are in the
spermogonium paraphyses in addition to the flexuous hyphae but in no
case did Buller find that they served as receptive hyphae. (Fig. 129.)

Just how the sperm nuclei reach the primordium of the aecium is still

a matter of controversy. Miss Allen (1933a) suggested that they passed

into the receptive cells which elongated inward and formed a mycelium

which grew down to the basal portion of the aecial primordium. In Me-

lampsora lini (Pers.) Lev., where no receptive hyphae are present either

piercing the epidermis or passing through the stomata or emerging from

the spermogonia she reported (1934a) that the sperm cell dissolves a hole

through the outer epidermal wall of the host {Linum usitatissimum L.)

through which it gains access to the lumen of the cell where it germinates

or through which a slender germ tube passes. In either case a slender

branched mycelium arises which grows toward the aecial primordium

where abundant pairing of hyphae occurs, resulting in the formation of

the basal cells from which the chains of aeciospores are produced. The

suggestion has been made that the sperm nuclei possibly initiate a general

diploidization of the mycelium of the sorus so that all of the mycelium

between the point of union of sperm and receptive hypha to the aecial

primordium is diploidized, as Lehfeldt (1923) and Buller (1930) showed

to be the case in some of the Eubasidiae. Savile (1939) showed that in

several rusts investigated by him the cells of this intervening mycelium

are not diploidized, suggesting that the sperm nuclei travel from cell to

cell, without accomplishing diploidization, until the aecial primordium is

reached. He also demonstrated that sori close together in the leaf tissue

can fertihze one another without intervention of the sperm cells, probably

by contact of the mycehal cells and mutual diploidization. Andrus (1933)

obtained differential staining of the sperm nuclei and of those of the

ORDER UREDINALES (tHE RUSTS) 391

vegetative hyphae in the sorus and was able to confirm Savile's suggestion
that the former pass through the hyphae without bringing about their
diploidization. Apparently there is lacking a standard method of diploid-
izing the fertile cells of the aecial primordium. Miss Allen (1932b) has
demonstrated that in Puccinia coronata Corda the sperm cells unite
abundantly with the receptive hyphae and that mycelium (carrying the
sperm nuclei) grows from these down to the primordium. In the latter the
union of basal cells, to be described in the next paragraph, rarely occurs.
In the case of Puccinia sorghi Schw., whose spermogonia and aecia occur
on species of Oxalis, Miss Allen (1934b) reported that functional receptive
hyphae occur in the spermogonium, and also project out of stomata.
Sometimes sperm cells germinate on the outside of the leaf and the slender
hyphae enter the stomata. Within 24 hours after the " spermatization " of
a sorus with compatible sperms 60 per cent of the mycelial cells of the
mycelium are found to possess more than one nucleus, through rapid
division and migration of the sperm nuclei. Six days after spermatization
the first aecia set free their aeciospores.

Prior to the discovery of the active participation of the sperm cell
Blackman (1904), Christman (1905, 1907), Mme. Moreau (1913), Colley
(1918), and many others described the manner by which the aeciosporic
chains are originated. A layer of uninucleate cells in the basal portion of
the aecial primordium either shows fusion by twos, the walls between the
upper portions of these cells dissolving out, resulting in a binucleate two-
legged cell, or nuclei pass from adjacent cells through small pores to form
a dicaryon cell. From this by conjugate division of the nuclei and the
formation of successive cells comes a chain of aeciospores and disjunctor
or intercalary cells. Just how the two sets of uniting cells originated was
not made clear by these authors. It is now apparent that the two uniting
cells contain respectively a nucleus originating from the sperm and one
from the mycelium developed from the sporidium. Wang and Martens
(1939) do not believe that "Christman" conjugations at the base of the
aecium are the normal mode of diploidization but that it occurs earlier,
perhaps as far back as spermatial fusions with one another or with recep-
tive cells of the rust. (Fig. 130.)

From the foregoing accounts it is 'clear that in some and probably all
rusts which produce sperm cells they are functional. Furthermore, the
same haploid mycelium produces both sperm cells and receptive hyphae
so that both male and female structures are present, yet self-fertilization
does not occur. Like the condition in Schizothecium (Pleurage) reported by
Ames (1932), in A'eiirospora as demonstrated by Shear and Dodge
(1927), and in Stromatinia (Sclerotinia) gladioli (Drayton) Whetzel studied
by Drayton (1934), two sexual phases are present, each hermaphroditic
but incapable of self-fertilization.

392

CLASS BASIDIOMYCETEAE

Fig. 130. Subclass Teliosporeae, Order Uredinales. Origin of chains of aeciospores
from union of basal cells in the aecium; Cronartium rihicola Fischer. (A) Basal cell of
future aeciospore chain. (B) Chain of aeciospores and intercalary cells. (C-G) Steps
in the formation of a chain of aeciospores and intercalary cells in Phragmidium specio-
sum (Fr.) Cke. (A-B, courtesy, Colley: /. Agr. Research, 15(12) :619-660. C-G,
courtesy, Christman: Botan. Gaz., 39(4):267-275, Univ. Chicago Press.)

ORDER UREDINALES (tHE RUSTS) 393

The binucleate aeciospores are capable of remaining viable for a long
time and can be carried great distances by the wind and still cause infec-
tion. Falling upon a suitable host plant the spore germinates in a drop of
water (dew, rain, etc.) forming a stout germ tube which usually seeks out
and enters a stoma. Pady (1935) showed that in Gymnoconia peckiana the
germ tube penetrates the epidermis directly as do the germ tubes from
the sporidia, but this seems to be an exception. Just within the stomatal
opening the hypha enlarges to form a "substomatal vesicle" containing
many nuclei from which arise hyphae of dicaryon cells which penetrate
the host tissue in various directions, growing intercellularly and sending
haustoria into some of the cells between which they pass. Since the normal
aeciospore is binucleate this mycelium arising from it is dicaryotic and
has two nuclei in each cell, one descended from that of the sperm and
one from that of the parental monocaryon mycelium. Usually the size of
the infected area from one aeciosporic infection is limited, i.e., the fungus
in this phase is not capable of indefinite growth. When its full extent of
development is nearly attained a subepidermal mass of hyphae is formed
making a pseudoparenchymatous layer of dicaryon cells from which grow
short upright two-celled branches. These raise and eventually rupture the
epidermis, thus producing a uredial sorus or uredium. The outermost of
the two cells enlarges to form a binucleate urediospore while the other
elongates to form its long stalk. The urediospores are colorless or more
usually yellow to orange red in color, mostly finely verrucose or echinu-
late. There are several germ pores for each urediospore and their number
and position on the spore are of great assistance in the identification of
the various species. They break loose from the stalk and like the aecio-
spores may be carried long distances by the wind. They germinate and
infect the host through the stomata in exactly the same manner as do the
aeciospores. As long as the host plant has not passed a certain stage of
development these new infections produce other crops of urediospores.
As the host plant becomes more mature tehospores begin to appear, often
at first intermingled with the urediospores in the same sorus, but eventu-
ally in sori containing only teliospores. They arise from the same type of
pseudoparenchymatous subepidermal mycelium as do the urediospores.
From this basal layer there grow upward series of dicaryon cells. Some of
these may differentiate into stalk cells and teliospores or all may become
teliospores. Malengon (1936) observed the production rarely of teliospores
in the aecial sori of Puccinia atropae Mont. In some of the most primitive
rusts no basal layer is formed and the tehospores are produced singly or
by twos or threes as enlarged cells of the mycelium in the interior of the
leaf as occurs in Uredinopsis. The young teliospores, whatever their shape
or location, are at first binucleate. The two nuclei unite to form a diploid
nucleus. Usually the teliospore becomes thick-walled (mostly with one or

394 CLASS BASIDIOMYCETEAE

more germ pores) and more or less colored, from light yellow-brown to
almost black. In some of the rusts on ferns they are colorless and thin-
walled.

From the mature uninucleate teliospore the promycelium emerges at
once (various species of Cronartium, Puccinia malvacearum, etc.) or only
after over-wintering {P. graminis Pers.) or after certain unfavorable en-
vironmental conditions have passed. It emerges through one of the germ
pores and into it may pass the undivided nucleus of the teliospore or the
first of the two divisions of the nucleus may occur before the nuclei enter
the promycelium. The final division usually takes place in the promyce-
lium whereupon septa are formed dividing it into four cells. Sometimes a
further septum cuts off the teliospore from the basal cell of the pro-
mycelium. From the promycelial cells sterigmata grow bearing at their
tips the sporidia which are shot off violently at maturity. In the case of a
compound teliospore, such as is characteristic of Puccinia, Phragmidium,
Ravenelia, etc., each of the component cells produces its own mycelium,
thus demonstrating that each cell is to be considered a teliospore, not the
whole compound structure.

Rusts may be either autoecious or heteroecious. In the former the
sporidial and aeciosporic infections take place on the same or closely
related species of host while in a heteroecious species the host infected
by the sporidia belongs to a family not at all closely related to that con-
taining the host for the aeciosporic infection. Heteroecism was first proved
by actual inoculation experiments by de Bary in 1865 for Puccinia
graminis and by Oersted for Gymnosporangium sabinae (Dicks.) Wint. in
the same year. The former showed that the aeciospores produced on the
common barberry {Berheris vulgaris L.) would not infect that species but
would infect the small grains such as wheat, barley, etc. The urediospores
would infect the same and often also closely related species of grains. The
sporidia from the teliospores on the overwintered straw or stubble would,
on the contrary, infect only the barberry. Thus was brought the scientific
explanation of a phenomenon known for a hundred years or more that the
presence of barberry plants was detrimental to small grains. This much
had been definitely proved by careful observation and experiment long
before, but the actual connection of the rust on barberry with that on the
grain was left for de Bary to prove. Plowright (1889) gives an excellent
account of these early observations and beliefs as to the harmful effect of
barberry on grain. Since that time the heteroecism of hundreds of species
has been demonstrated. It is worthy of note that Waterhouse (1929)
discovered one physiologic race of wheat rust that is incapable of infecting
barberry.

In an autoecious species the aeciospores infect the same or closely
related host species as do the sporidia. Thus Pi/cama helianthi Schw., the

ORDER UREDINALES (tHE RUSTS) 395

rust of the sunflower {Helianthus annuus L. and other species) has its
sporidial infection on the sunflower with the production of a monocaryon
mycehum which produces the spermogonia and the aecial primordia. In-
fection by aeciospores produces in the same host species or even in the
same plant the dicaryon mycelium from which arise the urediospores and
teliospores. It may happen that both types of infection may occur on the
same leaf.

Of the rusts of more or less economic importance the hosts are indi-
cated below for the different stages of a few species, showing that
both heteroecious and autoecious rusts may be enemies of cultivated
plants. The list contains only a very few of the many rusts that attack
important crop plants. The customary symbols are used, viz., 0, spermo-
gonial development, I, aecial stage, II, uredial stage and III, telial stage.

Heteroecious Species

Puccinia graminis Pers., Black stem rust of small grains: 0 and I on Berberis vul-
garis L., barberry; II and III on small grains (wheat, rye, barley, oats) and
various other grasses.

P. rubigo-vera tritici (Erikss. & Henn.) Carl., leaf rust of wheat: 0 and I on species
of Thalictrum; II and III on wheat (Triticuni).

P. coronata Corda, crown rust of oats: 0 and I on various species of Rhamnus; II
and III on oats (Avena sativa L.).

P. sorghi Schw.: 0 and I on Oxalis strida L.; II and III on maize (Zea mays L.).

Gymnosporangium juniperi-virginianae Schw. : 0 and I on apple {Malus sylvestris
Mill.); Ill on red cedar (Juniperus virginiana L.).

Uromyces dianthi Niessl. {U. caryophyllinus Wint.): 0 and I on Tithymalus sp.;
II and III on carnation (Dianthus caryophyllus L.).

Cronartium ribicola Fischer, white pine blister rust ; 0 and I on white pine (Pinus
strobus L.) ; II and III on various species of currant (Ribes) and gooseberry
(Grossularia) .

Uredinopsis spp.: 0 and I on species of fir (Abies); II and III on various ferns.

Autoecious Species

Gymnoconia peckiana (Howe) Trotter: 0, I, and III on blackberry, dewberry,

and black raspberry (Rubus spp.).
Uromyces phaseoli typica Arth. : 0, I, II, and III on American beans (Phaseolus

vulgaris L.).
Puccinia asparagi DC: 0, I, II, and III on asparagus (Asparagus officinalis L.).
P. helianthi Schw. : 0, I, II, and III on sunflower (Helianthus annuus L. and some

other species).
Phragmidium spp.: 0, I, II, and III on various species of rose (Rosa sp.).

In the case of heteroecious rusts efficient control can be obtained by
the elimination of the alternate host in case the rust is unable to over-
winter on the host that is of economic importance. Thus in the northern
portions of the United States and of Europe the extermination of the
barberry has greatly reduced the ravages of the black stem rust because
this rust cannot survive the winter except as teliospores on the over-

396 CLASS BASIDIOMYCETEAE

wintering wheat or rye or other grains. In the southern portions of the
United States and Europe the rust is not killed out by the cold of winter
and so perpetuates itself by its urediospores. Under such conditions the
eradication of the barberry has little effect. The apple rust has been found
to be susceptible to control by removal of its alternate host, the red cedar,
over an area to a distance of a mile or more from the orchard that is to be
protected. White pine blister rust requires the destruction of all currants
and gooseberries to a distance of half a mile (one mile for Ribes nigrum L.)
from the trees it is desired to protect. Puccinia ruhigo-vera tritici, although
heteroecious, cannot be controlled in this manner for the rust that infects
the wheat in the fall survives the winter on this host and produces uredio-
spores in the spring from which the disease is spread. In fact this rust is
exceedingly abundant in many parts of the United States where the host
for the aecial stage does not occur.

The rust life cycle described above is the typical one. Rusts possessing
such a cycle are called macrocyclic or long-cycle rusts. Arthur and his col-
laborators (1929), Jackson (1931), and others believe that these represent
forms with the more primitive life cycle so far as present rusts are known.
Many species of rusts have shortened their life cycle by the omission of
one or more stages. Dietel (1928), Olive (1908), and Grove (1913), on the
contrary, consider the forms with the short life cycle to be more primitive.
As examples of the omission of certain stages the following may be men-
tioned. Most species oi Gymnosporangium produce no binucleate repeating
spores (urediospores) from the dicaryon stage of growth but they do occur
in G. nootkatense (Trel.) Arth. Some rusts produce only spermogonia and
telia while still others omit the spermogonia also (e.g., Puccinia malva-
cearum Bert.). These last two are properly speaking microcyclic rusts.
There are also rusts in which no true telia are produced but whose aecio-
spores germinate in the manner of teliospores by the formation of a
promycelium. These rusts also are microcyclic. Kunkelia nitens (Schw^)
Arth., on Ruhus spp. is of this type as are the various species of Endo-
phyllum. The microcyclic species are of especial interest as regards the
origin of the binucleate condition of the young teliospore. In Puccinia
arenariae (Schum.) Wint., liindfors (1924) described the formation of a
two-celled promycelium, each cell giving rise to a binucleate sporidium.
This produces a dicaryon mycelium and no monocaryon mycelium occurs.

Another anomaly in the life cycle of a short-cycle rust is described by
Thirumalachar (1946) for Uromyces aloes (Cke.) Magn. In this species the
spermogonia appear to be normal and exude drops of sweet liquid filled
with sperm cells. However, there are no flexuous hyphae. The appearance
of the spermogonia is followed immediately by the development of teha.
The monocaryon mycelium within the sorus shows here and there con-
tacts between two adjacent hyphae where the intervening walls are dis-

]

ORDER UREDINALES (tHE RUSTS) 397

solved and a nucleus passes from one cell into the other. From this cell
arise dicaryon hyphae which occupy that portion of the sorus while the
monocaryon hyphae degenerate. These dicaryon hyphae give rise to bi-
nucleate teliospores on slender binucleate stalks. The nuclei unite in the
teliospore and the pedicel breaks, so that the teliospores are distributed by
air currents. The zygote nucleus passes into the apical promycelium where
the two normal divisions of meiosis occur. Usually but one septum is
formed dividing the promycelium into a basal uninucleate cell which soon
degenerates, and a large terminal cell with three nuclei. This sends out a
slender germ tube into which the nuclei enter and infect the host. After
the epidermis is penetrated the nuclei divide further and septa are formed,
dividing the resultant mycelium into uninucleate cells. In this species the
terminal cell of the promycelium takes the place of a sporidium in infect-
ing the host while the distribution of the teliospore by air currents makes
distribution by means of sporidia unnecessary. Apparently the spermo-
gonia and sperm cells do not function.

In other microcyclic forms there are other ways in which the life cycle
is completed. Miss Allen (1933b) found that in Puccinia malvacearum the
teliospores arise from dicaryon mycelium. The two nuclei unite and then
divide in the promycelium in the usual way. The nucleus then undergoes
division in the sporidium but Miss Ashworth (1931) finds that it gives
rise to a monocaryon mycelium. According to her investigations certain
cells in the telial sorus show nuclear migrations producing the dicaryon
phase. Miss Allen (1935) reported that occasional conidia are formed at
the tips of hyphae emerging from the stomata and suggested that possibly
these may have a part in the diploidization of the mycelium that forms
the telial sorus. However, in the main this appears to result from the
intermingling and fusion of hyphae from mycelia produced from two
separate but adjacent sporidial infections. In Puccinia prostii Duby, ac-
cording to I. M. Lamb (1934), no receptive hyphae occur in the spermo-
gonia which are produced. The mycelium within the host (Tulipa sp.)
remains monocaryotic. The teliospores arise from the lateral fusion of two
adjacent hyphal tips or from the passage of a nucleus through a small
opening in the septum from a basally placed cell to the cell above. Al-
though normal teliospores are produced, apparently infection rarely if
ever occurs by means of sporidia, the mycelium being carried over from
year to year through the bulbs.

In most of the short-cycled rusts studied in which normal spermogonia
are produced (spermogonia and telia, or spermogonia and aecia whose
spores function as teliospores) it has been shown that the mycelium is of
monocaryon type until the telium or aecium is formed, when dicaryon
cells appear. It is probable that this dicaryon phase arises in the same way
as is described above for macrocvclic rusts.

398

CLASS BASIDIOMYCETEAE

The transfer of the production of the mycehum back from the teho-
spore to the aeciospore has apparently taken place independently several
times so that the microcyclic rusts of the formula 0, I are not necessarily
closely related but have probably developed in separate lines from 0, I,
II, III forms. It is clear that Kunkelia nitens, the microcyclic orange rust
of Ruhus, is derived from Gymnoconia peckiana, a long-cycle form (0, I,
III) on the same hosts. The genus Endophyllum represents a similar series
of cases. Some of the species of this genus correspond in aecial host and

Fig. 131. Subclass Teliosporeae, Order Uredinales,
Family Pucciniaceae. A-D, Endophyllum sempervivi (A.
& S.) de By. (A) Binucleate aeciospore. (B) Aeciospore with
the nuclei united. (C) Aeciospore germinating. (D) Pro-
mycelium formed. (E, F) Endophyllum euphorbiae-
sylvaticae (DC.) Wint. (E) Aeciospore germinating with-
out union of nuclei. (F) Promycelium and sporidia. (After
Moreau and Moreau: Bull. Sac. Botan. France, 66:14-44.)

structure to the aecial stage of known macrocyclic species. These abbrevi-
ated forms may even lack the fusions of the two nuclei within the spores
before the formation of the promycelium. Dodge and Gaiser (1926)
showed that in Kunkelia nitens the two nuclei pass out into the promy-
celium where each divides again to form the four nuclei which enter the
sporidia. The Moreaus (1919) have found the same to be true for Endo-
phyllum euphorbiae-silvaticae Lev., but in other species of the genus they
demonstrated nuclear fusion in the aeciospore before the promycelium
began to be formed. Dodge (1924) reported that in one form of Kunkelia
nitens no dicaryon mycelium was produced at all and the aeciospores re-

ORDER UREDINALES (tHE RUSTs) 399

main uninucleate and produce a two-celled promycelium. The spermo-
gonia in this race are never more than rudimentary and no receptive
hyphae are formed. (Fig. 131.)

A third type of life cycle is the one usually designated as 0, II, III,
i.e., a cycle in which the typical aecial structure is lacking. The spermo-
gonia are succeeded by sori containing spores exactly resembling typical
urediospores. In the same way as are produced the dicaryon basal cells of
the chains of aeciospores Christman (1907) showed that there are pro-
duced dicaryon cells giving rise to urediospore-like structures. The latter
give rise to dicaryon mycelium from which may arise later another series
of urediospores. The usual interpretation of this phenomenon is that this
is really a macrocyclic rust of the formula 0, I, II, III in which the aecio-
spores are not produced in chains but singly on stalks, like urediospores.
The primary (first produced) urediospores are therefore modified aecio-
spores while the secondary ones are true urediospores.

The aecium may be cup-shaped (cupulate) with a well-developed pe-
ridium or it may be very tall so as to make a horn-like (cornute) structure.
The peridium may be lacking so that the aecium is diffuse. Other forms
are known, the most curious of which is the "hyphoid" aecium of Dasy-
spora foveolata B. & C. (D. gregaria (Kze.) Henn.) in which a branching
dicaryon mycelium emerges through various stomatal openings, forming
a colorless mass of hyphae. These are terminated by single, not catenulate,
aeciospores which drop off while the hypha elongates sympodially and
produces another spore and so on. The cytology of this type of aecium
needs careful investigation to determine where the diploidization occurs.
Sydow (1925) interprets these as urediospores but Arthur and co-authors
(1929) consider them to be aeciospores. (Fig. 132.)

The uredium may be merely a cluster of stalked urediospores bursting
through the epidermis of the host or it may be surrounded by paraphyses.
In Cronartium and Puccmiastrum and closely related genera the uredium
possesses a true peridium. In some genera, e.g., Coleosporium the uredio-
spores, like the aeciospores, are produced in chains but they arise from a
dicaryon mycelium and so differ from the latter. In a few rusts the ured-
ium is cupulate and resembles the aecium but spermogonia are lacking.
They are sometimes called secondary aecia. Cummins (1937) reported
that the uredium of Prospodium is a salver-shaped structure whose slender
stalk about three cells thick emerges through a stoma and spreads out as a
flat plate with upright fringe-like marginal paraphyses. On the flat surface
of the salver arise the stalked urediospores. In some species of the genus
the telial sorus is quite similar. (Fig. 133.)

The telium is the most variable structure in the order. Properly speak-
ing a teliospore is a single cell, binucleate at first but becoming uninucleate
by the fusion of the two nuclei and giving rise immediately or after a delay

400

CLASS BASIDIOMYCETEAE

Fig. 132. Subclass Teliosporeae, Order Uredinales. Various types of aecia. (A)
Cupulate aecium of Uromyces erythronii (DC.) Pass. (B) Margin of caeomoid aecium
of Phragmidium rubi (Pers.) Wint., showing absence of peridium but presence of
paraphysate hyphae. (C) Hyphoid aecium of Dasyspora foveolata B & C. and a com-
pound teliospore. (A-B, after Sappin-Trouffy : Le Botaniste, 5:59-244. C, after
Sydow: Mycologia, 17(6):255-262.)

ORDER UREDINALES (tHE RUSTS)

401

Fig. 133. Subclass Teliosporeae, Order Uredinales. Types of uredia. (A) Uredium
with peridium in Melampsoridium beiulinum (Pers.) Kleb. ; on either side a portion of a
tehal sorus. (B) Uredium without peridium but with numerous capitate paraphyses
among the urediospores, in Melampsora hehoscopiae (Pers.) Cast. (C) Urediospores of
Coleosporium soUdaginis (Schw.) Thlxm., in a chain. (D) Extrastomatal uredium of
Prospodiurn plagiopus (Mont.) Arth. (A-B, after Sappin-Trouffy : Le Botaniste,
5:59-244. C, after Christman: Botan Gaz., 44(2):81-101. D, after Cummins: Ann.
ilfycoL, 35(1) :15-21.)

402 CLASS BASIDIOMYCETEAE-

to a promycelium. The teliospores are produced under the epidermis or in
the epidermal cells or in the mesophyll, or very rarely (Cystospora, Go-
plana) the basal cells from which they arise may push out through a stoma
so that the teliospores are then produced externally. The stalked types of
teliospores are formed subepidermally and become external by the rupture
of the epidermis. In some genera they are formed in separable chains and
the telium is surrounded by a peridium which bursts the epidermis and
opens to allow the teliospores to escape. In the fern rusts of the genus
Uredinopsis the teliospores are produced in the mesophyll of the leaf
singly or united, two, three, or four together, into a compound teliospore,
each of whose component cells gives rise to a separate promycelium which
emerges from the leaf surface. In some species of Pucciniastrum the telio-
spores are produced in groups of two to four cells in the epidermal cells of
the host, but in other species these clusters of teliospores may be aggre-
gated laterally into a subepidermal crust. In Cronartium the teliospores
are joined laterally and longitudinally into a tall waxy column which
pushes out through the epidermis to a length of up to 6 or 8 mm. In
Melampsora and Coleosporium the teliospores are crowded laterally into a
subepidermal or subcuticular crust. Among the stalked forms Uromyces
has but a single teliospore at the apex of its stalk, Puccinia has two united
teliospores on the single stalk, in Phragmidium one stalk bears a row of
three to eight or more teliospores. In Pucciniosira the telium has a perid-
ium and the teliospores are formed on a stalk in chains which break apart
into units of two teliospores each. In Ravenelia the stalk bears a head of
laterally united teliospores, below which hang colorless cells, the so-called
"cysts." Usually in systematic literature these various types of compound
teliospores are spoken of as single teliospores although properly each cell
from which a promycelium arises is a teliospore.

The order has been divided into many families or into two families.
The author follows Dietel (1928) and the later works of Arthur (1929,
1934), both of whom recognize but two families, each divided into several
tribes.

Family Melampsoraceae. Teliospores without stalks, produced
singly or united in groups of two to four in the mesophyll or just below or
within the epidermal cells, or united laterally into subepidermal or sub-
cuticular crusts, or united into separate vertical chains or into chains
which are united laterally into a waxy column which bursts through the
epidermis. Aecia mostly on species of Family Pinaceae. There are 15 to 20
genera and about 300 species. This family clearly includes the most primi-
tive living representatives of the order. The genus Uredinopsis with its
colorless thin-walled teliospores, single or united by twos, threes, or fours,
in the mesophyll of the leaves of ferns, its two kinds of colorless uredio-
spores, thick-walled and thin-walled, the sori surrounded by peridia, and

ORDER UREDINALES (tHE RUSTS)

403

Fig. 134. Subclass Teliosporeae, Order Uredinales, Family Melampsoraceae.
(A, B) Uredinopsis struthiopteridis Stormer. (A) Urediospore. (B) Section through
leaf of host showing uredial sorus with peridium, and teliospores scattered through the
mesophyll. (C) P ucciniastrum goeppertianum (Klihn) Klebahn; telial stage in epi-
dermis of Vaccinium sp. (A-B, after Dietel: Ber. deut. botan. Ges., 13(7):326-332.
C, after Hartig: "Lehrbuch der Baumkrankheiten," Berlin, J. Springer, 1889.)

with its aecia and spermogonia on the needles of species of Abies probably
combines the greatest number of primitive characters of any rust, viz.,
telial hosts in the ancient group Pteridophyta, colorless urediospores and
teliospores, the latter scattered in the mesophyll, uredia surrounded by
peridium etc. Two other genera {Hyalopsora and Milcsia) with colorless
teliospores formed in the epidermal cells are also found in the ferns. P uc-
ciniastrum goeppertianum (Klihn) Klebahn is of interest because it is one
of the few rusts in which the sporophytic mycelium is perennial. The aecia
occur on the leaves of the fir (Abies) as is true of the other genera men-
tioned above. The telial stage (no uredia are known) occurs in Vaccinium,
its presence causing the development of a sort of witches' broom with an
upright thickened stem and small distant leaves. The mycelium grows out

Fig. 135. (See legend on facing page.)
404

ORDER UREDINALES (tHE RUSTS) 405

into the epidermal cells of the stem, there forming clusters of two to four
closely united teliospores which send forth their promycelium almost im-
mediately, through the epidermal cell walls. In Cronartium the gameto-
phytic (monocaryon) mycelium is perennial in the twigs and cortex of
older limbs of pine (Pinus), often causing the formation of galls which may
attain great size and age, up to the size of a human head in C. quercuum
(Berk.) Miyabe. The uredia are small, with a peridium. The teha are
rows of teliospores united laterally into a waxy column 6 to 8 rows in
thickness and bursting through the epidermis and projecting several milli-
meters. These columns continue to grow at the base for some time. Each
of the hundreds of teliospores of the column germinates immediately by a
curved promycelium producing almost spherical sporidia. The dicaryon
stage is found in various Flowering Plants (Anthophyta), apparently only
in Dicotyledoneae. In Chrysomyxa the tehospores are in separate chains
of three or more spores each which arise at the base of the sorus. The
urediospores are also in chains. The sporophytic phase occurs in the
Family Ericaceae in the wider sense and in a few other families. Coleo-
sporium and Melampsora both produce subcuticular or subepidermal
crusts one cell thick of laterally united teliospores. In the former the
urediospores are produced in short chains and the promycelium is "in-
ternal." In the latter the urediospores are single and the promycelium is of
typical structure. Coleosporium solidaginis (Schw.) Thiim., on Aster spp.
and Solidago spp. is abundant even in those parts of the United States
where the aecial hosts (species of Pinus) are not found. It is apparently
able to maintain itself by overwintering mycelium or urediospores. Me-
lampsora medusae Thiim. causes the spotting of leaves of various species
of poplars iPopulus) with the dark-colored telial sori, these being pre-
ceded by the small powdery yellow uredial sori. The aecial host is the
larch {Larix sp.) on whose young needles the almost white aecia appear.
Most of the species of this genus have their aecia on Pinaceae, but M.
lini (Pers.) Lev., on flax (Linum usitatissimum L.) is autoecious as is M.
euphorUae (Schub.) Cast. In M. rihesii-purpureae Kleb., the tehal stage
is on Salix but the aecial stage occurs on species of Ribes and Grossularia,
while three other species with Salix as their telial host have as their aecial
hosts respectively Saxifragaceae, and Larix and Ahies in the Pinaceae.
One microcychc species M. farlowii (Arth.) J. J. Davis, occurs in the
United States on Tsnga canadensis (L.) Carr. of the Pinaceae. (Figs. 134,

135.)

Family Pucctniaceae. Teliospores usually stalked, simple or com-
pound, sometimes without stalks and produced successively as simple or

Fig. 135. Subclass Teliosporeae, Order Uredinales, Family Melampsoraceae, (A)
Cronartium flaccidum (A. & S.) Wint. ; teliospore column growing up through uredium.
(B) Coleosporium sonchi-arvensis (Pers.) L6v.; vertical section through telium. (After
Sappin-Trouffy: Le Botaniste, 5:59-244.)

Fig. 136. Subclass Teliosporeae, Order Uredinales, Family Pucciniaceae. Various
types of teliospores, all compound except Uromyces and Goplann. (A) Uromyces fabae
(Pers.) de By. (B) Puccinia graminis Pers. (C) Phragmidium ruhi-idaei (DC.) Karst.

(Continued on facing page.)
406

ORDER UREDINALES (tHE RUSTS) 407

compound teliospores which escape from the sorus dry or embedded in
sHme. Aecia only very exceptionally on Pinaceae. The stalked forms of
this family are easily distinguished from the Melampsoraceae but the
forms with tehospores produced in loose, quickly fragmenting chains are
a sort of connecting group difficult to segregate definitely from one or the
other family. Dietel (1928) recognized 83 genera and about 3000 species,
while some authors add at least 20 genera by the segregation of such
larger genera as Uromyces and Puccinia. These two are among the more
important genera of the family with, respectively, about 600 and over
1800 species. The location of the spermogonia is of importance taxonomi-
cally. They are subcuticular and rather flattened or subepidermal and
then more spherical. The uredia may have paraphyses or these may be
lacking. The aecia may be cupulate, or cornute, or hyphoid, or caeomoid,
or uredioid. The teliospores may be single on the pedicel {Pileolaria,
Trachyspora, Mainsia, Uromyces) or several on one pedicel (compound
teliospores). In Puccinia there are two; in Phragmidium, Xenodochus, etc.,
three to many in a single row on the simple pedicel. In Ravenelia the telio-
spores are numerous in a head on a usually compound pedicel, being sub-
tended by hyaline "cysts." As in the Melampsoraceae many species are
autoecious and long-cycled, others are heteroecious, and many lack one or
more spore forms. In the case of Puccinia and Uromyces the relationship
is very close. The spermogonia are subepidermal, the aecia are cupulate
and the uredia are without peridium with urediospores single on long
stalks, in both genera. The teliospores are brown and stalked, emerging
from a ruptured epidermis. In Uromyces they are simple, in Puccinia
compound, formed of two teliospores closely united in a row with a one-
celled stalk. A number of cases are known where the aecia of heteroecious
species of the two genera are borne on the same host and the urediospores
and teliospores on the same alternate host, the aeciospores and uredio-
spores of the respective species of Uromyces and Puccinia being practically
indistinguishable. The only essential difference is that in the one genus
the pedicel is topped by a single teliospore and in the other by two.
Arthur (1934) believes that the frequency of such cases indicates a very
close relationship of the two genera. In some species of Puccinia, inter-
mingled in the same sorus may be found a varying number of Uromyccs-

FiG. 136 — {Continued)

(D) Dicheirinia binata (B. & C.) Arth. (E) Prospodium plagiopus (Mont.) Arth. (F)
Ravenelia acaciae-micranthae Diet. (G) Section through portion of tehum of Goplana
dioscoreae (B. & Br.) Cummins. (A, after Engler and Prantl: Die Xatiirlichen Pfianzen-
familien, Leipzig, W. Engelmann. B, after Sappin-TroufTy: Le Botaniste, 5:59-244.
C, courtesy, Cummins: Mycologia, 23(6):433-445. D, courtesy, Cummins: Mycologia,
27(2):151-159. E, courtesy, Cummins: Lloydia, 3(l):l-78. F, after Dietel: Botan.
Centr. Beihefte, Zweite Abt.^, 20 :SA3-4\3. G, courtesy, Cummins: Mycologia, 27(6) :605-
614.)

408 CLASS BASIDIOMYCETEAE

like teliospores ("mesospores") among the typical compound teliospores.
In Phragmidium the row of teliospores is longer, three to eight or more in
a row, and the stalk is long and enlarged toward the base. In Gymnospo-
rangium (with 40 or more species) the stalks are very long and their walls
as well as the outer walls of the usually two-celled compound teliospores
swell when wet so that the masses of teliospores are extruded from the
telial galls as gelatinous tongues sometimes 2 to 3 cm. in length. The aecial
hosts of this genus with a few exceptions belong to the Malaceae and the
telial hosts are species of Juniperus or closely allied genera. This genus
forms rather an exception in that it is the dicaryon stage that is perennial,
the galls on the host persisting sometimes for several years. In Dicheirinia
two, rarely three, verrucous teliospores are borne side by side at the apex
of a common pedicel whose upper cell is divided into two or three short
cells from which the teliospores arise. Prospodium, like Puccinia, has two
teliospores in a single row but they differ in minor points. Goplana is some-
times placed in this family because of its stalked spores. These grow up
into a gelatinous matrix and, in the manner of Coleosporiinn produce an
internal promycelium. Ravenelia is a genus whose tehospores form a head
of one layer of fertile spores subtended by colorless cysts which possibly
represent sterile tehospores. The head is supported by a centrally attached
stalk usually several cells in thickness. The species of this genus are
mostly tropical and subtropical and are autoecious so far as known.
Gymnoconia resembles Puccinia in its two-celled compound teliospores
but the aecia are diffuse (i.e., caeomoid) without peridium, and the
spermogonia are subcuticular. Urediospores are lacking. (Fig. 136.)

In addition to the genera and species assigned to these two families
there are over 1000 species of which the telial stage is unknown or its
connection with the other stages not determined. Peridermium, Aecidium
(600 species) and Caeoma represent different types of aecia. Uredo (450
species) consists of species of which the uredial stage only is known. In
most of these cases there are probably other stages as yet unknown or
whose connection with these has not yet been demonstrated, but it is
possible that in some of these species the other stages have been omitted
during the course of evolution.

The Uredinales are of great economic interest on account of their
harmful effects on many important crops. The following may be men-
tioned since they frequently cause great damage on small grains: Puccinia
graminis Pers., several varieties on wheat, rye, oats, barley; P. ruhigo-vera
(DC.) Wint., several varieties on wheat, barley, rye; P. coronata Corda, on
oats; P. glumarum (Schm.) Erikss. & Henn., on wheat, rye, barley; P.
sorghi Schw., on corn (maize); Gymnosporangium juhiperi-virginianae
Schw., on apple; Tranzschelia pruni-spinosae (Pers.) Diet., on peach;
Gymnoconia peckiana (Howe) Trotter and Kunkelia nitens (Schw.) Arth.,
on Ruhus spp.; P. asparagi DC, on asparagus; Uromyces phaseoli typica

ORDER USTILAGINALES (tHE SMUTS)

409

Arth., on Phaseolus vulgaris L.; Melampsora lini (Pers.) Lev., on flax;
Cronartium ribicola Fischer, on white pine {Pinus strohus L.). Several
species of rusts are harmful to ornamental plants.

Order Ustilaginales (The Smuts). The Smuts are parasitic, but ca-
pable of growth as saprophytes on substrata rich in organic material, e.g.,
well-manured fields. Flerov (1923) and Sartoris (1924) grew several species
of Ustilago through to teliospore production on artificial culture media
apart from their hosts. Recently Leach and Ryan (1946) grew Ustilago
striiformis (West.) Niessl. from teliospore to teliospore. Kniep (1911) ac-
complished this with Urocystis anemones (Pers.) Schroet. and Wernham
(1938) with U. gladioli (Req.) Sm. In the host plant the mycelium is at
first intracellular (Kolk, 1930) and later intercellular, with or without
haustoria, and actually growing in and keeping pace in its growth with
that of the meristematic regions of the host, usually dying in the portions
that have passed that stage. A few species parasitic on perennial hosts
live over winter in the crown of the plant so that the new growth becomes
diseased. Seyfert (1927), investigating several smuts in Europe and Stak-
man and Christensen (1927) studymg Ustilago zeae (Beckm.) Unger, and
Hanna (1929b), investigating the same species seem to have demon-
strated clearly that clamp connections are often present in the dicaryon
stage of growth. Sleumer (1932) however, claims that these are not true
clamp connections but abortive branches. The nuclear behavior as re-
ported by Seyfert would support the idea that they are typical clamp

Fig. 137. Subclass Teliosporeae, Order Ustilaginales. (Left and center) Family
Ustilaginaceae, Ustilago levis (Kellerm. & Swingle) Magn. (Left) Mycelium in meri-
stem of growing tip of seedling of Avena sativa L. (Center) Piece of intercellular
mycelium. {Right) Family Tilletiaceae, Urocystis anetnones (Pers.) Wint.; haustorium.
(Courtesy, Lutman: Trans. Wisconsin Acad. Sci., 16(2):1191-1244.)

410 CLASS BASIDIOMYCETEAE

connections, especially since Stempell (1934-1935) grew various species
of Entyloma in culture and obtained typical clamp connections there. The
teliospores are produced largely in the flowers, fruits, and inflorescences,
but in many cases are produced in the leaves or stems. More often the
masses of teliospores are dusty at maturity, and with the rupture of the
host tissues are set free for distribution by the wind or other means.
The sporidia are shot from the promycelium with considerable force in the
Family Tifletiaceae accompanied by the formation of a droplet of water
at the apex of the sterigma a moment before the discharge of the spore.
Conidia are mostly produced on the saprophytic mycelium but in Tubur-
cinia and Entyloma they are produced abundantly on the surface of the
Hving leaves of the host plant on long conidiophores which emerge from
the epidermis and give a whitish appearance to the affected leaf. When a
conidium arises from a cell of monocaryon mycelium the nucleus of the
cell divides and one of the daughter nuclei passes out into the conidium
which grows out from one side of the hyphal cell. If the cells are binucleate
both nuclei divide simultaneously and one daughter nucleus of each pair
passes out into the developing conidium, or but one nucleus enters the
conidium (Paravicini, 1917). Thus it is possible for a conidium from a
monocaryon mycelium to produce a new mycelium only of the same sexual
phase, but that from a dicaryon mycelium may produce a dicaryon
mycelium or monocaryon mycelia of one or the other sexual phase depend-
ing upon which nucleus entered the conidium. (Fig. 137.)

Liro (1935-1938) described conidia in the anthers of plants of Carda-
mine hellidifolia L., in whose pods the seeds were destroyed by Ustilago
cardamines Liro. The conidia cover and fill the discolored anthers. He
proposed for this conidial stage the name Rhombiella cardamines Liro. He
also described, under the name Crotalia cifitractiae-fischeri, the conidial
stage of Cintradia fischeri (Karst.) Liro, parasitic on Car ex canescens L.

Infection of the host plant takes place only in meristematic tissues, by
means of sporidia or conidia or by a germ tube produced in place of a
sporidium. The Smuts fall roughly into four groups Avith reference to the
manner of infection:

1. Infection takes place as the seed germinates, either from sporidia or
germ tubes produced on promycelia from teliospores adhering to the seed
or from sporidia or conidia present in the soil. This type of infection can
be controlled by treating the seed with suitable disinfectants before
planting them in soil free from smut. Examples are Ustilago avenae (Pers.)
Jens., oat smut; Tilletia foetida (Wall.) Liro {T. levis Kuhn) and T. caries
(DC.) Tul. {T. tritici (Bjerk.) Wint.), both of which cause stinking smut
or bunt of wheat; Urocystis occulta (Wall.) Ilab., causing stem smut of rye.

2. Any actively growing meristem may be infected by sporidia or by
conidia. In the case of maize smut, Ustilago zeae, the infection may occur
on young roots, at any joint of the stem (i.e., the meristem at the base of

ORDER USTILAGINALES (tHE SMUTs) 411

each internode), on young, not yet unrolled leaves, on the male inflores-
cence (tassel) before it emerges, on the young ear or certain grains of the
ear and even on the elongated styles ("silks") on which N. F. Petersen
has shown that it may cause small galls. ^ Treatment of the grain with
disinfectants is of little value in controlling this type of infection. Planting
must be done in soil free from the fungus.

3. Infection of the flower takes place at times of blooming. This was
worked out by Brefeld and Falck (1905). It occurs in the loose smuts of
wheat and barley, Ustilago tritici (Pers.) Rostr. and U. nuda (Jens.)
Kellerm. & Swingle, respectively. When the host plant has headed out the
flowers are normally self-pollinated before opening. When they open the
teliospores from nearby diseased plants germinate and produce their
sporidia or germ tubes on the stigmas and infect them, the germ tubes
growing down into the ovary and entering the developing embryo, in the
growing point of whose stem the mycelium becomes dormant until the
grain is planted. Then it grows rapidly in the apical meristem, apparently
causing Httle injury to the host plant until the head is being produced.
Within this developing head the mycelium grows very vigorously and
reduces it to a skeleton surrounded by the powdery masses of spores which
are set free at just the time the healthy plants are coming into flower.
Control is possible by soaking the infected grain and then dipping it into
hot water at a temperature and for a length of time that will kill the con-
tained mycelium without killing the grain.

4. Another type of infection has been shown by Mundkur (1943) to
occur in the case of Neovossia indica (Mitra) Mundkur. In this smut the
spores or the smutted grains fall to the ground and under favorable condi-
tions the long promycelia produce up to 150 sporidia which are wind
borne and infect the kernels in the dough stage or a little earlier. These
kernels then become smutted that season.

Smuts may cause large gafls consisting in part of host tissues and in
part of fungus tissues. The galls of maize smut ( Ustilago zeae) may attain
a large size and are edible when young. Various leaf smuts cause the pro-
duction of galls, e.g., Doassansia on the leaves of Sagittaria. Many smuts
on the other hand do not show their presence until their teliospores are
formed. Ustilago violacea (Pers.) Fuckel, attacks several species of Dian-
thaceae and Alsinaceae. Its teliospores are formed only in the anthers.
When the female plant of a dioecious species of Lychnis is infected the
presence of the fungus causes the flower to produce stamens, within which
the fungus produces its spores although the normal female flower lacks
stamens.

The order is divided into two famihes which are almost certainly
closely related and to which a third family, Graphiolaceae, is probably
related.

^ In a letter to the author.

412 CLASS BASIDIOMYCETEAE

\ ^:^

ri;.Ti-sa>^-i'i<.^^'ii.J"-'..---»^'/'.-.'>-/^-.*p.-".vA.i'-^-«''^ "»"'**'"'"'-

,f „ ,i..S*>-=

Fig. 138. Subclass Teliosporeae, Order Ustilaginales, Family Ustilaginaceae,
Ustilago zeae (Beckm.) Unger. (A) Germination of teliospore to form promycelium
and sporidia. (B) Two sporidia of opposite sexual phase, infecting epidermal cell of
host {Zea mays L.), the slender germ tubes uniting to form a stout multinucleate
hypha. (Courtesy, Hanna: Phytopathology, 19(5):415-442.)

Family Ustilaginaceae. Promycelium transversely septate into
several, mostly four, cells. The teliospores arise from transformed di-
caryon cells of the mycelium. Sometimes the cell walls of the usually
tangled hyphae swell and undergo gelatinization, thus separating the
protoplasts to some distance. Around these then, within the gelatinized
walls, new walls are laid down and the cells enlarge while the gelatinized
walls disappear. At about the same time the two nuclei unite and the
teliospores come to maturity. The teliospores are produced singly or
united into spore balls. They are usually dark-colored, with the wall more
or less thickened and smooth or rough. From each cell of the promycelium
usually several sporidia are produced, for, unlike the majority of Uredi-
nales, the nucleus of the cell divides several times and one of the daughter
nuclei enters each sporidium. In some cases each promycelial cell grows
out into a slender germ tube instead of producing sporidia. This difference
in behavior is connected for some species with differences in temperature,
moisture, etc. In Ustilago nuda (Jens.) Kellerm. & Swingle and U. tritici
(Pers.) Rostr. sporidia are apparently never found. (Fig. 138A.)

Sexual reproduction in this family is accomplished in various ways. In
several species it has been determined that two of the promycelial cells
represent one sexual phase and the remaining two the other phase. The
distribution of the two sexual phases in the four cells of the promycelium
may occur apparently in every possible order. In Ustilago zeae (Beckm.)
Ung., Hanna (1929b) showed that there occur four sexual phases. Prob-
ably there are two allelomorphic pairs of genes on separate chromosome
pairs that control this sexual behavior. If both chromosome pairs undergo
their reduction (disjunction) division in the first or in the second of the
two divisions occurring in the production of the promycelium two of the
promycelial cells will be of one sexual phase and two of the opposite

ORDER USTILAGINALES (tHE SMUTs)

413

phase. This is the common case. But if one chromosome pair undergoes
disjunction in the first nuclear division and the other chromosome pair
waits until the second division before undergoing disjunction the result
will be four nuclei with four different combinations of the sexual factors,
i.e., four sexual phases in the same promycelium. This doubtless occurs in
the other smut species also. The sexuality of smuts has been studied also
by Paravicini (1917), Bauch (1922 and later publications), Dickinson
(1927, 1928), Kniep (1926), Sleumer (1932), and others. Liro (1924) de-
scribed a peculiar method of sexual reproduction in Ustilago vuijckii Oud.
& Beijer., on Luzula multiflora (Hoffm.) Lej. This should be reinvestigated

Fig. 139. Subclass Teliosporeae, Order Ustilaginales,
Family Ustilaginaceae. Germination of teliospores by two
or more promycelia. (A) Sphacelotheca columellifera (Tul.)
Yen. (B) Sphacelotheca schweinfurthiana (Thiim.) Sacc.
(C) Sorosporium consanguineum E. & E. (After Yen:
Rev. mycol, 2(2):76-84.)

and similar studies undertaken on other species for it is very different
from the method usually accepted for this family. The sporogenous hy-
phae in the ovaries of the host are slender and more or less dichotomously
branched. The cells are binucleate. From the base toward the apex of
these hyphae the cells swell successively, usually leaving a slender portion
containing the septum between each cell and the one next above it. At the
upper part of a cell a branch is produced, containing one nucleus. This
branch is separated from the cell below by a septum. It pierces the cell
above by a very fine tube and the cytoplasm and nucleus pass into it,
making it 3-nucleate. In the meantime a similar process from this cell
takes one of the two original nuclei and transfers it to the next overlying
cell, and so on. Then the two nuclei unite and a thick wall is formed,
within and free from the original wall. Sometimes an "antherid," as Liro
calls these small branches, from another hypha fertilizes a cell and in that

414 CLASS BASIDIOMYCETEAE

case the antherid from the cell below does not fertilize it but degenerates,
unless it finds an adjacent hypha one of whose cells it can fertilize. Liro
calls the receptive cells "oogones," and suggests the similarity to the
sexual process in the Peronosporaceae. Oudemans (1895) saw these bridg-
ing branches and called them clamp connections, but that can hardly be
correct since they grow upward from the base toward the apex, and con-
nect two dicaryon cells.

Occasionally more than one promycelium may grow from a single
teliospore. This has been observed by Brefeld (1895) and by Yen (1937),
both in Sphacelotheca schweinfurthiana (Thiim.) Sacc. Yen observed a
similar development of two promycelia from a teliospore of Sorosporiuni
consanguineum E. & E. On dilute beer malt the sporidia are produced in
clusters terminally (as in Tilletia) and in lateral groups, not one at a time
from each cell of the promycelium. The sporidia in liquid media often
bud in the manner of yeasts, forming a monilioid chain or a cluster of
yeast-like cells. (Fig. 139.)

Hiittig (1933) has shown that in Ustilago avenae (Pers.) Jens, the
temperature has considerable effect upon the proportions of disjunction
in the first and second meiotic divisions. Disjunction in the first division
(called by him pre-reduction) occurred in 14 per cent of the cases at 9° C,
17.4 per cent at 19°, 31.5 per cent at 25.5°, and 18.7 per cent at 29.5°.
Various chemicals also modify these proportions.

Union of opposite sexual strains may take place by the conjugation of
two sporidia, the nucleus of one passing into the other sporidium. When
this germinates it gives rise to a dicaryon mycelium. Both Dickinson
(1927, 1928) and Boss (1927) showed that the dicaryon phase initiated by
the union of two sporidia is often only transitory in cultures of the fungus,
the further growth of the mycelium consisting of monocaryon hyphae,
some of one, some of the other sexual phase. For such species it seems
probable that only when the union occurs in the tissues of the host is the
dicaryon phase permanent. The binucleate sporidium may produce a
binucleate conidium. Rawitscher (1912) and others have shown that
instead of producing sporidia the promycelial cells of the opposite sexual
phase may conjugate by short conjugation tubes, through which the
nucleus of one of the cells passes into the other. This latter then produces
binucleate sporidia. It may happen that uninucleate sporidia germinate
and produce monocaryon mycelium. When two hyphae of such mono-
caryon mycelia of opposite sexual phases meet they unite and a dicaryon
mycelium results. A uninucleate conidium from one mycelium may unite
with a hypha of a mycelium of opposite sexual phase, etc. Bauch (1922)
showed that in Ustilago violacea (Pers.) Fuckel, the two sexual phases of
mycelium differ in their reaction to various nutrients present, as evinced
by the degrees of growth and distribution of the mycelia in various culture

ORDER USTILAGINALES (tHE SMUTS)

415

Fig. 140. Subclass Teliosporeae, Order Ustilaginales, Family Ustilaginaceae,
Ustilago hordei (Pers.) Kellerm. & Swingle. Normal and abnormal germination of
teliospores. (A-G) Stages in the normal development. (H, I) Conjugation between
promycelial cells of opposite sexual phase. (After Htittig: Z. Botan., 24(6):529-577.)

media. Both of these differ from the mode of growth of the dicaryon
mycehum produced when the two cultures are allowed to grow together.
Dickinson (1927, 1928) showed for Ustilago levis (Kellerm. & Swingle)
Magn., causing smut on oats, that infection does not take place unless
mycelia of two opposite sexual phases are present. Monocaryotic struc-
tures do not infect the host plants. It must be noted that Flerov (1923)
studied a strain of U. avenae (Pers.) Jens., also causing a smut on oats, in
which a monocaryon sporidium brought about infection by a monocaryon
mycelium which eventually produced uninucleate teliospores in which no
nuclear fusion occurred. From such teliospores arose a two-celled pro-
mycelium. Boss (1927) found the same to be true in Sphacelotheca ischaemi
(Fuckel) Clinton {Ustilago ischaemi). These are similar to the monocaryon
strain of Kunkelia nitens studied by Dodge (1924). (Fig. 140.)

Hanna (1929b) demonstrated for U. zeae on maize that single sporidia
or conidia from cultures from single sporidia are able to infect the tender
meristem wdth a very slender germ tube composed of monocaryon cells.
This infection is usually of very limited extent and except in rare strains
no smut galls or teliospores are produced. When two sporidia of opposite
sexual phase infect the tissue in rather close proximity the slender mono-
caryon hyphae approach one another and unite and thenceforth develop
as a stout mycelium of dicaryon cells which penetrates the meristem in
all directions. These hyphae show numerous clamp connections. Eventu-
ally a smut gall is produced, filled with numerous teliospores. Similar
conditions exist according to this author in Sorosporium reilianum (Kiihn)
McAlpine, another species producing smut galls on maize. Christensen
and Stakman (1926) studied various mutations of corn smut. Christensen
(1929) reported three strains in which infection and production of large
galls occurred with monosporidial cultures. He did not follow the cyto-

416 CLASS BASIDIOMYCETEAE

logical phenomena of these strains within the host. Similar results were
obtained by Eddins (1929) in his studies of the same species. (Fig. 138B.)
Aside from the distribution of the two sexual phases in the promycelial
cells of Ustilago levis Dickinson studied the distribution of the factors for
color of the mycelium and for form of the colony when grown in culture.
These studies were made by isolating and culturing the sporidia, noting
their position on the promycelium. The smut used was a cross of strains
producing yellow and white mycelium and corrugated or depressed colo-
nies. Out of 22 such isolations the sexual strains A and B occurred in the
following order from the apex toward the base of the promycelium:
A ABB (3 times), BBAA (5 times), ABAB (6 times), BAB A (twice),
ABBA (3 times), and BAAB (3 times). The distribution of the other two
sets of allelomorphic factors was apparently entirely independent of the
distribution of the sexual strains and of each other.

Hybridization has been brought about in cultures and also appears to
occur in nature. E. and J. Hirschhorn (1935) have shown that in Argentina
there may be present in the same smut gall on maize the three species
Ustilago zeae (Beckm.) Unger, U. fischeri Pass., and Sorosporium reilianum
(Kiihn) McAlpine, and that there occur crosses between all three, so that
all types of intermediate forms may occur together. Only very rarely are
U. zeae and S. reilianum found alone in the pure state in Argentina.
Fischer and Holton (1941) report the result of crosses between U. avenae
(Pers.) Jens, on oats (Avena saliva L.) and U. perennans Rostr. on Ar-
rhenatherum elatius (L.) Mert. & Koch. The race of the former that was
used had naked sori with indurated spore masses while the latter had
covered sori (i.e., the glumes were not entirely destroyed by the fungus)
and powdery spores. The Fi grew on A. fatua L. but not on Arrhenath-
erum. The F2 generations grew on A. saliva but still not on the tall oat-
grass. It was demonstrated that the covered character is recessive to
naked and the indurate recessive to powdery. Other hj^brids between
species of Ustilaginaceae have been reported by Dickinson (1927-1928),
Hanna and Popp (1931), Kniep (1926), and others.

The occurrence of geographic races has been studied intensively in
Ustilago longissima (Sow.) Tul., by Bauch (1930, 1931). He showed that
multiple allelomorphy occurs in both the A and B factors.

That the Ustilaginaceae are degenerate forms seems to be indicated by
the fact that their types of sexual reproduction are not at all standardized.
In the majority of cases studied the teUospore upon germination produces
a four-celled promycelium, from each cell of which one or more sporidia
are produced, but this is not universal. In some species germ tubes take
the place of the sporidia. In Ustilago striiformis (West.) Niessl. several
races are recognized on different hosts (Davis, 1935, Fischer, 1940).
Fischer has shown that the teliospores of forma Hordei Fisch. germinate

ORDER USTILAGINALES (tHE SMUTS) 417

by the production of promycelia and sporidia, and that the latter pair by
twos and then give rise to actively growing hyphae. On the contrary in
forma Poae-pratensis Davis it was shown by Leach and Ryan (1946) that
the teliospores germinate by the production of branched germ tubes of
indeterminate length. The earlier divisions of the teliospore nucleus are
regarded as meiotic so that this branching mycelium is considered to be
haploid. Here and there in this mycelium the nuclei begin to assort in
pairs and to undergo fusion. These hyphae wdth diploid nuclei grow and
the cells divide but soon the cells separate and form teliospores by thicken-
ing and becoming fusiform or limoniform with thickening and darkening
of the spore wall. These phenomena can be followed in artificial cultures
on agar.

Another indication of the degeneration of this family is the fact that
in some species of Ustilago no promycelium is formed but from the telio-
spore there grow out cells that form short branching hyphae of yeast-like
cells. Bauch (1923) found that in Ustilago longissima and its variety
macrospora Davis the meiotic divisions of the diploid nucleus .of the ma-
ture teliospore take place in the teliospore itself and that then successively
sporidia are budded off from the latter. Into the first such sporidium two
nuclei enter while into those subsequently produced one nucleus may enter
or two, the latter being made possible by division of the nuclei remaining
in the teliospore. More often the two nuclei in the first sporidium are of
opposite sexual phase. In that case the sporidia become two-celled by the
formation of a septum and then these two cells unite through conjugation
tubes and grow out as slender dicaryon hyphae. The other sporidia if
uninucleate mostly remain one-celled but conjugate with sporidia of op-
posite sexual phase and then they too form the slender dicaryon "Such-
faden." Onl}^ under exceptional circumstances is a typical four-celled
promycelium formed, from the cells of which are budded off the sporidia.
The teliospores of the Ustilaginaceae vary in their longevity. Fischer
(1936) reports viable spores in herbarium specimens up to 3-10 years in
many cases and as high as 23 years for Ustilago hordei (Pers.) Kellerm. &
Swingle.

Mutant forms have been observed frequently in smuts. Johnson et al.
(1940) report the occurrence of a mutant race of Sorosporium syntherismae
(Pk.) Farl., in which the teliospores are almost colorless but still retain
their ability to infect the host.

When grown in liquid culture media with an abundance of soluble
carbohydrates the mycelium of smuts may break up into separate cells
resembling and multiplying like the asporogenous yeasts. Teliospores
sown in such media will frequently produce hyphae which become yeast-
like, instead of typical promycelia.

The family includes over 450 species in about 12 genera. About 61 per

418

CLASS BASIDIOMYCETEAE

cent of the North American species occur in members of the grass family
(Poaceae) and about 14 per cent on sedges (Cyperaceae) with about 19
per cent in Dicotyledons, particularly in the Polygonaceae and the Aster-
aceae. In the majority of species the spores are produced in the ovaries or
other parts of the flower, or the whole inflorescence may be involved. Some
species produce their sori in the leaves or stems of their hosts.

The largest genus is Ustilago with over 300 species. Its spores are pro-
duced singly in a powdery mass. Many serious enemies of cultivated
plants are found in this genus, e.g., U. avenae (Pers.) Jens, and U. levis
(Kellerm. & Swingle) Magn., destroying the spikelets and inflorescence of
oats {Avena saliva L.), U. zeae (Beckm.) Unger, producing smut galls on
maize {Zea mays L.), U. hordei (Pers.) Keflerm. & Swingle and U. nuda
(Jens.) Kellerm. & Swingle on barley {Hordeum sativum L.), U. tritici
(Pers.) Rostr. on wheat {Triticum spp.), U. striiformis (West.) Niessl on
Poa pratensisL., U. violacea (Pers.) Fuckel on Dianthaceae, etc. Sphacelo-
theca differs from Ustilago in having the powdery mass of teliospores sur-
rounded by a pseudoparenchymatous layer of fungus tissue. S. sorghi
(Link) Clinton is injurious to the ovaries of Sorghum vulgare Pers. and
related species. In Schizonella the teliospores are in twos, otherwise much
as in Ustilago. S. melanogramma (DC.) Schroet. produces long black sori

Fig. 141. Subclass Teliosporeae, Order Ustilaginales, Family Ustilaginaceae. (A)
Spore ball of Sorosporium saponariae Rud. (B) Spore Imll of Tohjposporium junci
(Schroet.) Woron., some of the spores germinating to form promycelia. (C) Ustilago
kuehneana Wolff, teliosporo germinating with whorls of sporidia at the septa of the
promycelium. (D) Schizonella melanogramma (DC.) Schroet. Tc^liospores joined in
twos. (A, after Dietel, in Engler und Prantl: Die Natiirlichen Pflanzenfamilien,
Zweite Auflage, vol. 6, pp. 1-98, Leipzig, W. Engelmann. B-C, after Brefeld: Unter-
suchimgen aus dem Gesammtgcbiete; der Mykologie, vol. 12, pp. 99-236. D, after
Clinton: Connecticut State Geological and Natural History Survey Bull, 6:1-44.)

ORDER USTILAGINALES (tHE SMUTs) 419

in the leaves of various species of Carex. In Sorosporium, Tolyposporium
and other genera the teliospores are united into more or less firm balls.
They are largely parasitic on grasses. (Fig. 141.)

Family Tilletiaceae. Promycelium nonseptate, the sporidia being
formed at its apex in a dense cluster or in a whorl. The number of sporidia
varies usually from 4 to 30-50, but in Neovossia indica (Mitra) Mundkur
may reach 150 (Mundkur, 1943). The teliospores arise as terminal cells of
hyphae or of short lateral branches or as intercalary cells. The two nuclei
unite and the cells round up and secrete a heavier wall which may be dark
or light in color and is smooth or more often reticulately marked or spiny.
The teliospores occur as single spores in a dusty mass or united into small
or large balls of spores with or without a covering or a core of sterile cells.
These spores or spore balls may escape as a powdery mass or remain
within the host tissue. The sporidia are fusiform or sickle-shaped and are
inclined to unite by twos while still attached to the promycelium or after
they have been shot off. A binucleate conidium may be produced directly
from a pair of united sporidia. As in the preceding family the sporidia
appear to be of at least two sexual phases. From one of the sporidia of the
united pair a germ tube of dicaryon cells may grow out and infect the host
plant. Infection may take place from a dicaryon conidium set free from
the united sporidia or arising on a dicaryon mycelium. The sexual cycle
which is initiated by the fusion of the sporidia is completed by the union
of the nuclei in the teliospore. (Fig. 142A, B.)

About 13 genera and over 250 species are recognized in the family.
Tilletia (about 40 species) corresponds to Ustilago in producing its telio-
spores as single cells in a powdery mass. T. caries (DC.) Tul. (T. tritici
(Bjerk.) Wint.), with rough teliospores and T. foetida (Wallr.) Liro {T.
levis Ktihn), with nearly smooth teliospores, cause stinking smut or bunt
of wheat. Flor (1932) has crossed these two species by picking off single
sporidia and allowing the two monocaryon myceha produced from them
to unite. These hybrid mycelia were used successfully to inoculate wheat
plants. The teliospores produced on these plants resembled most closely
those of T. foetida. Certain species of Tilletia have been described from
the capsules of Sphagnum and Anthoceros, both in the Bryophyta, but
Bauch (1938) studying T. sphagni Nawaschin has demonstrated that it
does not belong to this order, the supposed teliospores actually represent-
ing the conidial stage of Helotium schimperi Naw., one of the Pezizales.

Entyloma produces its teliospores singly and in the tissues of the host
from which they do not escape as a powdery mass, but germinate within
the host sending their elongated promycelia out through the epidermis
and forming the sporidia externally. Conidia are also formed on conidio-
phores which emerge through the stomata. Kaiser (1936) observed clamp
connections on the mycelium of E. calendulae (Oud.) de Bary. Frequently

Fig. 142. Subclass Teliosporeae, Order Ustilaginales, Family Tilletiaceae. (A, B)

Tilletia caries (DC.) Tul. (A) Teliospore with sporidia united in H form. (B) The

conjugated sporidia have produced a dicaryon conidium. (C) Doassansia sagittariae

(West.) Fisch., with some of the teliospores germinating. (D) Section through a portion

of a spore ball of Doassnnsiopsis martianoffiana (Thiim.) Diet. (E, F) Tuburcinia

trientalis (B. & Br.) Wor. (E) Section of portion of infected leaf with external conidia.

(F) Germinating spore ball. (G) Germinating spore ball of Urocystis violae (Sow.)

Fisch. de Waldh. (A-B, after Plowright: A Monograph of British Uredineae and

Ustilagineae, London, Kegan, Paul, Trench, & Co. C, G, after Brefeld: Untersuchungen

Gesammtgebiete der Mykologie, Heft 12, pp. 99-236; D, after Diotel, in Engler und

Prantl: Die Natlirlichen Pflanzenfamihen, Zweite Auflage, vol. 6, pp. 1-98, Leipzig,

W. Engelmann. E-F, after Woronin: Abhandl. Senckenberg. Natur. GeselL, 12:559-

591.)

420

ORDER USTILAGINALES (tHE SMUTs) 421

the teliospores arise as outgrowths of these structures. Stempell (1935) ob-
tained cultures of Entyloma on agar media from conidia on infected leaves.
The mycelia were of two types, monocaryon and dicaryon, but clamp
connections were produced only on the latter. He reported that sickle-
shaped, uninucleate conidia were formed on the first type of mycelium
and lunate, binucleate conidia on the dicaryon type. Both types of conidia
are discharged violently with the formation of a droplet of liquid at the
apex of the sterigma, just as occurs in the discharge of the sporidia from
the promycelium. Hanna (1938) found that in several species of Entyloma
two types of conidia were produced, sickle-shaped, uninucleate conidia
which are discharged violently and slender filiform or needle-shaped
conidia which are not shot off. In the species studied by Hanna the sickle-
shaped spores were always uninucleate and lunate binucleate spores were
not observed. The 100 or more species are found on Grasses (Poaceae),
Ranunculaceae and other families, but particularly on the Asteraceae.
Urocystis (about 60 species) produces its teliospores in balls of from two
(rarely one) to four or five, the ball being partly or completely surrounded
by a layer of small sterile cells. The mass of spore-balls is powdery and
they escape upon rupture of the host tissue. U. occulta (Wallr.) Rab.
causes longitudinal, lead-colored, slightly raised sori on the stems and
leaf sheaths of rye {Secale cereale L.), the head being killed by the presence
of the fungus in the stalk below, it being rarely entered by the fungus.
The very similar U. tritici Korn, parasitic on wheat {Triticum aestivum
L.) has been shown by Yu and his associates (1936) to occur in several
physiologic races in China. U. violae (Sow.) Fisch. de Waldh. and U .
anemones (Pers.) Wint. form their sori in the leaves, respectively, of Viola
and of various species of the Ranunculaceae. In Tuhurcinia all the cells of
the spore ball are fertile spores. It must be noted that Liro (1922) and
others combine the genera Tuburcinia and Urocystis under the former
name which is the earlier. About 20 species of Doassansia produce their
sori in the leaves of various Alismataceae and related aquatic plants. The
large spore balls have very numerous teliospores and an external layer of
hyaline sterile cells. Doassansiopsis, also on Alismataceae, produces large
spore balls made up of a central core of hyaline pseudoparenchymatous
cells surrounded by a single layer of larger dark teliospores, these in their
turn being surrounded by a filamentous sheath. (Fig. 142C-G.)

As in the preceding family, the teliospores in the Tilletiaceae may live
a long while. Fischer (1936) obtained germination from specimens pre-
served in the herbarium for 10 years in the case of Entyloma dahliae Syd.
and for 25 years for Tilletia foetida (Wallr.) Liro.

Family Graphiolaceae. Parasitic in the leaves of palms. Sori formed
under the epidermis and immediately underlying tissues and tearing these
so as to permit the emergence of the spores. They consist of a thick, cup-

422

CLASS BASIDIOMYCETEAE

like, dark outer peridium surrounding the sporogenous central portion
which may project some distance above the rim of the opened peridium.
A thin hyaline inner peridium may surround this projecting mass of
sporogenous hyphae. The spores are formed in parallel chains and bud
laterally to form two to four (or more) sporidia which become more or less
colored, with somewhat thickened walls. Bundles of sterile hyphae scat-
tered throughout the chains of spores probably serve the same function
as the capillitium in Mycetozoa, Lycoperdaceae, etc. They are lacking in
one genus. (Fig. 143.)

Graphiola has been carefully studied as to its morphology by Eduard
Fischer (1883, 1920, 1922) and by Kilhan (1924) who showed the cyto-

FiG. 143. Subclass
Teliosporeae, Order Us-
tilaginales (?), Family
Graphiolaceae. (A, B)
Graphiola phoenicis
(Moug.) Poit. (A) Sorus
on leaf of palm. (B)
Sporogenous hyphae, the
cells near the top produc-
ing sporidia. (C, D)
Graphiola thaxteri Fisch.
(C) Group of four sporidia
from one spore, three of
them once septate. (D)
Bundle of sterile hyphae
in sorus. (A-B, after
Fischer: Botan. Ztg., 41
(45) -.745-756. C-D, after
Fischer: Ann. Mycolog.,
20(3-4) :228-237.)

logical features. The sporidia are uninucleate and produce a monocaryon
mycelium in the host leaf. Eventually a mass of monocaryon hyphae is
formed beneath the epidermis and immediately underlying layers of cells
and from the marginal portiiju of the cushion arise vertically closely
packed, branched, thick- walled monocaryon hyphae that form the outer
peridium. This arches over the whole structure at first. From the central
portion of the hyphal cushion arise the sporogenous hyphae and, if pres-
ent, the sterile hyphal bundles. The cells are all at first monocaryon, and
this remains true of the cells of the hyphal bundles. The cells of the closely
packed sporogenous hyphae elongate rapidly and become multinucleate,
but very soon cross walls divide these hyphae into chains of dicaryon cells.
These soon show nuclear fusion. At this stage the sporogenous hyphae

RELATIONSHIPS WITHIN THE SUBCLASS TELIOSPOREAE 423

show at their base monocaryon cells, a little further up cells with several
nuclei, followed by a series of dicaryon cells, while the upper portion con-
sists of cells each with a single diploid nucleus. These, according to Kil-
lian, are the cells which correspond to the teliospores of the Ustilagin-
aceae. In them the nucleus undergoes two divisions and the four nuclei
pass out into the sporidia which bud out of the teliospores. The latter may
remain attached, so that the upper portion of the sporogenous hypha
shows teliospores with sporidia in various stages of development, or the
teliospores may break apart before the sporidial development is com-
pleted. The sporidia may remain one-celled or may become two-celled by
the formation of a septum. In their germination they may bud like yeasts
or form germ tubes. The true relationship of this family is not a matter of
agreement among mycologists. Fischer and Killian incline to the idea of
kinship with the Ustilaginaceae in some of which (e.g., Sphacelotheca) a
peridium of hyphal tissue surrounds the sporogenous part of the sorus.
The budding of the sporidia from the teliospores instead of the formation
of typical promycelia caused Killian to refer to the somewhat similar case
in Ustilago longissima referred to previously in this chapter. Fischer
(1922) recognized two genera: Graphiola, with bundles of sterile hyphae
among the sporogenous hyphae, the latter separating into their individual
cells at maturity, and Stylina which lacks the sterile hyphae and whose
sporogenous hyphae do not separate into individual cells. He described
the four species of Graphiola studied by him and the one species of Stylina.

Relationships Within the Subclass Teliosporeae

It is very apparent that there is not a very close relationship between
the more highly developed Uredinales and Ustilaginales. In vegetative
structures the similarities are greater, the mycelium in both orders being
intercellular with nucleated haustoria. Both types of mycelium are
present, monocaryon and dicaryon, although the former may have a
very brief life, especially in the parasitic life of the Ustilaginales. Eventu-
ally in both orders the dicaryon mycelium produces special cells, the
teliospores, within which the nuclei unite to form a diploid nucleus. This
nucleus undergoes two meiotic divisions to form four haploid nuclei
which apparently in both Rusts and Smuts are two of one sexual phase
and two of the other. The meiotic divisions may occur in the teliospore
but most often occur in the promycelium. In Ustilaginales the dicaryon
phase of the mycelium frequently bears clamp connections but these have
been demonstrated rarely in the Uredinales. In the sexual reproduction
the Ustilaginales produce no definite male gametes. Any two cells of
opposite sexual phase may unite to initiate the dicaryon phase, be it near-
by promycelial cells, sporidia, conidia, or mycelia. In the Uredinales two
mycelia of opposite sexual phase may diploidize one another when they

424 CLASS BASIDIOMYCETEAE

come in contact within the host but the usual mode of reproduction is by
the union of sperm cells with special receptive hyphae of the opposite
sexual phase. The Rusts have reached a much higher stage of specializa-
tion of spore forms and of adaptation to alternate hosts, while the Smuts
are parasitic but able to grow saprophytically, thus eliminating the neces-
sity for two growth phases in the same or different species of hosts. Be-
cause of the formation of sperm cells in typical spermogonia and the pro-
duction of receptive hyphae the Rusts in this regard hark back to more
primitive ancestral forms among the Ascomyceteae where such struc-
tures are present. Clearly the Ustilaginales must have branched off from
the earlier Rusts, with loss of these special organs, but at an early stage
when the inherited tendency to produce clamp connections had not been
lost. Jackson (1931) points out the similarity in life cycles of many Rusts
and Florideae. This is worthy of further careful consideration. The rela-
tionships of the Teliosporeae to the Subclass Heterobasidiae are discussed
in the next chapter.

Key to the Orders and Families of Subclass Teliosporeae

Obligate parasites in Pteridophyta, Strobilophyta (Coniferae) and Anthophyta
(Angiospermae) . Teliospores single or united into crusts or columns or
several together in compound spores, remaining within the host tissue or
bursting through the epidermis or cuticle. Spermogonia normally produced,
the sperm cells diploidizing special receptive hyphae. Typically three types of
spores are produced, aeciospores, the product of the diploidization of mono-
caryon mycelium which arises from the sporidia; urediospores (repeating
spores) and teliospores from which arise the promycelia and sporidia. Sporidia
always expelled violently. Order Uredinales

Teliospores without stalks, produced singly or in groups of two to four in the
mesophyll or just below or within the epidermal cells or united laterally into
subepidermal or subcuticular crusts or united into separate vertical chains
or into chains that are joined laterally into a waxy column which emerges
through the epidermis. Aecia mostly on species of Pinaceae.

Family Melampsoraceae

Teliospores usually stalked, simple or compound, sometimes without stalks and
produced successively as simple or compound teliospores which escape from
the sorus dry or embedded in slime. Aecia only very exceptionally produced
on Pinaceae. Family Pucciniaceae

Obligate parasites in Anthophyta or in many cases facultative saprophytes. Telio-
spores single or united in columns or balls, remaining within or bursting out
of the host tissue, mostly distributed by air currents. No spermogonia or
special receptive hyphae. Diploidization by means of union of compatible
spores, hyphae, etc. Typically only teliospores and oftien hyaline thin-walled
conidia are produced. Sporidia expelled violently in one family, not so in the
two others. Order Ustilaginales

Promycelium transversely septate into several, mostly four, cells. Teliospores
arising in the tissues of the host from transformed hyphal cells, and mostly
distributed by air currents. Sporidia not expelled from the promycelium.

Family Ustilaginaceae

KEY TO THE COMMONER NORTH AMERICAN GENERA OF FAMILY PUCCINIACEAE 425

Promycelium not septate, the four to many sporidia at its blunt apex. Telio-

spores mostly arising as lateral outgrowths from hyphal cells or intercalarly.

Sporidia expelled from the promycelium. Family Tilletiaceae

Promycelium lacking, but the teliospores bud directly to form four sporidia

which form thick, dark walls. Teliospores in vertical rows in compact sori.

Parasites in leaves of palms. Sporidia not expelled violently.

Family Graphiolaceae

Key to the Commoner North American
Genera of Family Melampsoraceae

(Based upon Arthur, 1934)

Teliospores single or united laterally into groups of two or more.
Telia in ferns (Polypodiaceae and Osmundaceae), aecia in Abies.
Teliospores subepidermal. Uredinopsis

Teliospores in the epidermal cells.

Aeciospores and urediospores colorless. Milesia

Aeciospores and urediospores with yellow contents.

Hyalopsora
Telia in Anthophyta (Angiospermae) .

Teliospores with brown walls. Pucciniastrum

Teliospores with colorless walls.

Peridium of uredium delicate, with a central pore.

Melampsorella
Peridium of uredium firm, with long-pointed ostiolar cells.

Melampsoridium
Teliospores united laterally into subcuticular or subepidermal crusts.
Promycelium external.

Teliospore walls colored. Melampsora

Tehospore walls colorless. Aplospora

Promycelium within the teliospore. Coleosporium

Teliospores in chains which are united laterally into long columns emerging
through the epidermis. Urediospores borne singly on stalks.

Cronartium
TeUospores in chains which are not united laterally and not emerging as a column.

Urediospores in chains. Chrysomyxa

Teliospores united laterally into crusts of two or more layers, mainly tropical.
Life histories not well known. Bubakia, PhysopeUa,

Cerotelium, etc.

Key to the Commoner North American Genera of Family Pucciniaceae

Teliospores borne singly.

Teliospores colorless; aecia peridermioid; autoecious. Mainsia

Teliospores colored.

TeUospores flattened vertically; aecia uredioid; autoecious.

Pileolaria
Teliospores spherical or short ellipsoidal, aecia uredioid, autoecious; spermo-

gonia subcuticular.' Trachyspora

TeUospores spherical or short ellipsoidal, with thickened apex; aecia cupulate
(rarely uredioid); autoecious or heteroecious; spermogonia sub-
epidermal. Uromyces

426 CLASS BASIDIOMYCETEAE

Teliospores compound, two to a pedicel (see also Earlea).

Teliospores easily separable, not surrounded by a conspicuous common mem-
brane, pedicels fascicled at base; spermogonia subcuticular.

Tranzschelia
Teliospores not separable, surrounded by a distinct common membrane.
Outer membrane gelatinizing when wet; spermogonia subcuticular.

Uropyxis
Outer membrane not gelatinizing when wet (except in the case of
Gymnosporangium) .
Spermogonia subcuticular; aecia caeomoid; no uredia. Gymnoconia
Spermogonia subepidermal.

Pedicel and teliospore Avail gelatinizing when wet; aecia cupulate or
cornute; uredia wanting except in one species. Gijmnosporangium
Pedicels not gelatinous when wet; aecia cupulate or (more rarely) ured-
ioid.; apex of terminal teliospore thickened or papillate.

Puccinia
Teliospores compound, several in a row (one species of Earlea has simple telio-
spores and several only two teliospores to a pedicel); spermogonia
subcuticular.
Teliospores with two to three lateral pores, smooth or more often verrucose;
basal portion of pedicel inflated in water; uredia present; aecia
caeomoid. Phragmidium

Like the preceding but basal portion of pedicels not inflated in water; uredia
absent except in one species; aecia caeomoid (often included in
Phragmidium) . Earlea

Terminal teliospores with one pore, the others with two, near the top; mostly
smooth, pedicel short, not swelling in water; uredia wanting; aecia
caeomoid. Xenodochus

Teliospores with one pore in each cell, firmly united, smooth; pedicel elongated,

not swelling in water, aecia uredioid. Frommea

Teliospores with one pore in each cell, smooth, easily separable, pedicel very
short; aecia uredioid. Kuhneola

Teliospores compound, consisting of a basal teliospore attached to a long pedicel
and two horizontally lying teliospores at its top; aecia uredioid,
spermogonia subcuticular.
Teliospores verrucose; uredia with paraphyses. Triphragmium

Teliospores spiny; uredia wanting. Nyssopsora

Teliospores numerous in a head, subtended by hyaline cysts.

Ravetidia

Key to the Commoner Mostly North American
Genera of Family Ustilaginaceae

{Based on Clinton, 1906)

Teliospores separate.
Sori dusty at maturity.

Without definite false membrane. Ustilago

With false membrane of fungus cells. Sphacelotheca

Spores more or less agglutinated at maturity.

Spores firmly agglutinated into tubercular nodules. Melanopsichum

Spores developed around a central columella (rarely dusty).

Cintradia

LITERATURE CITED

427

Teliospores mostly adhering in pairs.

Agglntinated, in leaves. Schizonella

Dusty, in inflorescences. Mycosyrinx
Teliospores in balls.
Sori dusty or granular.

Spore balls often evanescent; olive-brown or black-brown. Sorosporium

Spore balls rather permanent; yellowish or reddish. Thecaphora
Spore balls rather permanent; spores adhering by folds or thickenings of the

outer coat. Tolyposporium
Sori agglutinated.

Spore balls of thick-walled spores. Tolyposporella

Spore balls of central sterile cells surrounded by the ])eripheral functional

teliospores. Testicularia

(It must be noted that Ustilago and Sphacelotheca are in some cases distin-
guished with difficulty. Furthermore, in some species of Sorosporium the telio-
spores do not remain adherent in the spore balls, so that they are not easily
distinguishable from the other genera. In some genera the germination of the
teliospores has not been studied, or perhaps in only a few species of the genus.)

Key to the Commoner Genera of Family Tilletiaceae

Teliospores single.

Spores dusty and escaping at maturity. Tilletia

Spores remaining embedded in the host tissue. Entyloma

Teliospores in balls.

Sori dusty ; spore balls more or less surrounded by an adhering layer of sterile

cells; escaping from the ruptured sorus. Uroajstis

Spore balls more or less permanently remaining in the host tissue.
Spore balls lacking a cortex of sterile hyphae.

Spore balls dark, no sterile core. Tuhurcinia

Spore balls light-colored, with or without a core of sterile pseudoparen-

chymatous cells. Burrillia

Spore balls light-colored, with a core of septate hyphae. Tracya
Spore balls with filamentous cortex.

No central core of sterile cells. Doassansia

A single layer of spores surrounding a large core of pseudoparenchymatous
cells. Doassansiopsis

Key to the Genera of Family Graphiolaceae

Bundles of sterile hyphae among the sporogenous hyphae; the latter separating at
maturity into their individual teliospores. Graphiola

No bundles of sterile hyphae present; the sporogenous hyphae do not separate at
maturity into their individual teliospores. Stylina

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LITEKATURE CITED 435

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CLASS BASIDIOMYCETEAE:
SUBCLASS HETEROBASIDIAE

IN THE basidial structure of one of its orders, the Auriculariales, this
subclass shows many points of similarity with the more typical forms
of Subclass Teliosporeae. By some authors (e.g., Patouillard, 1900) the
latter are included in this order. Taking all their characteristics into con-
sideration it seems to the author to be a more satisfactory arrangement to
keep them apart in separate, but more or less closely related, subclasses.
Thus limited the Heterobasidiae consist of fungi most often living as
saprophytes upon wood or other plant materials but in some cases para-
sitic upon living plant tissues or even upon insects (e.g., Septobasidium) .
They form their spore fruits as thin or thick layers or projecting cushions
or clubs or shelves of basidium-forming tissue. The basidia are formed
mostly in a more or less recognizable layer, the hymenium, often crowded
side by side in a palisade-like arrangement, and shoot their basidiospores
off into the air with the exception of one or two genera. The basidiospores
frequently become septate as they mature or after they are discharged
and as a general rule (to which there are exceptions), germinate by the
formation of numerous small buds when they fall into water, though
capable of producing germ tubes under favorable conditions. In many
cases a basidiospore will form a small sterigma and a secondary spore will
be shot off, just as occurs frequently in the formation of secondary spo-
ridia in the Uredinales. The spore fruits are more often waxy, or gelatinous
when wet, and frequently dry down to rather inconspicuous horny masses
when dry. This is made possible by the fact that the outer portion of the
wall of each hypha swells greatly upon absorbing water.

The basidial characters that further distinguish this subclass from
Subclass Eubasidiae are the following: In the Eubasidiae the basidium
remains as one cell usually bearing the Ijasidiospores at its upper end on
short or long sterigmata. Tulostoma and one or two related genera form
an exception in that the sterigmata may be formed laterally on the

436

SUBCLASS HETEROBASIDIAE 437

basidium. In the Heterobasidiae the basidium may be (1) elongated and
transversely divided by three septa into four cells from each of which a
sterigma arises and produces a single basidiospore, or (2) it may be
rounded and divided into four cells by vertical septa, each cell producing a
sterigma and a basidiospore, or (3) elongated and forked into two prongs,
at the apex of each of which a sterigma bears one basidiospore, or (4) the
rounded basidium produces (usually) four rounded cells at its upper end,
these usually being separated by a septum from the collapsed basidium
and frequently falling free from it. Each of these cells produces a sterigma
and a basidiospore. In all of these four types, and in the Eubasidiae, the
basidia arise as enlarged binucleate cells on mycelium of the secondary
(dicaryon) type. Mostly these cells are terminal to the hyphae bearing
them but sometimes (Sirohasidium) several cells at the terminal portion
of the hypha become transformed into a chain of basidia. In a few cases a
single intercalary cell may become a basidium.

In this subclass the author recognizes four orders, based upon the four
types of basidia. It must be noted that Martin (1944) considers these to
be more closely related and unites them into one order, Tremellales. The
orders recognized by the author are:

Auriculariales : with elongated basidium divided into a row of four cells by
cross septa. Nuclear divisions stichobasidial.

Dacrymycetales : with nonseptate forked basidium ("tuning fork" type) bear-
ing two basidiospores. Nuclear divisions stichobasidial.

Tremellales: with rounded basidium divided cruciately into four cells by ver-
tical septa. Nuclear divisions chiastobasidial.

Tulasnellales : with rounded basidium bearing large rounded epibasidia, usually
separated from the hypobasidium by septa and often falUng free. Nuclear
divisions chiastobasidial.

In these four orders there is a marked parallelism of evolution of spore
fruit structure from effused, sometimes almost film-like, structures to
cushions, shelves, stipitate pilei, etc.

The arrangement followed below is not to be considered as represent-
ing a single progressing line of evolution. On the contrary the groups have
been produced by a more or less parallel evolution from primitive forms
that have been lost. The orders Auriculariales and Tremellales are some-
times set apart from the other orders as the Phragmobasidiae, since the
basidium is divided into four cells. The Dacrymycetales and Tulasnel-
lales with undivided basidia are then united with the remaining orders of
the class under the name Holobasidiae. Gilbert (1921) has studied the
nuclear behavior in the Heterobasidiae and finds that in their essentials
they are identical with those occurring in the Eubasidiae.

Within this subclass the basidium in many genera has been con-
sidered as being made up of two parts, hypobasidium and epibasidium

438 CLASS BASIDIOMTCETEAE

or epibasidia (Juel, 1898; Neuhoff, 1924; Rogers, 1934). These are then
homologized with the tehospore and promycehum (as hypobasidium and
epibasidium respectively) of the Uredinales and Ustilaginales. Onto-
genetically, where this distinction can be made, the hypobasidium is a
more or less spherical dicaryon cell within which the fusion of the nuclei
occurs. It may remain thin-walled and proceed immediately with the
formation of the epibasidium or may develop a thicker wall and become a
resting cell with diploid nucleus, from which, under proper conditions the
epibasidium then grows.

The difficulty with the foregoing interpretation is that in some of these
fungi there is no such distinction in some species of a genus while it is
pronounced in other species. Furthermore, different structures have been
designated as hypobasidium and epibasidium. In the Auriculariales, in the
genus Auricularia, the somewhat elongated dicaryon basidium primor-
dium elongates after the nuclei have united and then meiosis occurs and
the four nuclei become separated by septa. There is no hypobasidium nor
epibasidium that can be distinguished. In some species of Septohasidium
the basidium primordium becomes a somewhat thick-walled resting cell,
and when favorable conditions come on this sends out a short stout hypha
like the whole basidium of Auricularia, within which meiosis occurs and
septa are formed. In other species no hypobasidium is formed at all and
within the basidial primordium occur the union of the two nuclei, and
their meiotic division, followed by septation to form the four celled
basidium. In still other species of this genus Avhether the resting stage,
usually interpreted as a hypobasidium, develops or not depends upon the
environment. Thus it is apparent that this is perhaps not such a funda-
mental character as has been supposed. In other words in the development
of the basidium in this group the presence or absence of a hypobasidium
and consequently of an epibasidium depends upon whether the ontogeny
of the basidium is interrupted or not.

In the type of basidium found in the Dacrymycetales the basal portion
is interpreted as the hypobasidium and the two arms as epibasidia. But
here the meiotic divisions occur in the "hypobasidium," not in the "epi-
basidium" as in Septohasidium. In Tremella the "hypobasidium" is the
rounded basal portion within which meiosis occurs before the vertical
walls divide it cruciately into four cells. The four arms at whose tips the
sterigmata produce the basidiospores are interpreted as "epibasidia." In
TulasncUa the same structures are found, except that the "hypobasid-
ium" is not septate but is separated from the four epibasidia by septa.

Order Auriculariales. The fungi included in this order are some of
them cosmopolitan but many are confined to the Tropics. They are largely
saprophytes on wood. A few are true parasites, such as some species of
Eocronartium and Jola, occurring on mosses; Septohasidium and Ured-

ORDER AURICULARIALES 439

inella, parasitizing upon scale insects; Hei-pohasidium filicinum (Rostr.)
Lind and Platycarpa upon ferns; etc. According to Boedijn and Steinmann
(1931) the East Indian species of Helicohasidium are parasitic upon roots,
while H. purpureum (Tul.) Pat. is parasitic on various plant roots and
crowns in Europe.

Clamp connections have been recognized in Auricularia, Phleogena,
Helicohasidium, Jola, Helicogloea, etc. H. L. Barnett (1937) reported that
the mycelium of Auricularia derived from single spores, and therefore
monocaryotic, lacks clamp connections but that when two such mycelia
of opposite sexual phase come into contact the resulting dicaryotic myce-
lium may be recognized immediately by the presence of these structures.

The spore fruits are external and vary from a more or less felt-like
film to a thin crust or to firm shelf-like structures standing out from the
substrata. In Phleogena they are upright and stipitate with an enlarged
head. In size they vary from a few millimeters to several centimeters.
Septobasidium forms a more or less felted layer over the twigs and larger
branches of the host with various tunnels and chambers in which the scale
insects are protected or through which the larvae may travel. The basidia
are formed on the outer surface of the felted mycelium. In some genera
the spore fruits are gelatinous when wet, drying down to a horny crust or
cushion. Other genera on the contrary, such as Phleogena and Septo-
basidium, do not become gelatinized with moisture.

In the gelatinous types of spore fruits, e.g., Auricularia, the basidia
may be situated below the surface or at least with only the upper end
reaching the air. This necessitates the formation of tube-like extensions
from the four cells of the basidium to the surface where each develops a
sterigma and bears a spore. This is very similar to the structure of the
germinating teliospore of Coleosporium. Where the structure is not highly
gelatinized the basidia are superficial and the sterigmata are very short,
without any elongated supporting tubes, e.g., Phleogena.

The basidia are usually the terminal cells of the hyphae but in some
species of Helicogloea {Saccoblastia) the basidial primordia may be inter-
calary according to Miss Baker (1936).

No distinction of hypobasidium and epibasidium is apparent in Au-
ricidaria, Phleogena, and Herpobasidium. Within the basidial primordium
the two nuclei unite and then undergo "the two meiotic divisions, the
spindles of the dividing nuclei being parallel to the long axis of the enlarg-
ing cell, thus producing a stichobasidial structure. In Helicogloea {Sacco-
blastia) the primordial cell of the basidium produces a lateral sack-like
"hypobasidium" in which the union of the nuclei occurs. Then sometimes
at one end of this sack, but more often from the apex of the primordium
the "epibasidium" grows, and in it the meiotic divisions occur. Since this
genus includes species with greater or less gelatinization of the vegetative

440 CLASS BASIDIOMYCETEAE

hyphae there arise from the four cells of the epibasidium shorter or longer
arms reaching to the surface and there bearing the sterigmata and spores.
The basidial primordium may be an intercalary cell of the hypha although
more often it is terminal. In the genus Septohasidium it was pointed out by
Couch (1938) that the basidia among the more than 160 species represent
a number of different types. In the majority there is a distinct rather thin-
walled and hyaline hypobasidium which may send out the epibasidium
immediately or the hypobasidium may have a thick, colored wall and
serve as a resting spore until favorable conditions are present. The epi-
basidium is more often four-celled but in some species it is three-celled, in
others two-celled, and in two species it is one-celled. The basidiospores
become segmented into two to several cells and usually germinate by
sprout cells, when placed in water.

Perhaps three families should be recognized in this order, although
Gaumann and Dodge (1928) recognized four.

Auriculariaceae: parasitic on plants or saprophytic on dead plant material,
usually wood. Spore fruits often gelatinous but not so in some genera. Basidia
with no distinction into hypobasidium and epibasidium or these well dis-
tinguished. Clamp connections present in some genera, absent in others.

Phleogenaceae: saprophytic on wood, bark, etc. No distinction into hypo-
basidium and epibasidium. Spore fruit stalked with a head of radiating, more
or less coiled hyphae, among which the curved basidia are formed, bearing
the four spores without visible sterigmata. Clamp connections observed.

Septobasidiaceae (Order Septobasidiales, according to Couch, 1938): parasitic
upon scale insects with some of which they live in symbiotic relation. Basidia
usually with well-developed thin-walled or thick-walled hypobasidia and
one- to four-celled (mostly the latter) epibasidia, but hypobasidia sometimes
lacking. Basidiospores produced on distinct sterigmata. Conidia often pro-
duced. Clamp connections apparently lacking.

Family Auriculariaceae. In the Auriculariaceae the following gen-
era should be mentioned as illustrating the various types of structure.
Eocronartium, with a single species E. muscicola (Fr.) Fitzp. is a perennial
fungus parasitic in the gametophytes of many mosses (see Fitzpatrick,
1918a, b). The cells of the mycelium are always binucleate and there are
no clamp connections. The mycelium is intracellular, passing from cell to
cell of the host but apparently doing little harm except that the produc-
tion of the sporophyte appears to be suppressed in the infected plants. At
the apex of the stem the hyphae pass out into the spaces between the
leaves and grow upward and parallel and form a gelatinous, club-shaped
sporophore on the outer surface of which the basidia are produced in great
abundance. The longitudinal hyphae of the sporophore turn outward to
the surface and there give rise to the basidia which as they elongate bend
almost at right angles so as to lie nearly parallel to the surface. The basid-
ial primordium is at first cylindrical or clavate, the major portion (epi-

ORDER AURICULARIALES 441

basidium) becoming separated by a septum from the short cylindrical
thin-walled hypobasidium which is not enlarged as in some genera. The
epibasidia are transversely three septate and from each cell is produced a
long tube at whose apex a sterigmatic structure is formed on which arises
the one-celled uninucleate basidiospore. This fungus is found on many
species of mosses in Europe and America. (Fig. 144 A.)

Jola is also parasitic upon mosses but mostly in the sporophytes. It
seems to be almost exclusively tropical. It forms its small spherical or
elongated spore fruit at the apex of the sporophyte. It appears to be more
or less gelatinous. The binucleate hyphal cells at the surface enlarge at the
ends and in these terminal cells (hypobasidia) the nuclei unite. From the
apex of each emerges the epibasidium within which the meiotic division
of the nucleus takes place. After the three transverse septa have been
formed short or long tube-like hyphal growths reach the surface of the
hymenium and bear the uninucleate spores at their tips. Below the ter-
minal basidium the next cell grows out sympodially and produces another
basidium and this process is repeated until a dozen or more basidia are
produced (see Gaumann, 1922). Closely related, but growing on fungi
instead of mosses is the likewise tropical genus Cystobasidium. (Fig.
144 B.)

Herpobasidium filicimim (Rostr.) Lind was studied by Jackson (1935).
It is parasitic upon the leaves of ferns in which the mycelium is inter-
cellular, producing massive coiled haustoria in the cells of the host. The
internal mycelium emerges through the stomata, forming small white
patches. On the surface the basidia arise as terminal cells of upright hy-
phae. They are usually slightly bent. No distinction of hypobasidium and
epibasidium is observable. The fusion nucleus divides once, by the first step
of meiosis, so that the two nuclei now have the haploid number of chromo-
somes. A single septum is formed and one basidiospore develops on the
tip of a sterigma on each cell. The spore is uninucleate and no nuclear
division has been observed. It may germinate by repetition. The origin of
the dicaryon phase is unknown as is also the case in the two foregoing
genera. No clamp connections have been observed. This species occurs in
Europe and North America.

Another genus, Platycarpa (Couch, 1949), is parasitic upon tropical
ferns. The fruit body is resupinate, very small, dry to subcartilaginous,
separable from the host at maturity. The vegetative mycelium forms
coiled haustoria within the epidermal and mesophyll cells or in the sporog-
enous cells of the host. No clamp connections have been observed. Exter-
nally a more or less felty mass of hyphae occurs. These are more or less
wavy or loosely coiled near the surface and produce terminally on short
branches the ovoid probasidia. These are hyaline-walled and germinate
by the production of a straight or, more often, curved and mostly four-

Fig. 144. Order Auriciilariales, Family Auriculariaceae. (A) Eocronaiiuim. muscicola
(Ft.) Fitzp. Basidium with tubular extensions terminated by sterigmata bearing spores.
(B) Jola javensis Pat., basidiophoric hypha showing sympodial mode of growth, with
almost mature basidium at the tip. (C) Helicohasidiuin cotnpadtnn Boedijn, section
through the hymenium. (D, E) Helicogloea lagerheimi Pat. {Saccohlastia) . (D) Fructi-
fication on wood showhig basidia in various stages of development. (E) Mature basid-
ium shedding spores, the hypobasidium collapsed. (A, courtesy, Fitzpatrick: Phyto-
pathology, 8(5):197-218. B, after Gaumann: Ann. Mycolog., 20(5-6) :272-289. C, after
Boedijn and Steinmann: Bull. Jardin Botan. Buitenzorg, Serie III, 11(2) : 165-219. D-E,
courtesy, Baker: Ann. Missouri Botan. Garden, 23(1):69-128.)

442

ORDER AURICULARIALES 443

celled epibasidium. The somewhat allantoid basidiospores are produced
on distinct sterigmata. In structure of the sporiferous portion of the fruit
body the fungi of this genus show considerable similarity to Septohasid-
ium, to which Couch suggests that they may be transitional forms.

Helicobasidium is parasitic on the roots or crowns of trees and shrubs
or perennial herbaceous plants, and also may grow as a saprophyte. H.
purpureum (Tul.) Pat. has been studied by Buddin and Wakefield (1927).
On the surface of the substratum it forms a nongelatinized, effused, felted
layer. The basidial primordia are often more or less coiled. They become
hypobasidia from whose apex grows out the terminal hooked epibasidium
which produces sometimes rather long sterigmata. In H. candidum
Martin (1940) the hypobasidium is small, forming practically only a short
slender stalk for the stout curved basidium. Clamp connections are pres-
ent. Syzygospora, with a single species S. alba was described by Martin
(1937a). It forms a white gelatinous spore fruit up to 10 cm. long and 2.5
cm. thick, the surface covered by the hymenium. The basidia occur in
sympodial fascicles. They have but a single septum with a basal cell more
or less elongated and clavate and the terminal cell spherical. A single
spherical basidiospore is produced from each cell, near the base of the
apical cell of the basidium and near the top of the basal cell. Thus the
spores come into contact and unite to form an ellipsoid spore. Martin
compares this with the union of two sporidia while still attached to the
promycelium of some smuts. The hyphae are abundantly provided with
clamp connections. (Fig. 144 C.)

Platygloea is waxy or gelatinous, resupinate, and effused, mostly grow-
ing on wood. Usually there is no distinction into hypobasidium and epi-
basidium. Helicogloea (Saccohlastia) has many of the same characteristics
but is more gelatinous. It also grows on dead wood. A distinct, usually
lateral and hanging pyriform hypobasidium is conspicuous. The basidial
primordia are usually terminal but may be intercalary. (Fig. 144 D, E.)

Auricularia forms the largest and most conspicuous spore fruits of this
family. They are borne on branches or trunks of trees and are especially
abundant in the Tropics. The common species of the Temperate Zones is
A. auricularis (S. F. Gray) Martin {A. auricula- judae or Hirneola auricula-
judae of most authors). The spore fruits are gelatinous and more or less
ear-shaped when moist, and of a translucent brown color, but dry down to
small dark horny masses. The basidia form a hymenium on the lower
surface, standing parallel in a sort of palisade immersed in the gelatinous
matrix. The elongated basidial primordia become divided by three trans-
verse septa and from each of the cells grows a tube-like extension to the
surface of the matrix where a sterigma is formed bearing a single basidio-
spore. The tropical genus Tjibodasia is waxy, and more or less pezizoid in

444

CLASS BASIDIOMTCETEAE

Fig. 145. Order Auriculariales, Family Auriculariaceae. Auricularia auricularis (S.
F. Gray) Martin. (A) Expanded, moist spore fruit. (B) Stages in the development of the
basidium. (C) Tubular extensions from basidium, each with its sterigma and basid-
iospore. (D) Stages in germination of basidiospore. (A, after Buller: Researches on
Fungi, vol. 2, p. 162, London, Longmans, Green and Co. B-D, after Brefeld: Unter-
suchungen aus dem Gesammtgebiete der Mykologie, Heft 7, pp. 1-178.)

appearance. No sharp distinctions into hypobasidium and epibasidium are
apparent. (Fig. 145.)

Family Phleogenaceae. In this family the fleshy or gelatinous
spore fruit is a stalked structure of ascending hyphae. At the top the
hyphae flare outward to form a head. The outer hyphae form a sort of
loose peridium. Within this the straight or curved basidia arise. They are
transversely one to three septate and the basidiospores are practically
sessile. There is no distinction between the hypobasidium and epibasid-
ium. Clamp connections are abundant. The only common genus in the
temperate regions is Phleogena (Pilacre of some authors), with a single
species P. decorticata (Schw.) Mart. (P.faginea (Fr.) Link). This grows on
dead stumps, logs, etc., and forms colonies of stalked structures 5 to 7 mm.
tall, including the head which is 1 to 3 mm. in diameter. When young they
are fleshy but at maturity dry. The spores are yellow-brown. Superficially
they resemble the Ascomycetous genus Onygena. Shear and Dodge (1925)
described the life history and cytology of this species very fully. Other
genera more or less probably belonging in this family are Pilacrella, fleshy,
with a disk-shaped head and hyaline spores; Hoehnelomyces, tropical,
slimy cartilaginous, or even waxy, with hyaline spores and with a round
head with loose wavy hairs; and perhaps Stilhum, in which the stalk and
head are fleshy but the surrounding peridial hyphae are lacking. The ba-

ORDER AURICULARIALES

445

Fig. 146. Order Auriculariales, Family Phleogenaceae. Phleogena decorticata
(Schw.) Mart. (A) Habit study of the fungus. (B) Basidiophoric hypha with apex
extended to form part of the so-called peridium. (C-H) Stages in the development of
the basidium and basidiospores. (A-B, after Brefeld: Untersuchungen aus dem
Gesammtgebiete der Mykologie, Heft 7, pp. 1-178. C-H, courtesy, Shear and Dodge: J.
Agr. Research, 30(5):407-417.)

sidia are two-celled and the spores hyaline. There is one species in Europe
and North America and three others in the tropics. There is uncertainty
as to whether the genus really belongs here. (Fig. 146.)

Family Septobasidiaceae. This is considered by Couch (1937 and
1938) to be worthy of ordinal rank. The two genera included in the family
are parasitic upon scale insects, with which they live in a sort of symbiotic
relationship much as the lichen fungus does with the imprisoned algae.
For the majority of these insects the fungus provides a home and shelter
where they feed upon the woody host plant and produce their young.
Some of the insects, however, are parasitized and continue to feed upon
the host plant but give up their food to the fungus which penetrates their
bodies with numerous coiled or knotted haustoria. These insects may out-

446

CLASS BASIDIOMYCETEAE

Fig. 147. Order Auriculariales, Family Septobasidiaceae. Septobasidium hurtii
Lloyd. (A) Vertical section through a portion of the fungus showing a parasitized living
scale insect {Aspidiotus) and the hymenium on the upper surface. {In upper portion of
figure) (h, ha) Epibasidium; {ph) hypobasidium ; {Sp) spore; {ys) young scale insect;
{Fs) fruiting surface of fungus; {tl) top layer of fungus; {hi) bottom layer of fimgus;
{arrow) tunnel in fungus. {Referring to host plant) (6) Bark; (c) cambium; {nc) medul-
lary ray. {Referring to parasitized insect and sxirrounding fungus) {Fm) Enveloping
fungus mat; {ct) fungus thread connectmg fungus mat with insect; (C) coiled hausto-
rium within insect; {ss) spindle-shaped threads within insect; {St) stylet; {sh) sheath
secreted around stylet. {01, Stg, Ph, rp, gl, ov) Various organs of the insect.

ORDER AURICULARIALES

447

Fig. 14:7— (Continued). (B) Section of the hymenium showing hypobasidia and
fully developed basidium with basidiospores. (Courtesy, Couch: The Genus Septo-
basidium, Chapel Hill, Univ. North Carolina Press.)

448 CLASS BASIDIOMYCETEAE

live the nonparasitized ones, but never reproduce. The newly hatched
young from the unparasitized insects, in creeping around to find a place to
settle down come in contact with the germinating basidiospores and the
yeast-like buds from the latter adhere to the body or appendages of the
insects. Many of the young escape infection so that there are always
enough left to reproduce the species while the others feed the fungus which
provides the sheltering homes for them. The chief genus is Septobasidium,
with over 160 species and subspecies. They are reported from both the
Old World and the New World, especially in tropical and subtropical
regions, but extend up into the warmer temperate regions. One species
grows in Canada. None are known in the colder parts of Europe and Asia.
Vast areas of the world are still only slightly studied with this genus in
view. The typical basidium consists of a well-developed, rather firm-
walled hypobasidium from which grows out the straight or curved epi-
basidium which becomes four-celled. In some species new basidial
primordia proliferate in the empty hypobasidia. The basidiospores are
borne on well-developed sterigmata. They usually become septate after
discharge and then apparently germinate only by yeast-like buds and not
by hyphae. In many species the hypobasidium is dark-colored and serves
as a resting spore until favorable conditions arise, in others the wall is
hyaline and the epibasidium develops immediately. Boedijn and Stein-
mann (1931) were the first to report that in some species of the genus no
distinction of hypobasidium and epibasidium can be observed. In many
species the basidium is one-celled, two-celled, or three-celled, with or
without a distinct hypobasidium. The mycelium lacks clamp connections,
both on the hyphae external to or within the bodies of the insect hosts. In
the latter the hyphae are a series of slender spindle-shaped cells with or
without thick, coiled or clumped haustoria. The fungus forms a two-, or
more, storied structure with chambers in which the insects live, and tun-
nels. In some species definite pillars support the successive layers of the
fungus. Growth may cease in unfavorable times and become renewed
when favorable weather recurs. The presence of a scale insect often in-
duces the fungus to form a house or tent above it, but often only a low
vaulted cavity is developed over the insect. The tunnels and openings to
the surface provide means of egress for the young insects. The upper layer
consists of more or less vertically growing loose hyphae, usually branched
at intervals, and bearing the basidia at the surface. In a number of species
conidia have been observed, usually produced on the floor of the tunnels
or chambers or on short branches from the ascending hyphae of the upper
layer. (Fig. 147.)

The Septobasidium colony may be but a few millimeters in diameter or
up to 20 or 30 cm. or more. It may be a fraction of a millimeter in thick-
ness or up to 1 cm., depending upon the species. It often resembles a lichen

ORDER DACRYMYCETALES 449

and the color varies from very light gray to dark brown, purple or even
almost black. The infected trees or branches are injured, perhaps not by
direct action of the fungus itself but by the protection it offers to the
many scale insects it harbors.

The genus Uredinella is perhaps best placed in this family. Like Septo-
hasidium it is parasitic upon scale insects and forms small circular dark-
colored spots on the bark. The fungus is annual, not perennial as is
Septohasidmm. The top layer is a hymenium of ovoid to club-shaped,
brown basidial primordia (called " teleutospores " by Couch, 1937). These
are binucleate when young and nuclear fusion occurs within them. The
mature cells have two or three thick layers of wall especially at the apex,
and a distinct germ pore. When the fungus is wet with rain an epibasidium
emerges through the germ pore and forms a straight four-celled structure
on which four basidiospores are produced on distinct sterigmata. The
epibasidium may break off from the hypobasidium and apparently can
float around in a film of rain water. The insects are infected through the
mouth region and the dicaryon mycelium within the host produces coiled
haustoria much like those of Septohasidium. Besides the basidia somewhat
similar "uredo-mother cells" are produced and from their apices grow out
binucleate, elongated ellipsoid, slightly bent spores, called by Couch
"uredospores." It is suggested by Couch that this genus may well repre-
sent a stage intermediate between the Uredinales and Septohasidium.

Order Dacrymycetales. As in the preceding order the spore fruits are
mostly gelatinous or waxy, drying down to a thin sheet or horny mass.
They vary from thin, broadly effused sheets to cushion-like, cupulate or
pileate structures, or cornute, coralloid or spatulate upright forms. No
species are known to be parasitic. They are almost exclusively confined to
dead wood, with or without bark. With very few exceptions the spore
fruits are colored some shade of yellow or orange to deep brown and the
basidiospores are mostly yellowish in mass. Conidia are frequently pro-
duced. Whether these correspond to the "oidia" of some of the other
orders is uncertain, for cultural studies and attempts to match and dip-
loidize different strains are as yet much to be desired. The basidiospores
are one-celled when ready to be discharged but in most species septa are
formed immediately after discharge, di■\^^ding the spore into 2, 4, or even
up to 12 cells. In a specimen of Dacrymyces studied by the author the
basidiospores became once septate before they were discharged from the
sterigmata. They germinate by the formation of hyphae or of yeast-like
buds from the various cells of the spore.

The basidia are formed in a close hymenium or intermingled with
sterile hyphae, on the outer surface of resupinate forms or on definite
surfaces in various other forms. They start as binucleate terminal cells
soon thicker than the rest of the hypha. They are at first long cylindrical

450

CLASS BASIDIOMYCETEAE

or somewhat clavate. The two nuclei unite and then divide once sticho-
basidially, and then a second time in the same direction. The broadened
apex of the basidium becomes lobed to produce two "epibasidia" of the
same diameter and often of almost the same length as the "hypobasid-
ium." The upper two nuclei migrate into the two epibasidia and then
through the sterigma at the tip of each into the basidiospores. The two
nuclei remaining in the hypobasidium degenerate. The mature basidium
often resembles a tuning fork. Clamp connections are frequent in the spore
fruits but in the primary mycelium produced by the germination of the
spores they are lacking. Just how and when diploidization occurs has not
been demonstrated. Eight to ten genera are recognized, in temperate as
well as in tropical regions.

Fig. 148. Order Dacrymycetales, Family Dacrymycetaceae. (A) Daciymyces
lulescens Bref., habit sketch. (B) Dacrymyces deliquescens Duby, various stages in the
development of basidia and basidiospores. (C) Guepiniopsis sp., habit sketch. (D)
Basidiospores of Guepiniopsis. (A, after Brcfeld: Untersuchungen aus dcm Gesammt-
gcbiete der Mykologie, Heft 7, pp. 1-178. B, after Dangeard: Le Botaniste, 4:119-
181. C-D, courtesy, Martin: Mycologia, 24(2):215-220.)

ORDER TREMELLALES 451

Among the effused forms is Cerinomyces Martin, 1949 {Ceracea of
authors), forming a thin waxy to fleshy layer, without definite mycelial
roots. The basidiospores may remain nonseptate or in some species may
become transversely septate. The genus Arrhytidia forms tough waxy, at
first discoid, then broadly effused spore fruits with a centrally rooting
base. The spore fruits of Dacrymyces are sessile, attached by a point or
rhizoids, or substipitate. They are gelatinous to waxy, pulvinate, discoid,
or cerebriform, or even cupulate. The entire exposed area is covered by
the hymenium which may become wrinkled or folded when older. Martin
(1944) recognizes seven species in the United States and Canada. Guepi-
niopsis {Heterotextus of earher works) produces substipitate, cupulate spore
fruits, with the cup often turned downward so that the hymenium is in-
ferior. The cortex consists of swollen, thick-walled cells. The interior
hyphae are strongly gelatinized. Femsjonia is also discoid or cupulate,
sometimes substipitate, but is white-villous or tomentose externally.
Dacryopinax (Martin, 1948, a name substituted for the more familiar but
preoccupied name Guepinia), also may be cupulate and erect when young,
becoming spatulate, fan-shaped or petaloid, with hymenium on the lower
side. The spore fruits are definitely stipitate and tough or cartilaginous.
Dacryomitra also is stipitate, with a distinct pileus more or less morchel-
loid in appearance and gelatinous. The hymenium covers all sides of the
head. Calocera is cornute to coralloid, with hymenium on all sides. It
resembles Clavaria but differs in the basidial structure — gelatinous to
tough. (Fig. 148.)

Order Tremellales. Spore fruits varying from adhering, waxy or
gelatinous sheets to fohose or cushion-like or pileate structures, sometimes
upright and branched, forming more or less leaf-like lobes, or funnel-
shaped. In the genus Hyaloria the spore fruit is soft with a rounded head.
The consistency of the fruit body in this order may be very soft-gelatinous
to almost leathery or waxy and may become horny when dry. The colors
range from white to yellow, brown or almost black.

The basidial primordium is terminal and rounded but in Sirobasidium
the basidia are produced basipetally from the apex in the same hypha.
The young basidium is binucleate. After the fusion of the nuclei in the
typical cases the diploid nucleus undergoes its two meiotic divisions at
right angles to the axis of the cell (chiastobasidially) and then a vertical
septum is formed, followed almost immediately by another vertical sep-
tum at right angles to the first in each of the two cells. At the top of each
of the four cells so formed a sterigma may be produced bearing one ba-
sidiospore. More often, since the majority of the species are gelatinous
with the basidia embedded a short distance below the surface, a tube-like
extension of the basidial cell grows upward to the surface, there producing
the sterigma and basidiospore. These tubular outgrowths are homologous

452

CLASS BASIDIOMYCETEAE

Fig. 149. Order Tremellales, Family Tremellaceae; (A) Tremella reticulata (Berk.)
Farl. (B) Tremella mesenterica (S. F. Gray) Pers. Stages in the development of basidia
and basidiospores. (A, courtesy, Atkinson: Studies of American Fungi, Ithaca, N. Y.,
Andrus and Church. B, after Dangeard: Le Botaniste, 4:119-181.)

ORDER TREMELLALES 453

to the similar structures growing out of the cells of the basidia of Auricu-
laria and cannot rightly be called epibasidia, for the epibasidia of Auricu-
lariales and the promycelium of Uredinales are probably homologous, but
are entirely different from these tubular extensions to bring the spore pro-
duction to the surface. The basidiospores are hyaline or only slightly
colored. They are without septa when set free. They may then become
once septate, but more often not. Frequently they produce a sterigma and
a secondary spore, a process several times repeated. In abundance of
moisture the spores bud out innumerable "oidia" on their surface. They
may, if on a proper substratum, germinate by hyphae. The primary
mycelium produces an abundance of oidia and lacks clamp connections.
Eventually clamp connections appear (secondary mycelium) and oidial
production usually ceases. Barnett (1937) showed for several species of
Exidia that the spores from a given spore fruit exist in two sexual phases.
Multiple allelomorphy of the compatibility factors was demonstrated
also, as occurs in Auricularia and in various Ustilaginales.

Over 100 species in 17 or more genera are recognized in this order, of
which about half are found in the North Central United States and
Canada (Martin, 1944).

Three families are usually distinguished:

Family Tremellaceae. Basidia are single and terminal on the sup-
porting hyphae, with a more or less elongated extension from each cell,
terminated by a sterigma. They are gelatinous, waxy or somewhat dry.
Basidia normally are cruciately four-celled but sometimes three-celled
or even two-celled. The primary septum is vertical or oblique. Hymenial
surface is exposed. Spore fruits may be flat, cushion-shaped, lobed, or
pileate. Clamp connections present in secondary mycelium of many
species. (Fig. 149.)

Family Sirobasidiaceae. Gelatinous, cushion-shaped, hymenium ex-
ternal. Basidia are formed in chains by successive transformation of the
cells of the basidiogenous hyphae into basidia, beginning at the apex.
Basidia are four- or two-celled, the septum in the latter case oblique. Ba-
sidiospores are sessile and therefore probably not discharged violently.
Mostly tropical but known in the United States from North Carolina.
Contains a single genus, Sirobasidium. (Fig. 150.)

Family Hyaloriaceae. Spore fruits are stalked, with a head; stalk
and head somewhat gelatinous externally or sessile, forming a filmy layer
on the substratum. Basidia are two- to four-celled with tubular extensions
which may taper to a long fine thread, the basidiospore being borne sym-
metrically at the apex and breaking off with part of the supporting thread
attached, not discharged from a sterigma. Clamp connections are present.
There are two genera : Hyaloria, with one tropical species, H. pilacre A.
Moll, (see Martin, 1937b), and one European one, H. europaea Killer-

454

CLASS BASIDIOMYCETEAE

Fig. 150 {Left). Order Tremellales, Family Sirobasidiaceae. Sirobasidium albidum
Lagerh. & Pat. (A) Chain of two basidia, the terminal one almost mature. (B) Chain
of several basidia, the upper three collapsed, the fourth approaching maturity. (After
Lagerheim and Patouillard: /. Botan., 6(24):465-469.)

Fig. 151 (Right). Order Tremellales, Family Tremellaceae. Pro/o/iz/d/ium geZaiznosw?«
(Fr.) Karst. {Tremelodon of most authors). Habit sketch. (After A. MoUer, from
Killermann, in Engler und Prantl: Die NatiirUchen Pflanzenfamilien, Zweite Auflage,
vol. 6, pp. 99-290, Leipzig, W. Engelmann.)

mann (1936) and Xenolachne, growing as a parasite on a minute Dis-
comycete in Oregon (Rogers, 1947). Hyaloria is stalked and externally
gelatinous with the basidia on the head, much overtopped by long hairs
(or cystidia). Xenolachne forms a thin film on the apothecium of the host
and lacks a gelatinous coat or cystidia but the extremely long extensions
of the two-celled basidia, with the basidiospores at the apex give a felty
appearance to the fungus. The type of basidium in these two genera
resembles that characteristic of most of the Gasteromycetes and for this
structure Rogers adopts the name apobasidium proposed by Gilbert

(1928).

In the Tremellaceae the genus Stypella produces a small felty mass of
tangled, more or less gelatinous hyphae in whose upper layer the basidia
arise. These spore fruits are clustered, separate or anastomosing on a dry
fioccose subiculum, the whole patch sometimes reaching a diameter of
several centimeters. In Sehacina there is a more or less waxy or gelatinous
crust with the hyphae of the upper portion directed perpendicularly to
the surface, the terminal cells of these rather closely packed hyphae
being the basidia. Cushion-like or lobed, gelatinous spore fruits are char-
acteristic of Exidia and TremeUa. Gland-like dots occur in the spore fruit
of the former and are absent in the latter. The basidiospores of the former
are mostly allantoid, those of the latter straight and ellipsoidal to nearly
spherical. When these spores germinate the sprout-conidia of Exidia are
mostly curved, those of TremeUa yeast-like.

ORDER TULASNELLALES 455

Phlogiotis (Gyrocephalus) has funnel-shaped spore fruits with the
hymenium on the outer surface only. Tremella is found in all parts of the
world. T. reticulata (Berk.) Farl. forms large white masses of gelatinous
leaf-like lobes, the whole mass sometimes being 10 to 12 cm. in diameter.
It is considered edible. Other species are usually smaller and some are
bright-colored. Other genera less common in the temperate zones or con-
fined to the tropical or subtropical regions are among those described
below. Patouillardina has the basidia spindle-shaped. The first septum
is oblique and in each cell thus formed another septum is produced at
right angles to the first one. Because of the shape of the basidium these
two septa do not intersect the first septum opposite one another. In
Protomerulius the soft fleshy or waxy spore fruit is resupinate and rather
thin. Its hymenium is poroid. Protodontia and Protohydnum are waxy or
gelatinous and resupinate or stalked, but the hymenium instead of being
poroid is borne on downward directed teeth. The earlier name Proto-
hydnum must, according to Martin (1948), be used instead of the more
familiar but later Tremellodon. In Protohydnum the stalk of the basidium
is separated from it by a septum but not so in Protodontia. Heterochaete
is somewhat similar, but the blunt teeth are peg-like and pierce the
hymenium, not being covered with basidia themselves. Tremellodendron
is erect, more often branched, resembling Clavaria or some species of
Tremella. Eichleriella {Hirneolina) is cupulate or broadly attached. (Fig.
151.)

The Tremellales must be considered as a group which has developed
with more or less parallelism to the Auriculariales. The low, felty or
gelatinous waxy crust, bearing basidia on the upper surface, seems to be
the most primitive in each order, and from this simple structure have de-
veloped the more complex forms of spore fruit. It must be emphasized
again that until the life histories have been more fully worked out the true
relationships are only a matter of conjecture. The rather frequent occur-
rence of conidial production in these orders would hint at relationship to
the Ascomyceteae in which conidia are produced abundantly.

Order Tulasnellales. The fungi making up this order are mostly found
on dead wood or on old fungi on which they are saprophytic. The spore
fruits are resupinate, gelatinous or dry, usually thin, sometimes being
only a slight film-like coating on the substratum. Clamp connections are
found on the hyphae of most species but are lacking in some. The char-
acteristic feature of the single family, Tulasnellaceae, is the structure and
developinent of the basidia. These are typical in the genera Tulasnella
and Gloeotulasnella. The genus Ceratohasidium (Rogers, 1935) was tenta-
tively placed by Martin (1944) in this family but later (1948) segregated
by him in a distinct family Ceratobasidiaceae, assigned to a position
close to the Tulasnellaceae. The basidium in all three genera is a holo-

456

CLASS BASIDIOMYCETEAE

basidium, i.e., is not divided by vertical septa as in the Tremellales nor by
transverse septa as in the Auriculariales. The basidial primordium is sub-
globose, pyriform or broadly clavate. From the upper portion of this cell
arise usually four stout cells which are narrowed at the tip to form a
sterigma upon which a single basidiospore is formed. These four cells are
considered by Rogers, Martin, and others to be homologous to the stout
arms growing out of the four cells of the basidium of the Tremellaceae,
and which they call "epibasidia."

In TulasneUa and Gloeotulasnella the "epibasidia" are separated from
the "hypobasidium" by a septum at the base of each. They may remain
attached or may fall off. At the apical end they elongate and bear a
sterigma on which is borne the nonseptate basidiospore. This spore when
set free germinates "by repetition," i.e., it produces a lateral or terminal
sterigma and a secondary spore into which the whole cytoplasm and
nucleus pass, this spore then being discharged as in the case of the parent
spore. These two genera are distinguished as follows: TulasneUa, "arid-
pruinose to waxy, basidia not imbedded in a gelatinous matrix; pro-
basidia globose to obovate, sessile or with a short scarcely differentiated
stalk; epibasidia with subulate tips merging into the sterigmata; gloeocys-
tidia never present" (Martin, 1944). A dozen or more species. Gloeotulas-
nella, "waxy gelatinous to mucous gelatinous, basidia imbedded in a
gelatinous matrix; probasidia clavate capitate, with a more or less
cylindrical stalk and a swollen head; epibasidia extended into cylindrical
tubular filaments sharply constricted at the base of the sterigmata;
gloeocystidia present or absent" (Martin). Ten or more species. The

Fig. 152. Order Tulasnellales, Family Tulasnellaceae. (A, B) TulasneUa violea
(Quel.) Bourd. & Galz. {PachysterigmafugaxSoh^in-Ol&en). (A) Mycelium with several
clusters of basidia. (B) Mature basidium. (C) Ceratobasidium cornigerum (Bourd.)
Rogers. Basidia and basidiospores in various stages of development. (A-B, after
Brefeld: Untersuchungen aus dem Gesammtgebiete der Mykologie, Heft 8, pp. 1-305.
C, courtesy, Martin: Univ. Iowa Studies in Natural History, 18(3): 1-88.)

SUMMARY OF SUBCLASSES TELIOSPOREAE AND HETEROBASIDIAE 457

cytology and taxonomy of these genera were studied by Rogers (1932,
1933). (Fig. 152 A, B.)

From the two foregoing genera Ceratobasidium is distinguished by the
stout "epibasidia," elongate cornute or flexuous, continuous with the
"hypobasidium" (rarely a cross septum). The spore fruit is arid or waxy.
Six or more species. By the usual absence of septa cutting off the "epi-
basidia" and their cornute shape they approach on the one hand Dac-
rymyces [in C. sterigmaticum (Bourd.) Rogers, in which only two such
horns are produced, while the remaining species with their four "epi-
basidia" approach Pellicularia in the Thelephoraceae (Eubasidiae).
From the latter they differ by the germination by repetition, of the
basidiospores. (Fig. 152 C.)

Summary of Subclasses Teliosporeae and Heterobasidiae

A comparison of these subclasses demonstrates more or less basic
similarities throughout, in basidium production. Between the Auri-
culariales, Uredinales, and Ustilaginales there are such basidial similari-
ties that they are often placed together in one group. On the other hand
the tendency for the septation of the basidium in the Tremellales to be
oblique instead of vertical is taken by some mycologists who have studied
these groups intensively to indicate gradation from one to the other.
Rogers (1934) suggested that the septa at the base of the "epibasidia" of
Tidasnella may be accounted for by displacement upward of the vertical
septa of the basidium of Tremella. The peculiar tuning-fork type of
basidium of Dacrymyces could be considered a derivation from the
Tidasnella type by the loss of their cross septa entirely and the reduction
of the "epibasidia" to two (as actually does occur in Ceratobasidium
sterigmaticum) .

The question of the phylogeny of these groups is treated more fully
in Chapter 17, but the following suggestions may well be made here:

Studies by Juel (1898), Neuhoff (1924), Martin (1931), and Rogers
(1934) have led the latter to an interpretation of the relationships in the
class somewhat different from that of the author. He holds in common
with some of the others mentioned, that the primitive basidium con-
sisted of two parts, the basal hypobasidium, binucleate at first, within
which the nuclear union occurs, and one or more outgrowths, the epi-
basidia. Into the latter the nuclei pass from the hypobasidium. Meiotic
division may occur either in the latter or in the epibasidium, if there be
but one. Each epibasidium produces a true sterigma which bears a
basidiospore. The genus Tulasnella is considered by Rogers to represent a
primitive form. In the ovoid or pyriform hypobasidium of this genus the
fusion nucleus divides into usually four nuclei. One of these passes out
into each of the four (sometimes fewer) stout epibasidia which usually
become separated from the now almost empty hypobasidium by a basal

458 CLASS BASIDIOMYCETEAE

septum. Each epibasidium produces a terminal sterigma and basidio-
spore. The nucleus may divide within the epibasidium and both nuclei
pass into the basidiospore. By producing but two epibasidia,not sepa-
rated by septa from the hypobasidium the typical tuning-fork basidium
of the Dacrymycetales can be evolved. By crowding the basal septa
down into the hypobasidium so as to divide that longitudinally into four
cells the basidium of the Tremellales may be derived. By reduction of the
size of the epibasidia until only the sterigmata are left, is developed the
basidium characteristic of the Hymenomycetes. Rogers attempts to ex-
plain the derivation of the hypobasidium and single, transversely septate
epibasidium of Septobasidivm and of the corresponding teliospore and
promycelium of the Uredinales and Ustilaginales as being due to the
delay in the meiotic division of the fusion nucleus until it passes out into
an epibasidium which naturally would be single for a single nucleus. He
believes that the hypobasidium of Tulasnella represents an ascus, perhaps
of some form resembling Ascocorticium (Order Taphrinales), in which the
ascospores have pushed out into pockets, germinating there to form
secondary spores. These pockets have become the epibasidia and the
secondary spores have become the basidiospores. Martin (1938) discusses
the morphology of the basidium in connection with Heterobasidiae and
Eubasidiae.

Linder (1940) would, on the other hand, derive the Uredinales from
the Ascomyceteae, in the vicinity of the Sphaeriales or Dothideales.
From the Uredinales he would derive the Auriculariales, Tremellales and
Dacrymycetales. The Corticiae (Family Thelephoraceae), and thence the
other Eubasidial families he would derive from the Tremellales.

On the other hand some students of these fungi consider the primitive
basidium to have been of the holobasidium type from which the forked
and septate types have been derived. The distinction of hypobasidium
and epibasidium are, in this viewpoint, specializations to meet the need
of holding the basidium over until a more favorable period. The thick-
walled hypobasidium (or "probasidium") such as is found in the Telio-
sporeae or in Septohasidium cannot, because of the thickened wall,
develop in the manner normal to basidia, and so a thin-walled new struc-
ture, the epibasidium or promycelium, was developed. The thickened
apical branches of the basidia of the Tremellales and Dacrymycetales and
of Tulasnella are not considered to be epibasidia but merely modified
sterigmata.

Key to the Families and More Important Genera of Order Auriculariales

(Modified from Martin, 1944)

Parasitic on plants or saprophytic on dead plant material, usually wood. Basidia
with distinct sterigmata. Division into hypobasidium and epibasidium

KEY TO THE FAMILIES AND GENERA OF ORDER AURICULARIALES 459

present or absent. Basidia forming a more or less distinct exposed, loose or
compact hymenium. Family Auriculariaceae

Parasitic on the gametophytes of mosses at whose apex a gelatinous, club-
shaped spore fruit is produced. Hypobasidium short, not enlarged, very
soon collapsing, and not conspicuous. No clamp connections.

Eocronartium
Parasitic on the sporophytes of mosses forming a felty or gelatinous, more or
less spherical spore fruit. Hypobasidium and epibasidium distinct. No
clamp connections. Tropical. lola

Parasitic on the leaves of ferns; effused; haustoria coiled; no distinct difference
of hypobasidium and epibasidium. Conidial stage sometimes present. No
clamp connections observed. Herpobasidium

Parasitic on the leaves of ferns. Definite coiled haustoria formed. No clamp
connections present. Definite ovoid hypobasidia give rise to distinct, usu-
ally four-celled epibasidia. Platrjcarpa
Parasitic on the roots or crowns of vascular plants or saprophytic on dead
plant tissues. Dry, floccose, effused. Hypobasidia cylindrical or not
obvious, epibasidia curved at the top. Helicobasidium
Saprophytic on plant tissues.

Resupinate; gelatinous, forming a large cushion; basidia two-celled, the
basal cell clavate; the terminal one spherical; the two spores uniting to
form an ellipsoid spore. Numerous clamp connections. Tropical.

Syzygospora
Resupinate; soft gelatinous, hypobasidia in the form of lateral, reflexed sacks.
Clamp connections sometimes present. Helicogloea

(Saccoblastia)
Resupinate; firmly gelatinous or waxy, basidia fusiform, no apparent hypo-
basidia; clamp connections sometimes present. Platygloea
Pileate or ear-shaped, tough gelatinous; clamp connections obvious; no dis-
tinction into hypobasidium and epibasidium. Auricularia
Saprophytic on wood, bark, etc. No distinction into hypobasidium and epi-
basidium. Spore fruit stalked with a head of radiating, more or less coiled
hyphae among which the curved basidia are found, bearing their two to
four spores without visible sterigmata. Clamp connections present in some
species. Family Phleogenaceae^
Stalked with the hyphae flaring at the top to form a head, the outer ends form-
ing a loose peridium-like structure. Basidia not borne on sterigmata.
Forming colonies of stalked spore fruits on dead wood, fleshy when young,
then becoming dry. Phleogena

(Pilacre)
Parasitic upon scale insects, with some of which they live in symbiotic relation.
Basidia usually with well-developed thin-walled or thick-walled hypo-
basidia and one- to four-celled (mostly the latter) epibasidia; but hypo-
basidium sometimes lacking. Basidiospores produced on distinct sterig-
mata. Conidia often produced. Clamp connections not observed.

Family Septobasidiaceae

1 Other genera sometimes assigned to this family are Pilacrella, fleshy, with disk-
shaped head; Hoehnelomyces, slimy-cartilaginous or waxy, with round head with
loose wavy hairs; and perhaps Stilbum with fleshy stalk and head but no surrounding
peridial hyphae.

460 CLASS BASIDIOMYCETEAE

Fungus perennial, forming a two- or three-storied structure with chambers and
tunnels within which the scale insects live. Basidia formed on the outside,
more or less felty, layer. Often lichen-like. Septobasidium

Fungus annual, not forming distinct "houses" for the parasitized scale insects.
Hypobasidia thick-walled with an apical germ-pore through which a
hypha grows to form a straight, four-celled epibasidium with definite
sterigmata. In addition conidia are produced. Uredinella

Key to the More Important Genera of Family Dacrymycetaceae

(Based on Martin, 1944)

Fructifications broadly effused.

Broadly effused from the first, without root-like bases; arid to waxy-gelatinous.

Cerinomyces
(Ceracea of authors)
At first discoid or pustulate, soon becoming effused, attached to radicating
bases, tough waxy or waxy-gelatinous. Arrhxjtidia

Fructifications remaining distinct even when anastomosis occurs.
Sessile and attached by a point or on a constricted root-like base.

Pulvinate or discoid or rarely pezizoid, often cerebriforra; hymenium opposite
substratum, usually inferior. Dacrymyces

Definitely pezizoid; hymenium concave, at least until very late.
Cortex concolorous; spores finally three to seven septate.

Giiepinionsis
Cortex conspicuously white-tomentose; spores tardily multiseptate.

Femsionia
Distinctly stipitate and pileate.

Cornute to coralloid, Clavaria-like; hymenium amphigenous.

Calo cera
Pileate, pileus much broader than stalk.

Tough or cartilaginous, spatulate or cupulate; hymenium unilateral,
inferior. Dacryopinax

(Guepinia of authors)
Gelatinous, pileus conical, subglobose, flattened or morchelloid; hymenium
amphigenous. Dacrijomitra

Key to the Commoner Genera of Family Tremellaceae

(Based on Martin, 1944)

Fructification of thickly clustered, more or less anastomosing papillae, borne on a

thin floccose subiculum. Stypella

Fructification continuous, at least from an early stage, frequently enlarged by
anastomosis.
Resupinate, broadly effused, with indeterminate margins, probasidia globose to
ovate to pyriform, the first septum mainly longitudinal.
Hymenium smooth or nearly so; arid or tough to waxy or gelatinous.

Sebacina
Hymenium with spines or spine-like structures.

Spines sterile, piercing the hymenium; texture coriaceous to waxy or
tough gelatinous. Heterochaete

LITEKATURE CITED

461

Spines fertile, texture soft to tough gelatinous.

Soft gelatinous; subiculum delicate; probasidia without stalk becoming

separated as a stalk cell. Protodontia

Tough gelatinous; subiculum thick; probasidia with stalk becoming
separated as a stalk cell. Protohydnum

Erumpent or pileate, or, if appearing effused, with determinate margins.
Tough or coriaceous to somewhat waxy when moist.

Cupulate to broadly attached with a free margin; aspect of Stereum.

Eichleriella

Erect, branched or rarely simple; aspect of Clavaria or Thelephora.

Tremellodendron

Gelatinous; horny when dry.
Erect-cerebriform to lobate.

Spores subglobose or ovate. Tremella

Spores allantoid; gloeocystidia lacking. Exidia

Spores allantoid; gloeocystidia present. Seismosarca

Pileate and stipitate or substipitate.

Stipitate or dimidiate; hymenium on teeth. Protohydnum

(Tremellodon of authors)
Infundibuliform; hymenium inferior, smooth or somewhat wrinkled.

Phlogiotis
(Gyrocephalus)
Key to the Genera of Family Tulasnellaceae

(Based on Martin, 1944)

"Epibasidia" at fu-st bluntly cyhndrical, at length fusiform, not separated by
septa from the "hypobasidium." Ceratobasidium

"Epibasidia" at first globose, becoming ovate, pyriform or ventricose-cylindrical,
separated by septa from the "hypobasidium."
Arid-pruinose to waxy; basidia short-stalked, not embedded in mucus; gloeo-
cystidia never present. Tulasnella
More or less gelatinous; basidia long-stalked, embedded in mucus; gloeo-
cystidia present or absent. Gloeotulasnella

Literature Cited

Atkinson, George F.: Studies of American Fungi. Mushrooms edible, poisonous,

etc., vi + 275 pp. 6 colored plates. 223 figs. Ithaca, N. Y., Andrus and

Church, 1900.
Baker, Gladys E.: A study of the genus Helicogloea, Ann. Missouri Botan.

Garden, 23(1):69-128. Pis. 7-14. 1936.
Barnett, Horace L.: Studies in the sexuality of the Heterobasidiae, Mycologia,

29(5) :626-649. Fi^s. 1-3. 1937.
BoEDiJN, K. B., ET A. Steinmann: Les especes des genres Helicobasidium et

Septobasidium des Indes N^erlandaises, Bull. Jardin Botan. Buitenzorg,

s6r. Ill, 11(2):165-219. Pis. 14-18. Figs. 1-31. 1931.
Brefeld, Oscar: Basidiomyceten II. Protobasidiomyceten, in Untersuchungen

aus dem Gesammtgebiete der Mykologie, Heft. 7, pp.i-ix, 1-17S. Pis. 1-11.

Leipzig, Arthur Felix, 1888.
: Basidiomyceten III. Autobasidiomyceten und die Begrlindung des

naturlichen Systems der Pilze, ibid., Heft. 8, pp. i-iv, 1-305. Pis. 1-12.

Leipzig, Arthur FeUx, 1889.

462 CLASS BASIDIOMYCETEAE

BuDDiN, W., AND E. M. Wakefield: Studies in Rhizoctonia crocorum (Pers.)

DC. and Helicobasidium purpureum (Tul.) Pat., Brit. Mycolog. Soc. Trans.,

12:116-140. P^s. 11-14. 1927.
Couch, John N.: A new fungus intermediate between the Rusts and Septobasid-

ium, Mycologia, 29(6):665-673. Figs. 1-30. 1937.
: The genus Septobasidium, ix + 480 pp. Frontispiece and 114 plates. 60

text figs. Chapel Hill, N. C, Univ. North Carolina Press. 1938.

The taxonomy of Septobasidium polypodii and S. album, Mycologia,

41(4):427-441. Figs. 1-25. 1949.
Dangeard, p. a.: M^moire sur la reproduction sexuelle des Basidiomycetes,

Le Botaniste, 4:119-181. Figs. 1-24. 1895.
FiTZPATRicK, Harry M. : The life history and parasitism of Eocronartium musci-

cola. Phytopathology, 8(5):197-218. PL 1. 4 text figs. 1918a.
: The cytology of Eocronartium muscicola. Am. J. Botany, 5(8):397-419.

Pis. 30-32. 1918b.
Gaumann, Ernst: tjber die Entwicklungsgeschichte von lola javensis, Ann.

Mycolog., 20(5-6) :272-289. PL 3. Figs. 1-36. 1922.
: Comparative Morphology of Fungi. Translated by Carroll William

Dodge, xiv + 701 pp. 406 figs. 43 diagrams. New York, McGraw-Hill Book

Co., 1928.
Gilbert, E.: Bribes mycologiques : VI. Conjectures sur la classification et la

filiation des especes, Bull. soc. mycologique de France, 44(3) :225-227. 1928.
Gilbert, E. N.: Cytological studies of the lower Basidiomycetes, Trans. Wis-
consin Acad. Sci., 20:387-397. PL 29. Fig. 1. 1921.
Jackson, H. S.: The nuclear cycle in Herpobasidium filicinum with a discussion

of the significance of homothallism in Basidiomycetes, Mycologia, 27(6) :553-

572. Portrait of author. Figs. 1-4. 1935.
JuEL, H. 0.: Die Kerntheilungen in den Basidien und die Phylogenie der Basidio-

myceten, Jahrb. wiss. Botan., 32:361-388. PL 4. 1898.
KiLLERMANN, S. : Unterklasse Eubasidii: Reihe Hymenomyceteae (Unterreihen

Tremellineae und Hymenomycetineae), in A. Engler und K. Prantl: Die

Naturlichen Pflanzenfamilien, Zweite Auflage, vol. 6, pp. 99-290. 5 pis. Figs.

81-157. 1928.
: Eine europaische Hyaloria Art, Ber. deut. botan. Ges., 54(2):165-167.

PL 25. 1936.
DE Lagerheim, G., et N. Patouillard: Sirobasidium, nouveau genre d'Hym^no-

mycetes h^t^robasidies, J. Botan., 6(24) :465-469. Figs. 1-2. 1892.
Leach, J. G.: Insect transmission of plant diseases, xviii + 615 pp. Frontispiece.

Figs. 1-238, New York, McGraw-Hill Book Co., 1940.
Linder, David H.: Evolution of the Basidiomycetes and its relation to the

terminology of the basidium, Mycologia, 32(4):419-447. Figs. 1-6. 1940.
Martin, G. W.: Notes on Iowa Fungi, 1929-30. The genus Tulasnella in Iowa,
Univ. Iowa Studies in Natural History, 13(5):4-10. 1931.

: On certain species of Heterotextus, Mycologia, 24(2) :21 5-220. P/. 5. 1932.

: The application of the generic name Guepinia, Am. J. Botany, 23(9) :627-

629. 1936.
: A new type of heterobasidiomycete, ./. Wash. Acad. Sci., 27(3) :1 12-114.

Fig. 1. 1937a.
-: New or noteworthy fungi from Panama and Colombia, I, Mycologia,

29(5):618-625. Figs. 1-29. 1937b.
— : The morphology of the basidium. Am. J. Botany, 25(9) :682-685. 1938.

LITERATURE CITED 463

— : Some Heterobasidiomycetes from Eastern Canada, Mycologia, 32(6) :6S3-

695. Figs. 1-9. 1940.
— : The Tremellales of the North Central United States and adjacent

Canada, Univ. Iowa Studies in Natural History, 18(3):l-88. Pis. 1-5. 1944.
— : New or noteworthy tropical fungi IV, Lloydia, 11(2):111-122. Figs. 1-5.

1948.

The genus Ceracea Cragin, Mycologia, 41(1) :77-86. Figs. 1-13. 1949.

MoLLER, Albert: Protobasidiomyceten, Botan. Mitt. Tropen, 8:1-179. Pis. 1-6.

1895.
Neuhoff, W.: Zytologie und systematische Stellung der Auriculariaceen und

Tremellaceen, Botan. Arch., 8(3-4) :250-297. Pis. 1-4. Figs. 1-7. 1 diagram.

1924.
Patouillard, N.: Essai taxonomique sur les families et les genres des Hym^no-

mycetes. These pour I'obtention du diplome de Docteur de I'llniversit^ de

Paris, Ecole Sup^rieure de Pharmacie, ann^e 1900-1901, No. 2. 184 pp. 74

figs. Lons-Ie-Saunier, 1900.
Rogers, Donald P.: A cytological study of Tulasnella, Botan. Gaz., 94(1) :86-

105. Figs. 1-79. 1932.
: A taxonomic review of the Tulasnellaceae, Ann. Mycolog., 31(3) :181-203.

Pis. 6-7. 1933.
: The basidium, Univ. Iowa Studies in Natural History, 16:160-183. PI. 7.

1934.

: Notes on the lower Basidiomycetes, ibid., 17(l):l-43. Pis. 1-3. 1935.

A new gymnocarpous Heterobasidiomycete with gasteromycetous

basidia, Mycologia, 39(5) :556-564. I fig. 1947.
Shear, C. L., and B. O. Dodge: The life history of Pilacre faginea (Fr.) B. & Br.,
J. Agr. Research, 30(5):407-417. Pis. 1-2. 1925.

CLASS BASIDIOMYCETEAE: SUBCLASS
EUBASIDIAE, "HYMENOMYCETEAE"

Subclass Eubasidiae

IN CONTRAST with the Heterobasidiae the Eubasidiae possess basidia
which are one-celled and which mostly do not show a sharp distinction
into hypobasidium and epibasidium. The nuclear divisions occur in the
body of the basidium and after the spores have arisen at the ends of true
sterigmata the nuclei pass through the latter into the spores. The nuclear
spindles may be parallel to the longitudinal axis of the basidium (sticho-
basidial) or at right angles to this axis (chiastobasidial). In a few species
both types of basidia may be present (Exobasidium and some Boletaceae).

In the genera included in the "Hymenomyceteae" the basidia occur
in a hymenium (in a few species they are scattered so that no continuous
hymenium is present), which becomes exposed to the air before the spores
are shot off from the sterigmata to whose tips they are obliquely attached.
These characters are in contrast to the situation in the group of orders
to which collectively the name " Gasteromyceteae " has been applied (see
next chapter). In these the spores are mostly attached symmetrically to
the tips of the sterigmata and are not thrown off. They are set free by the
opening of the basidiocarps to the air in various manners after the spores
are mature.

The fungi included as Hymenomycetes have been considered to form
one order, Agaricales; or two orders, Polyporales (or Aphyllophorales)
and Agaricales, or several orders (Heim, 1934). In the main two series
can be distinguished; those in which the basidia so far as studied are
stichobasidial (most of the Polyporales) and those in which the nuclear
division where investigated has been shown to be chiastic (Agaricales).
It must be noted that of the thousands of species in this subclass only a
few in each group have been studied cytologically.

It is not by any means settled whether the families considered to be-
long to this subclass really form a monophyletic series or whether some of

464

SUBCLASS EUBASIDIAE 465

them have arisen from one or more of the gasteromycetal orders. In the
arrangement here set forth the majority are beheved to form one series
with a few forms of more doubtful origin.

Apparently the most primitive forms produced resupinate spore fruits
with the basidia scattered or packed close together, and without a very
definite hmit of growth to the hymenium. By the formation of folds,
ridges, teeth, etc., the hymenial surface became increased in the various
families. At the same time a tendency appeared toward confining the
hymenium to the under side of laterally attached or centrally stalked
spore fruits. These spore fruits vary from felty to fleshy or leathery or
corky or woody in consistence and may function for only a few hours or
days, in some of the fleshy sorts, to many years in some of the corky or
woody species. The size may vary from a few millimeters to over a meter
in diameter and from a few layers of cells in thickness to 30 or 40 cm.
A few species are obligate parasites in the stems, leaves, fruits, and
flowers of Anthophyta (Angiosperms), e.g., Exobasidium. Some are para-
sitic or saprophytic depending upon the environment or opportunity,
e.g., Pellicularia filamentosa (Pat.) Rogers {Corticium vagum var. solani
Burt ex Rolfs). Many are parasitic upon the roots of plants, killing them
and some extend up into the stem, kiUing the bark. Some are saprophytic
on humus, decaying leaves, stems, etc. The nonliving woodcells of living
trees may be attacked and the wood rotted without actually any parasitic
action upon the living cells of the trunk. The weakening of the stem by the
decaying of its woody elements may cause it to break, thus leading to its
death. Schizophyllum commune Fr. may attack some trees as parasites
but may be saprophytic on others. Many of the more woody or corky or
leathery species are capable of culture in the laboratory but many of the
fleshy forms have resisted all such attempts as yet.

The hymenium may consist entirely of basidia all of the same age or
with younger basidia pushing up between the older ones. The newer
basidia may push out beyond those first formed so as to increase the
thickness of the hymenium which may then show definite or indefinite
layers. In a great number of forms where the development of the spore
fruit is not limited by the formation of a definite border of different struc-
ture the centrally produced basidia are the oldest and around them
additional basidia arise successively further and further from the center.
As a result in such species all ages of basidia may be found on making a
radial section, from the oldest ones near the center to those near the
margin which are just beginning to develop.

The basidia vary greatly in shape in the Hymenomycetes. They are
usuaUy round in cross section but when crowded laterally may become
somewhat angular. They may be cylindrical, tapering at the very base
and rounded at the apex, or urn-shaped or clavate or almost globose.

466 CLASS BASIDIOMYCETEAE

The sterigmata may be long in proportion to the length of the basidium
or short, stout or. slender, straight or curved. In some species they are
hardly different from those of Ceratobasidium, tentatively placed in the
Tulasnellales (see Chapter 13) . Indeed these may represent transition forms.

In a great many forms some of the potential basidia do not develop
far enough to produce sterigmata and spores. When they otherwise re-
semble not much modified basidia they are often spoken of as paraphyses.
It must be noted however that they arise from the same type of hyphae
that give rise to the basidia, so that in normal cases the paraphyses as
well as the young basidia are binucleate. In the Ascomyceteae, on the
contrary, the asci normally arise from dicaryotic hyphae and the para-
physes from monocaryotic hyphae. Thus the term paraphysis in the
Hymenomycetes is based on morphology and location of the structure,
not upon its phylogenetic implications (Ktihner, 1925a).

Besides the paraphyses some of the hyphae underlying the basidial
layer may insert themselves between the basidia in the form of much
modified terminal cells. Except for the fact that in some cases they come
from hyphae more deeply located than those from which the basidia arise
they differ from the paraphyses mainly by their greater differentiation.
They are called cystidia. These are given special names depending upon
location, shape, contents, or function. They may be simple or branched,
colorless or colored, thin-walled or with thick walls, obtuse or pointed,
barely projecting from the hymenium or far exserted. In some species of
Coprinus they serve to hold the gills apart (trabecular cystidia) while in
Hymenochaete the stiff, sharp-pointed, bristle-like cystidia probably pro-
tect the hymenium from snails, slugs, or other soft bodied animals that
otherwise might destroy the basidia. The term gloeocystidium is given to
cystidia containing mucilaginous or oleaginous contents, usually at the
ends of conducting hyphae underneath the hymenium. In a few Hymeno-
mycetes there are produced in the trama or in the hymenium stellately
branched, thick-walled cells which may be considered as being specially
modified cystidia. By their location cystidia may be called cheilocystidia,
when they develop at the edges of the pores or lamellae, or pleurocystidia
when they occur in the hymenium that lines the pores or the surfaces of
the lamellae. Sometimes cystidium-like structures that develop on the
upper side of the pileus are called pileocystidia and similar structures on
the stipe caulocystidia. It must be recognized that these last two cate-
gories although resembling cystidia are perhaps better considered special
types of pubescence, confining the use of the term cystidium to structures
in the hymenium.

The spore fruits show a very great variability of size and complexity
of structure. It seems probable that the simple forms in many cases repre-

SUBCLASS EUBASIDIAE 467

sent more primitive organisms phylogenetically, although as elsewhere
among the fungi there is ample evidence that retrogression from more
complex to simpler structures has occurred frequently. Assuming, as
appears to the author to be most likely, that in some cases at least the
simpler forms of the Hymenomycetes represent more primitive forms, the
original type of spore fruit may have been a thin structure, not many
layers of hyphae in thickness, adhering at all points to the outer surface
of the substratum within which the vegetative mycelium was actively
growing and accumulating food reserves. This young spore fruit spreads
from a central spot more or less radially and its size is limited by external
obstacles or by the exhaustion of the supply of food. Thus the sporocarp is
theoretically to be considered as of unlimited ability to extend itself.
From the external layer of hyphae there turn outward short branches
which produce the basidia as their terminal cells. These upright hyphae
may branch sympodially as the basidia approach maturity so that not
only are new basidia appearing on the new radial growth of the spore
fruit but also are arising among the older basidia. At the same time that
the basidia are developing, the paraphyses, cystidia, etc., make their
appearance but probably not to any great extent in the most primitive
forms. In the species in which the mycelial growth outside of the sub-
stratum is more rapid and widespread than the production of the basidia
the hymenium may consist of interrupted groups of sympodially produced
basidia, as occurs in some species of the genera PeUicularia {Botryo-
basidium) and Tomentella. The spore fruit may then be more or less cot-
tony with scattered clusters of basidia. Such fungi are indiscriminately
lumped in the older works, in the genus Hypochnus.

From the simple type of fruit body illustrated by the foregoing,
evolutionary development progressed in various directions. The sub-
hymenial structure became more complicated, often with two or three
distinct layers, each several or many cells in thickness {Stereum, etc.).
The sporocarps more and more developed on the substratum in such a
position that the hymenium faces downward, especially in those forms in
which the edges pull away from the substratum to form a kind of shelf,
the effuse-refiexed forms. This tendency persists until we find that in
many genera the spore fruits are shelf-like, with little or no resupinate
portion. These shelf-like structures may be narrowed toward the point
of attachment and often a distinct stipe occurs. This stipe may be
attached at the edge of the pileus or show a tendency to be attached
excentrically or even centrally to the under side of the pileus (many
Polyporaceae, Boletaceae, most Agaricaceae, and other related families).

In consistency the sporocarp may be cottony, papery, leathery, corky,
woody, or fleshy. It may be short lived or may persist for many years.

468 CLASS BASIDIOMTCETEAE

The colors may be white or shades of gray or bright-colored or almost
black.

The mycelium in perhaps the majority of the Hymenomycetes shows
clamp connections. These may be found generally in the vegetative
mycelium as well as in the spore fruit or may be lacking in the latter or in
both. In abnormal forms whose mycelium is of the monocaryon type only,
clamp connections are lacking, but their absence in any given species
does not necessarily indicate such an abnormal type. Cytological study is
required to determine whether hyphae without clamp connections are
monocaryotic or dicaryotic.

Tischler (1927) and others who have made cytological studies in the
Higher Fungi report that for the Hymenomycetes as well as most other
Basidiomycetes the haploid number of chromosomes is mostly two,
although in a number of species it may be four, six or eight.

The basidiospores are various in shape; globose to ellipsoidal to ovoid
and in some genera angular or knobbed (Rhodophyllus) . They are rarely
symmetrical in more than one plane, that which passes through the spore,
the sterigma and the center of the apex of the basidium. Apparently
almost without exception they are perched in a slightly oblique manner
on the tips of the sterigmata from which they are expelled with violence.
They are nonseptate but in Exobasidium may become transversely
septate before germinating or even before being discharged. Normally
they germinate by a germ tube which may arise from any point on the
spore wall or only from a specially located germ pore. They vary in color
from hyaline, pink, red, yellow, ochre, ferruginous, to purple and black.

Conidia are produced in this group but in a rather limited number of
species scattered throughout the two orders. They occur on various types
of conidiophores. When produced internally in the spore fruit in a cushion-
shaped or spherical structure they are usually placed in the "form genus"
Ceriomyces but when formed externally may be called Paramyces.
Chlamydospores are produced abundantly in Nyctalis asierophora Fr.
and elsewhere. Oehm (1937) concludes that there are no true conidia in the
Hymenomycetes but that they are all to be considered as various forms of
chlamydospores.

Besides the foregoing the monocaryon stage of the mycelium of very
many species produces oidia which appear to be capable of functioning as
sexual cells (see Chapter 12), but which in some cases serve as conidia.
More rarely they are produced as two-celled oidia on dicaryon mycelium
but usually the two cells then fall apart and function like the uninucleate
oidia from the monocaryon mycelium. This was reported for Plioliota
aurivella (Fr.) Quelet by Vandendries and Martens (1932).

In the author's earlier book all of the Hymenomycetes were in-
cluded in one order, the Agaricales, but the modern tendency is toward

SUBCLASS EUBASIDIAE 469

further division, even to the recognition of four or five orders. The author
conservatively recognizes two orders, Polyporales (Aphyllophorales) and
Agaricales. It must be confessed that the distinction between these two
groups is not sharp at some points, either from convergence in structure
of two otherwise very distinct orders or because one grades into the other,
representing a phylogenetic relationship.

The great Swedish mycologist Elias Fries made extensive studies upon
the Hymenomycetes as well as upon other fungi for about sixty years.
His original classification was based upon the studies of Persoon with
whose later work his earlier publications were contemporaneous. Per-
soon's chief works were his Synopsis Methodica Fungorum, 1801, and
Mycologia Europaea, 1822-1828. The work of Fries, which is used as the
standard upon which the nomenclature of many groups of fungi (includ-
ing the Hymenomycetes) is based, is his Systema Mycologicum, in four
volumes, 1821-1832. His latest important work was Hymenomycetes
Europaei, 1874. The majority of the students of this group of fungi have
followed the Friesian system with minor modifications. He recognized five
families within the limits of the Hymenomycetes as follows : >i\

Agaricaceae: hymenium on radiating gills or lamellae.

Polyporaceae : hymenium lining the surfaces of small pores or tubes.

Hydnaceae: hymenium spread over spines or protuberances.

Thelephoraceae : hymenium unilateral, spread over a firm, smooth or corru-
gated, under or upper surface. Mostly membranous, leathery, etc.

Clavariaceae: hymenium spread over the surface of smooth, simple or branched
clubs. Mostly fleshy.

Several families have been segregated from those above. In the au-
thor's first book in addition to the foregoing there were recognized the
Exobasidiaceae, segregated from the Thelephoraceae, and the Boletaceae
and Fistulinaceae, separated from the Polyporaceae.

As mentioned in the preceding chapter the studies by Juel (1896, 1916)
and of Maire (1900, 1902) showed that in the Basidiomyceteae a distinc-
tion could be made as to the position of the spindle in the meiotic division
of the diploid nucleus of the basidium. This appeared to be correlated
more or less with the relationship of the groups. In some the basidium is
more slender and the diploid nucleus occupies an approximately central
position. The spindle of the first meiotic division is parallel to the longi-
tudinal axis of the basidium as are the two spindles of the subsequent
divisions of the two daughter nuclei. The nuclei then mostly migrate to
the upper part of the basidium where the sterigmata are formed. Such
basidia are stichobasidial. In other Basidiomyceteae the basidium is
broader above and the diploid nucleus is located in the somewhat widened
upper portion. The first nuclear spindle is more or less transverse and the
next two spindles also transverse, usually at right angles to the axis of

470 CLASS BASIDIOMYCETEAE

the first spindle. Such basidia are called chiastobasidial. It was shown
that many of the Thelephoraceae and Clavariaceae and some of the
genera hitherto placed in the Agaricaceae were stichobasidial while most
of the Polyporaceae and Agaricaceae and Boletaceae were chiastobasidial.

As more and more species and genera of the Hymenomycetes became
known it was necessary to find other characters for their classification
than the external morphological ones which largely formed the bases
for the studies by Fries. The form and color and surface characters of the
spores showed their importance, but soon the anatomy of the spore fruit
and especially of the trama, the tissue upon which the hymenium is pro-
duced, proved to be of great value, as well as the mode of development of
the spore fruit. The old families had to be broken up and recombined in
order that a more logical systematic treatment could be expressed.
Patouillard (1900) recognized two main groups: "Aphyllophoracees,"
with the hymenium naked from the first and capable of continued ex-
pansion, and "Agaricacees" with the hymenium more or less lamellar
and hemiangiocarpic, i.e., at first enclosed by a more or less fugacious veil
which is variously ruptured at maturity. Later authors have called these
Polyporales and Agaricales respectively. The old families Boletaceae and
Agaricaceae made up the latter order, the Polyporales containing the
other Friesian families and one or two stichobasidial genera from the old
family Agaricaceae (e.g., Cantharellus) . In the main the Agaricales as so
delimited were chiastobasidial, but the Polyporales had both types of
basidia. This led Gaumann (1926) and others to divide the included
families into two series, stichobasidial and chiastobasidial. The fact that
in the same hymenium of Exohasidium both types may occur throws
doubt upon the validity of this as a fundamental character, although it is
apparently correlated sufficiently with other characters to make it
important.

Another character has been emphasized recently as of perhaps great
importance: the blue or violet coloration of the basidiospore exospore or
of the warts or network of lines on the spores upon treatment with a solu-
tion containing free iodine. In some cases even certain of the hyphae of the
spore fruit give the same reaction. Such spores and hyphae are said to be
amyloid (i.e., starch-like in their reaction). Just how far this can be used
in revising the arrangement of genera and families is uncertain for within
certain genera (e.g., Mycena) occur some species with amyloid and some
with non-amyloid spores.

Although the presence of clamp connections on the mycelium is well
known in a great many of the Hymenomycetes yet it has been found that
they may be absent in some genera on the hyphae in the interior of the
spore fruit. This has been used as a supplementary generic character but
cannot be considered as of fundamental importance. In the genus Coprinus

ORDER POLYPORALES (aPHYLLOPHORALES) 471

there occur species that are quite similar, some of which possess clamp
connections and some lacking them.

In the sequence of famihes followed below it must be considered that a
logical arrangement should be based upon the supposed phylogenetic
relationships of the group. In the absence of decisive fossil remains we
have to fall back upon a comparison of species and genera now existent.
The surmised origins of the different families are quite various. Thus
Singer (1936) and others have been led to believe that the Agaricales
have descended from the Gasteromyceteae while others (e.g., Heim,
1937) believe the reverse to be the case. In the following discussion the
writer follows somewhat the latter's interpretation although admitting
that the evidence for Singer's view is quite strong.

Order Polyporales (Aphyllophorales) . Hymenium always gymno-
carpic, i.e., from its inception exposed to the air and not enclosed by a
veil, with the possible exception of Cryptoporus volvatus (Pk.) Hulb.
Growth of the hymenium is more often without definite morphological
limits. In the supposedly more primitive forms (at least forms of simpler
structure) no highly developed morphological or anatomical differentia-
tion occurs beyond the resupinate layers of hyphae upon which arise the
more or less separate clusters of basidia or a continuous hymenial layer
which may continue to enlarge in all directions at the margin. In the
higher forms the subhymenial portion of the spore fruit becomes dis-
tinguished into several layers differing more or less morphologically and
anatomically and the tendency to develop upright or reflexed (shelf-like)
portions becomes more strongly marked. With evolution progressing in
various directions there appear clavate or dendroid structures or pileate
forms which are sessile or laterally, excentrically or even centrally
stipitate. Along with this external differentiation the incipient hymenium
may be increased in surface area by being thrown into folds or by the
production of emergent tubercles or teeth or by reticulate outward growth
leaving shallow or deep pits (pores) which are lined by hymenium. With
some of the simpler types of structure is correlated the stichobasidial
type of basidia, but the more complex structures mostly have the chiasto-
basidial type. The chief distinction in the following order, the Agaricales,
is the fact that in the latter the poroid or more often lamelloid hymenial
portion, though in its younger stages sometimes gymnocarpic, often
becomes secondarily enclosed (pseudoangiocarpic) or may arise from the
beginning as an internal development (angiocarpic). In both the latter
cases prior to the maturation of the basidia the spore fruits become
opened in a regular manner to permit the distribution of the spores.

In both of these orders the basidiospores are perched obliquely at the
tips of the sterigmata and are shot off violently so that spore distribution
is effected by air currents. This distinguishes the Hymenomycetes from

472 CLASS BASIDIOMYCETEAE

the Gasteromycetes in which the basidiospores are perched symmetrically
on the tips of the sterigmata and are not shot off, the spores reaching
their maturity before the spore fruit opens.

Key to Families of Pols^porales

Parasitic in the living tissues of leaves, growing stems, or fruits of Anthophyta
(Angiosperms), producing the clavate basidia externally in a continuous or
interrupted layer. Basidiospores becoming septate upon germination, each
cell giving rise to a few spindle-formed conidia. Family Exobasidiaceae
Saprophytic in plant tissues or on plant debris, less often parasitic. Basidia oval
to clavate, basidiospores germinating directly.

Hymenium interrupted or continuous, smooth (rarely slightly roughened by
small wart-like emergences) , entirely resupinate or reflexed or partially stipi-
tate, the hymenial surface being on the under side. Spore fruit a thin weft
of hyphae or a more definite structure, more often papery, leathery, or even
corky or slightly woody. Both stichobasidial and chiastobasidial types of
basidia present. Family Thelephoraceae

Spore fruits with short stipes, funnel-shaped to trumpet-shaped or almost
clavate, but somewhat broader and truncate at the top; fleshy; the hyme-
nium on the outer surface, smooth or reticulate or with low, broad, rounded
longitudinal ridges. Basidia stichobasidial. Basidiospores white or Hght-
colored. Family Cantharellaceae

Hymenium continuous on all sides of upright filiform or clavate or much ramose
spore fruits which are fleshy or leathery. Both stichobasidial and chiasto-
basidial types present. Family Clavariaceae

Spore fruit resupinate or reflexed or stipitate, both laterally or centrally.
Hymenium smooth with a number of projecting pegs or teeth or the latter
very numerous, being directed downward in all but a few of the resupinate
forms with few emergences. Spore fruits leathery, corky, woody, or fleshy.
The teeth are round in cross section or flattened. Basidia usually chiastic
but some stichic. Family Hydnaceae

Spore fruit resupinate or reflexed, or laterally or centrally stipitate. The
hymenial surface is increased by the development of shallow or deep pores
which may be round or elongated radially with greater emphasis on the
radial ridges so as to form lamellae with secondary cross connections. Papery,
leathery, corky, woody, or fleshy. Basidia always chiastic.

Family Polyporaceae

Spore fruits resupinate or reflexed or laterally stipitate, fleshy, the lower fruit-
ing surface growing out to form numerous, elongated, separate tubes which
are lined internally by the hymenium. Basidia chiastic.

Family Fistulinaceae

Spore fruits resupinate or partially reflexed, the hymenial surface at first smooth
and then thrown into thick shallow ridges which often anastomose to form
low-walled, broad, shallow pits. Hymenium continuous over the sides and
edges of the ridges as well as the floor of the pits. Fruit body often more or
less gelatinous, with the surface in almost any direction. Many species are
very destructive to wood both in trees and in structural timbers.

Family Meruliaceae

The division of families in this order is not always as above. Patouil-
laid (1900) makes an entirely different arrangement. In the older works

THELEPHORACEAE 473

the Boletaceae and Fistulinaceae were united with the Polyporaceae.
Maire (1937) and Singer (1936) removed the Cantharellaceae from the
Agaricales to the Polyporales, but Heim (1934) retains this family in the
Agaricales. He recognizes two more orders intermediate between the Poly-
porales and Agaricales: the Boletales and the Aster osporales (including
the Russulaceae).

Family Thelephoraceae. Hymenial surface smooth or at most only
slightly warty or folded. Spore fruits membranous, leathery, or in two or
more genera fleshy; closely appressed to the substratum or forming a
shelf or funnel or simple or divided pileus with hymenium on one surface
only. Twenty or more genera are recognized and probably about 1000
species. The most complete study of the North American species of this
family is that by E. A. Burt (1914-1926). Rogers and Jackson (1943)
have made a thorough nomenclatorial study of many of the resupinate
species of this family.

The genus Corticium forms a thin spore fruit growing closely appressed
to the substratum and not distinguishable into several layers. The hymen-
ium arises directly from the mycelium and consists of a layer of closely
packed basidia. The margin of the spore fruits may be definite or indefi-
nite. When dry the hymenium is often cracked. There are no true cystidia
among the basidia but gloeocystidia may be present in some species.
Most of the species of the genus are saprophytic on wood or bark, a few
are destructive to wood.

Miss Nobles (1937) demonstrated, by mating monocaryon cultures
of Corticium incrustansvon Hohn. &Litsch., that this species falls into two
sexual phases, i.e., is of the bipolar type of sexuality. Aerial hyphae of
monocaryon mycelium give rise to allantoid hyahne uninucleate conidia,
often many to a hyphal cell, which upon germination produce again the
monocaryon phase. The dicaryon aerial hyphae, which have a clamp
connection at every septum, produce a single binucleate conidium on each
hyphal cell, leaving two nuclei behind in that cell. These conidia upon
germination give rise immediately to dicaryon hyphae.

The old genus Corticium has been divided into several genera, the dis-
tinctions being based chiefly upon the structure of the basidia and the
nature of the hymenium whether loose or compact. One of these genera,
Ceratohasidium, with two to six sterigmata so long and so much thickened
as to be called "epibasidia" shows close relationship to the Tulasneflaceae
and has been considered in the discussion of that family in the preceding
chapter. It represents about a halfway step between the Thelephoraceae
in the Eubasidiae and the Tulasneflaceae and Dacrymycetaceae in the
Heterobasidiae. Another genus, Pellicularia {Botryohasidium) has been
segregated by Rogers (1943) for those fungi formerly included in Corticium
which have a thin film of short, broad-celled mycelium on the substratum

474

CLASS BASIDIOMYCETEAE

Fig. 153. Polyporales, Family Thelephoraceae. (A, B) Pellicularia isabeUina (Fr. )
Rogers {Tomentelln flava Bref.) (A) Loose weft of mycelium bearing below scattered
basidia and above conidiophores. (B) Basidium and spores. (C-E) Aleurodiscus
amorphus (Pers.) Rabenh. (C) Habit sketch. (D) Section of hymenium with basidia
and "paraphyses." (E) Basidiospore. (A, after Brefeld: Untersuchungen aus dem
Gesammtgebiete der Mykologie, Heft 8, pp. 1-305. B, courtesy, Rogers: Univ. Iowa
Studies in Natural History, 17(l):l-43. C-E, after Killermann, in Engler and Prantl:
Die Natiirlichen Pflanzenfamilien, Zweite Auflago, vol. 6, pp. 124-288, Leipzig, W.
Engelmann.)

THELEPHORACEAE 475

and cymose tufts of short, broad basidia with four, less often six to eight,
sterigmata. The commonest parasitic species of this genus, P. filamentosa
(Pat.) Rogers {Corticium vagum var. solani Burt, Hypochnus solani
Prill. & Del., etc.) occurs as a parasite upon the stems and roots of potato
(Solarium tuberosum L.), bean (Phaseolus vulgaris L.), and very many
other plants of economic value. It produces cankers at or below the sur-
face of the soil which kill or seriously injure the parts affected. Small
sclerotia are formed which enable the fungus to overwinter. On the stem
of the host plant the mycelium creeps up as a thin gray or white hyphal
layer on which the oval basidia are produced in groups. The sclerotial
stage is known under the name of Rhizoctonia. (Fig. 153A, B.)

The genus Tomentella {Hypochnus, as interpreted by Burt, 1916)
produces its basidia more or less scattered or in tufts on a loose cottony
mycelium. The basidiospores are nearly spherical and spiny. They are
mostly saprophytic.

The species of Corticium in which gloeocystidia occur, are placed by
many authors in a separate genus, Gloeocystidium, but Burt (1926) and
Rogers and Jackson (1943) do not approve of this segregation. Peniophora
is practically a Corticium with true, fusiform, pointed cystidia (not
gloeocystidia). Asterostroma is similar but has stellately branched, thick-
walled bristles or cystidia in the hymenium. Epithele has the hymenium
interrupted here and there by sterile projections or pegs consisting of
bundles of hyphae. These differ from the teeth of the Hydnaceae which
are covered with basidia. Coniophora is practically a Corticium with
ferruginous spores. Some species have cystidia {Coniophorella Karst.)
and others not. Donk (1933) and Singer (1944) place the Family Merulia-
ceae close to these genera (see p. 484).

In Aleurodiscus the spore fruit instead of remaining fiat against the
bark on which it develops curls up a little at the edge to form a fiat saucer
or shallow cup. The basidia and spores are rather large and there are
present various types of structures called by some cystidia, by others
paraphyses. Much similar is Vararia (Aslerostromella) but the modified
cells in the hymenium are many times dichotomously branched. (Fig.
153, C-E.)

All of the foregoing genera have a rather thin and not much differ-
entiated spore fruit below the hymenium. The following genera have a
subhymenial structure much thicker and often in several distinct layers.
At the edges the spore fruit bends away from the substratum to form a
sort of shelf, with the hymenium on the smooth lower surface. Even a
sort of central stipe may be developed in some species, the spore fruit
being more or less funnel-shaped in that case. In Stereum conspicuous
cystidia are lacking, but in Hymenochaete the hymenial layer has numer-
ous long stiff, usually brown, pointed setae, whose probable function is

476

CLASS BASIDIOMYCETEAE

Fig. 154. Polyporales, Family Thelephoraceae. (A, B) Thelephora terrestris (Ehrh.)
Fr. (A) Lower jview of lobe. (B) Section through hymenium. (C) Hymenochaete
cacao Berk. Portion of hymenium showing basidia and setae. (D) Solenia Candida
(Hoffm.) Fr., habit sketch. (A-D, after Killermann, in Engler und Prantl: Die
Naturlichen Pflanzenfamilien, Zweite Auflage, vol. 6, pp. 124-288, Leipzig, W. Engel-
mann.)

the protection of the hymenium from snails and other harmful animals.
In Thelephora (called by the name Phylacteria Pat. by Patouillard, 1887,
1900; Rea, 1922, and others) the more or less leathery fruit body is
upright, stalked, pileate or fan-shaped or much lobed or in an overlapping
series, the hymenium being on the under side and smooth or slightly
warty. In Cyphella and Solenia (united into one genus by some authors)
the spore fruit is cup-like or tubular, with the opening usually directed
downward, and lined internally by the hymenium. In some species of
Solenia many spore fruits arise close together, but separate from one
another, from a common mycelial mass, the subiculum. Some mycologists
place these two genera in Family Fistulinaceae (see p. 484). (Fig. 154,
A, D.)

Sparassis (often included in the Clavariaceae, but see Cotton, 1912)
is fleshy, much branched, with the terminal portions flattened and bearing
the hymenium on the under side only. The edible S. ramosa (Schaef.)
Schroet. occurs in Europe and North America, often attaining the size
and shape of a small, loose-leafed cabbage.

Family Cantharellaceae. Perhaps intermediate between the
Thelephoraceae and Clavariaceae is the genus Craterellus. This is fleshy,
club- or funnel- or trumpet-shaped with the hymenium on the outer side
which is smooth or more or loss longitudinally ribbed or reticulate. Several
species are edible. The genus is stichobasidial as is the genus Cantharellus
which is undoubtedly closely related but differs in having the longitudinal
ribs further developed so as to be low, thick lamellae. Perhaps, as Singer
suggests, the correct name is Gomphus. Cantharellus was formerly included
in the Agaricaceae but probably belongs here. Perhaps these two genera
should be united into a distinct family, the Cantharellaceae (as was done

CLAVARIACEAE

477

Fig. 155. Polyporales, Family Cantharellaceae. Cantharellus floccosus Schw. (Gomphus
floccosus (Schw.) Singer). (Courtesy, M. B. Walters.)

by Maire 1900; Rea, 1922). This family shows close affinities with the
following and may be really more closely related to it than to the
Thelephoraceae. (Fig. 155.)

Family Clavariaceae. Spore fruits fleshy or waxy or even gelatinous,
rarely leathery; upright, clavate or branched in a coralloid manner;
round or flattened; usually covered by the hymenium on all sides over
the whole spore fruit or over special more terminal and often enlarged
portions. A dozen or more genera and probably over 500 species. Almost
all are saprophytic and possibly some form mycorrhizae on tree roots.
A few species are parasitic upon plants. Many of the larger fleshy forms
are edible. Some of the genera seem to have their origins in the Thele-
phoraceae and some are perhaps nearer to the Cantharellaceae, and per-
haps some are related to Irpex in the Polyporaceae or to Hericium in the
Hydnaceae. In other words, the Clavariaceae probably do not represent
a phylogenetic unit. Some are stichobasidial and some chiastobasidial.
How great weight this should have upon the division into genera must

478

CLASS BASIDIOMYCETEAE

await further studies in an attempt to determine whether the structural
differences within the group show definite correlations with the stichic
or chiastic position of the nuclear spindle in the meiotic divisions of the
diploid nucleus of the basidium.

Pistillaria is a small fungus usually not over 2 to 5 mm. tall, with no
sharp distinction between stalk and hymenium-bearing portion, which
is clavate. No sclerotium is present. Some of the species have two-spored
basidia, others have four spores to each basidium. Growing on soil or
dead stems, leaves, etc. Also small, but much taller than the foregoing,
is the genus Typhula whose spore fruits grow from sclerotia and form
very slender filiform stalks, sometimes branched, with the terminal por-
tions thickened to form spindle-formed hymenophores. A number of
species are parasitic upon grasses, sugar beets, potatoes, etc. Miss
Remsberg (1940) studied the two genera and determined the presence
and absence of sclerotia to be the best distinguishing character.

Physalacria is also small, up to 2 cm. tall, consisting of a slender stalk
and a downturned, hollow, fleshy head on whose lower surface the
hymenium is most abundantly developed. For this reason McGuire (1939)
suggested that the genus should be placed in the Thelephoraceae as was

Fig. 156. Polyporales, Family Clavariaceae. Clavarindelphus pistillaris (Fr.) Donk.
(Courtesy, Coker: The Clavarias of the United States and Canada, Chapel Hill, Univ.
is'^orth Carolina Press.)

CLAVARIACEAE

479

Fig. 157. Polyporales, Family Clavariaceae. Clavariella subbotrytis. (Courtesy,
Coker: The Clavarias of the United States and Canada, Chapel Hill, Univ. North
Carolina Press.)

done for Sparassis by Cotton (1912) because the ultimate flattened
branches of the much ramose spore fruit of the latter bear their hymenium
on the lower surface only. Killermann (1928) retains both these genera in
the Clavariaceae.

The larger forms of this family are clavate and unbranched or only
slightly branched or very much branched in a more or less coralloid
manner. Among the clavate forms is the very large Clavariadelphus
pistillaris (Fr.) Donk {Clavaria pistillaris), often up to 10 to 15 cm. tall
with a thickness of 2 to 3 cm. It is edible. In many ways it resembles
some of the members of Family Cantharellaceae but differs in possessing
chiastic instead of stiehic basidia. Somewhat smaller and usually brighter-
colored are the unbranched species of Clavaria which are not enlarged
much upwards and have smaller spores. Some species are much branched
and fleshy. Where the branching consists of very slender dry cartilaginous,
cylindrical and tapering branches, forming a bush-like structure we have

480 CLASS BASIDIOMYCETEAE

the genus Pterula. Eriocladus (Lachnodadium) is larger, with flattened
or rounded branches which are hairy. Some of the much branched fleshy
species of Clavaria have been set off as separate genera: Clavulina and
Clavariella (Ramaria). The majority (but not ah) of the species of these
two genera are respectively stichobasidial and chiastobasidial. Gaumann
(1926) and Donk (1933) accordingly place Clavulina in a distinct order,
the Cantharellales, along with some segregates from the Thelephoraceae
and Hydnaceae and Agaricaceae. The genus Clavicorona, with apices of
the branches truncate or cup-shaped, and with gloeocystidia in the
hymenium layer has been set apart from Clavaria by Doty (1947).
(Figs. 156, 157.)

Family Exobasidiaceae. The fungi composing this family are para-
sitic in the leaves, green stems or even fruits of higher plants, often
distorting the affected parts or causing the formation of galls. The inter-
cellular mycelium apparently bears no clamp connections. It sends rod-
shaped or branched haustoria into the host cells. The basidia arise singly
or in tufts between the epidermal cells, eventually piercing the cuticle
and forming their usually four to eight spores externally. There are no
paraphyses or cystidia. Eftimiu and Kharbush (1927) have made an
extensive study of a species of Exobasidium. The mycelial cells within the
host are elongated but become shorter nearer the epidermis and there are
binucleate. The fusion nucleus in the basidium divides meiotically into
four nuclei, one passing into each basidiospore. The nuclei may divide
before the spores are formed and then the eight nuclei enter eight basidio-
spores. Sometimes one nucleus of the four degenerates and the remaining
three divide so that six spores are formed. Sometimes only two nuclei are
found in the basidium and then only two basidiospores. These spores
germinate by budding like yeasts often becoming once septate first. In
E. rhododendri Cramer, the spores divide by a septum and send out germ
tubes from each of the two cells thus formed. In this species the nuclear
divisions within the basidium may be either stichobasidial or chiasto-
basidial. Gadd and Loos (1948) report that in E. vexans the basidiospores
become once septate before they are discharged. These two-celled spores
may develop thick walls after becoming free. E. vaccinii (Fuckel) Wor. is
frequent upon the cranberry (Oxycoccos macrocarpus (Ait.) Pursh) and
related plants. The affected shoots become upright and take on a pink
color and the leaves are increased in size and the stem thickened. Some-
times only a small portion of a leaf may be infected or a spot on one side
of a fruit. Such spots are thickened and reddened. The basidia are club-
shaped with four basidiospores which become septate before germination
and send out short branching sterigmata bearing spindle-shaped spores.
About 30 species are known in this genus. The genus Kordyana probably
also belongs in this family. It is tropical. The basidia are two-spored,

HYDNACEAE

481

rarely four-spored and stichobasidial and emerge from the stomata of
the host, sometimes intermingled with long, slender, hyaline hyphae or
"paraphyses." This was described rather fully by Gaumann (1922).
(Fig. 158.)

The following genera have been assigned to this family but later study
has shown that they belong elsewhere : Micro-
stroma, Protocoronospora, and Urohasidium.
Wolf (1927, 1929) showed that the first is not
a Basidiomycete at all because its supposed
basidia are multinuclear, the mycelium lacks
conjugate nuclei, and the spores are not borne
upon the sterigmata in the manner typical of
the class. Wolf (1920) and Karakuhn (1923)
showed that the second, like the former,
belongs to the Fungi Imperfecti, with the
name properly Kahatiella. Urohasidium
described by Giesenhagen (1892) was shown
by Mason (1941) to be still another Imperfect
Fungus. Its proper name is Zygosporium. (Fig.
201B, C.)

The relationship of this family is not
certain. Perhaps it represents a line derived
from the simplest Thelephoraceae but highly
modified by its extreme parasitism.

Family Hydnaceae. These fungi are
mostly saprophytic. Some cause serious decay
of timber. The spore fruit may be small or
large and resupinate or shelf-like or with a
pileus borne on a lateral or central stipe. In
some forms the pileus is divided into many
small pilei. The consistency varies from fleshy
to woody and there is a great range of color. The under side of the
fruit body is at first smooth but as the hymenium develops it grows
out into hymenium-covered spines or teeth. Under the classification
of Fries all such fungi were classed in one family. However, Bourdot and
Galzin (1927) placed the genera with hyaline or light-colored spores in
the group Hydnes (except Irpex which they place in the Pores) and those
with brown spores in the Phylacteries (i.e., close to Thelephora or
Phylacieria). Donk (1933) places the stichobasidial forms in Tribe
Hydneae of the Cantharelloideae and the remaining, chiastobasidial
forms in the Phylacteroideae close to Thelephora, except some species of
Irpex placed by him in the Polyporoideae, Tribe Daedaleae. Irpex is
placed by Singer (1944) in the Polyporaceae, and by Murrill (1907) in

Fig. 158. Polyporales,
Family Exobasidiaceae. Ex~
obasidium vaccinii (Fuckel)
Wor. Basidia in various
stages of maturity emerging
through the epidermis of the
host leaf. (After Killermann,
in Engler und Prantl: Die
Nattirlichen Pflanzenfami-
lien, Zweite Auflage, vol. 6,
pp. 124-288, Leipzig, W.
Engelmann.)

482 CLASS BASIDIOMYCETEAE

the same family under the name Irpiciporus. Miller (1933) regards
Hydnochaete, Sistotrema, Irpex, and Echinodontium, whose teeth are
formed by the breaking up or unequal growth of the pore walls, as
properly placed in that family. Hydnum, as recognized by Fries in 1821,
included all of the genera now included in the family as well as some of
the above mentioned ones that may eventually have to be placed else-
where. In the following the generic distinctions of Miller (1933) are
followed in the main.

Grammothele Berk. & Curt, is resupinate with its porose-reticulate
surface covered with numerous small warts over which the hymenium
extends. Caldesiella Sacc. is in habit much like TomenteUa of the Thele-
phoraceae, but is covered with soft conical spines or teeth. The spores are
colored and rough. Asterodon Pat. is similar but has dark, simple or
stellately branched, setae, and the smooth spores are subhyaline. Gran-
dinia Fr. is also resupinate and resembles Corticium from which it differs
in the development toward maturity of hemispherical, cylindrical or
subulate warts or spines, covered with hymenium. Cystidia are lacking
and the spores are hyaline and smooth or roughened. Odontia Pers. differs
from the preceding by the presence of cystidia. It therefore resembles
Peniophora except for the conical to subulate or cylindrical spines.
Oxydontia Miller lacks cystidia and has long subulate and conspicuous
teeth. It is resupinate or effused-reflexed. This is the genus called Ada,
Karst., an untenable name because of its prior use for one of the Rosaceae.
Radulum Fr. (including Phaeoradulum Pat.) is resupinate or reflex, with
coarse, blunt, irregularly scattered or confluent teeth. Spores hyaline or
light-colored. Phlehia with hymenium covered with more or less notched
wrinkles is sometimes placed in this family. These last four genera suggest
how the Hydnaceae may have arisen from the resupinate Thelephoraceae
in which the smooth hymenium often shows little warts or even pro-
jecting bundles of hyphae. However, these latter are not covered by the
hymenium. Mucronella Fr. is essentiall}'' a cluster of subulate teeth
arising from a fugacious mycelium.

The following genera are attached laterally or have stalks. Steccherinum
S. F. Gray is sessile or substipitate and laterally attached. It has terete
or flattened spines and develops cystidia. The spores are white and small.
It is wood inhabiting. Auriscalpium S. F. Gray is laterally stipitate and
has short subulate spines with scarcely differentiated cystidia. Growing on
cones of conifers. The following three genera are centrally stipitate with
subulate spines, and grow on the ground: Dentinum S. F. Gray, fleshy,
pale, spores white and smooth, subspherical; Hydnum L. emend. S. F.
Gray, fleshy, dark-colored, spores subspherical, angular or echinulate,
brown; Calodon Qu(51. (Hydnellum and Phellodon of Karsten), fibrous,

HYDNACEAE

483

Fig. 159. Polyporales, Family Hydnaceae. Hydnum imbricatum L. ex S. F. Gray.

(Courtesy, M, B. Walters.)

tough and sometimes woody, dark-colored, spores shaped as in Hydnum,
brown or subhyahne. (Fig. 159.)

Usually placed in this family is the genus Hericium Pers. ex S. F. Gray
(Dryodon Quel, Manina Scop, ex Banker). This is fleshy and unbranched
or more often branched, with subulate spines mostly long and pendent.
Spores spherical or subspherical and amyloid (i.e., walls staining blue
with iodine). In H. coralloides Pers. ex S. F. Gray the pileus is but little
developed, the fruit body consisting essentially of branching stalks
bearing at their tips pendent tufts of pointed teeth. The writer questioned
in 1935 whether such a fungus belongs in this family at all and suggested
that it might be more closely related to the Clavariaceae. Singer (1936)
suggests its origin from much branched species of that family with which

484 CLASS BASIDIOMYCETEAE

they have in common the amyloid type of spore. For Irpex and Echino-
dontium see Polyporaceae (p. 494).

Family Fistulinaceae. This small family of only a few species and
two, possibly more, genera is distinguished by its fleshy spore fruits and
by the separate though closely crowded hollow tubes which hang down
underneath the pileus. The latter may be centrally or laterally stipitate
in Fistulina, or resupinate in other genera. Each separate tube is formed
as an open cup which elongates and becomes lined internally by the
hymenium. If one would imagine hundreds of spore fruits of Solenia of
the Thelephoraceae arising from a common pileus the characteristic struc-
ture of this family would be realized. Fistulina hepatica Fr., the beef-
steak fungus, is found in nearly all temperate regions of the world on
oaks and other deciduous trees. The spore fruit is more or less fan-shaped
with a short, thick, lateral stipe. It is brown-red, blood-red internally,
and the closely crowded but separate tubes are reddish brown. When
young it is edible. Porothelium perhaps belongs here. It forms a resupinate
membranous or crust-like spreading hymenophore on which develop
numerous scattered wart-like projections which elongate, leaving a central
pore in each. This perhaps represents a connecting link between Solenia
and Fistulina. The relationship of this family to the Polyporaceae is
doubtful. Lohwag and Follmer (1936) and Elrod and Blanchard (1939)
studied the development of Fistulina and showed that its tubes are of
essentially the same structure as the cup-like spore fruits of Solenia and
Cyphella. There is no similarity in the development to the Boletaceae to
which the easily separable pores of the latter had suggested relationship.

Family Meruliaceae. Like so many others of the Hymenomycetes
the members of this family are wood destroyers. The spore fruits at first
develop a smooth hymenium on which basidia develop to spore bearing
maturity. As this enlarges the surface produces low, rather thick folds
or ridges on the sides and rounded edges of which new basidia continue
to develop so that eventually basidia of all ages are to be found on the
ridges as well as on the intervening hymenium at the original level. The
ridges may anastomose so as to form a net-work, the pores that form the
meshes being shallow. Sometimes the ridges are more or less radiate or
may be interrupted. The spore fruits and the ridges are fleshy or waxy
or gelatinous. The low walls separating the pores do not necessitate their
pointing downward since air currents can carry off the spores when
discharged from the basidia regardless of the direction in which the pores
point. There is no agreement as to what genera should l)e included in this
family. Bourdot and Galzin (J 927) include seven genera, three of which
have a smooth hymenium (Coniophora, Coniophorella, and Jaapia), the
other four having projecting ridges or tubercles. Of these Phlehia is often
placed in the Hydnaceae and Plicatura (Trogia) in the Agaricaceae,

MEKULIACEAE

485

leaving Merulius (with colorless spores) and Gyrophana Pat. (with brown
spores). According to Singer (1944) this should be called Serpula Pers. ex
S. F. Gray. Rea (1922) has the same arrangement except that he does
not separate Gyrophana from Merulius. These last two genera may be
looked upon as of Thelephoraceous origin, with the added feature of the
folds, ridges or tubercles that increase the hymenial surface. The best
known species are M. lacrymans (Wulf.) Schum. {Gyrophana lacrymans

Fig. 160. Polyporales, Family Meruliaceae.
Merulius lacrymans (Wulf.) Schum. (Courtesy,
Falck, in Moller: Hausschwammforschungen in
amtlichem Auftrage, vol. 6, pp. 1-405.)

(Wulf.) Pat.) and its close relatives which have been given intensive
study by Falck (1912). This author believes that it is distinct enough as
a house fungus ("Hausschwamm"), both in habits and details of struc-
ture to deserve specific distinction under the name M. domesticus Falck.
It is a very destructive enemy of floor boards, beams and other wood
construction in buildings, causing a red-colored dry rot. It spreads be-
tween timbers in sheets and strands and in openings between the wood-
work forms great cottony masses of mycelium. It forms resupinate sheets
on the floors and walls and even ceilings of rooms. These enlarge, with

486 CLASS BASIDIOMYCETEAE

white margins and brown central portion on which the hymenium de-
velops. This rapidly becomes wrinkled and porose and sometimes in older
stages with flattened teeth much like those of Irpex. It occurs very
extensively over Europe and Asia but is comparatively rare in America.
Some species of MeruUus are laterally attached and form shelf -like spore
fruits. (Fig. 160.)

Family Polyporaceae. The fungi here included in one family are
placed in two or more families by some of the more modern authors
(Rea, 1922; Donk, 1933; Singer, 1944). Until further studies show the
definite hmits of these different families it may be well to take the more
conservative stand and retain the one family.

In the sense that the cells invaded by the hyphae of the fungi of this
family are mostly no longer Hving, i.e., wood fibers and tracheary tissue,
these fungi are saprophytes. Many of them, however, attack only the
sap wood of living trees in which living cells are intermingled with the
dead fiber and tracheary cells, bringing about a ''sap rot" and death of
the tree. Others, though confined to the heart wood, which contains few
if any living cells, attack this wood only in living standing trees. Still
others attack only the wood of dead trees or of structural timbers. So
there are all grades of practical parasitism even though the particular
cells invaded are not living. A very few species of this family are directly
parasitic upon other fungi. Although most species are wood inhabiting
some grow on the ground, obtaining their nourishment from buried
pieces of wood or from the vegetable matter in the soil. Such species are
true saprophytes.

The spore fruits may be fleshy or fleshy-leathery when young but at
maturity are, with few exceptions, papery, leathery, corky, or even
woody. They range in size from a few millimeters in width and 1 or 2 mm.
in thickness to a width of 75 cm. (specimens of Ganoderma applanatum
(Pers. ex Fr.) Pat. collected by the author) and 30 to 50 cm. thick (speci-
mens of Fomitopsis (Fomes) officinalis (Vill.) B. & S. seen by the author),
and a width of over two meters in Polyporus squamosus (Huds.) Fr.,
according to Clements (1910). They may be evanescent or may live
many years; according to Atkinson over 80 years in the case of specimens
of Phellinus {Fomes) igniarius (Fr.) Pat.

The spore fruits may be closely appressed to the sides of tree branches,
logs, boards, etc., without a free margin, or may grow out laterally like
a shelf or bracket, or may be stalked laterally or centrally. The underside
is usually smooth when young, as in the Thelephoraceae, but develops
unevenly so as to leave numerous pits (pores) of various shapes on whose
inner face the hymenium develops. With but few exceptions the pores
are directed downward so that as the spores are shot off from the sterig-
mata of the basidia that line the pore they drop down and out of the

POLYPORACEAE 487

pore into the open air, where they are carried off by currents of air.
Cystidia of various types may be present in the hymenium and in some
species stiff, pointed, brown setae similar to those in Hymenochaete of the
Thelephoraceae. This has led to the suggestion that the various species
with setae now distributed among several genera, but especially in
Phellinus, in the Polyporaceae, should be placed in the same group as
the above mentioned genus. With very few exceptions the hymenium is
confined to the sides of the pores, not usually being formed on the edges
as in Meruliaceae. Conidiophores are produced on the upper surface of
the sporophore in some species. Under certain environmental conditions
a large portion of the tissue of the spore fruit may be converted into
chlamydospores, a condition upon which were based the genera Cerio-
myces and Ptychogaster.

The vegetative mycelium is slender and branching, the individual
cells often being rather long. Some species of Polyporaceae produce large
tuber-like subterranean sclerotia the size of a man's head (e.g., Pachyma
cocos Fr.). From these the spore fruits arise when conditions are favorable.

The processes of sexual reproduction are known in only a few cases in
the family. The mycelium of the spore fruits of many species appears
mostly to have clamp connections as does to a large degree the vegetative
mycelium growing in the wood. The basidia eventually have four or eight
nuclei, four of which pass into the four basidiospores whose nuclei in
some cases divide so that the spores become binucleate. In such species
the germ tubes show by the presence of clamp connections that secondary
or dicaryon mycelium is produced immediately upon germination of the
spores. Cultures of uninucleate basidiospores produce monocaryon
mycelia which lack clamp connections. When compatible mycelia come
into contact diploidization occurs and the dicaryon mycelium may de-
velop clamp connections, or in rarer cases they may fail to develop
although the mycelium is dicaryotic.

Mounce and Macrae (1936) showed that in Gloeophyllum saepiarium
(Wulf.) Karst. {Lenzites saepiaria), Coriolopsis trabea (Pers.) B. & S.
(L. trahea) and Trametes americana Overh. the monocaryon mycelia
arising from germination of the basidiospores fall into only two mutually
compatible classes, therefore these species are of ''bipolar" sexual be-
havior. However, there is complete compatibility between all monocaryon
mycelia derived from sporophores collected in different geographic
regions. This indicates that geographical races exist in these fungi as has
been demonstrated in the Ustilaginales (see p. 378) and some of the
Heterobasideae (see p. 453). Robak (1936) reported that Hirschioporus
abietinus (Dicks, ex Fr.) Donk (Polystictus abietinus) is, on the contrary,
quadripolar in its sexual behavior. He also showed that the monocaryon
mycelium of Coriolellus serialis (Fr.) Murr. {Trametes serialis), a bipolar

488 CLASS BASIDIOMYCETEAE

species, is just as capable of causing rot when inoculated into wood blocks
as is the dicaryon mycelium. The production of oidia by the monocaryon
mycelia of Polyporaceae has been demonstrated by Vandendries (1936)
in Leptoporus adustus (Fr.) Quel. (Bjerkandera adusta (Fr.) Karst.). Such
oidia do not occur on the dicaryon mycelium of this species. This species
is quadripolar as are Leucoporus hrumalis (Fr.) Quel, and L. arcularius
(Batsch) Quel., neither of which produces any oidia.

The number of species in this family is very uncertain. Several
thousand have been described but it is probable that very many of these
are synonymous. The opinions as to the validity of described species vary
greatly. Thus in Gaumann-Dodge, Comparative INIorphology of Fungi
(1928) the genus Polysticius is credited with nearly 1000 species while
Killermann (1928) in the second edition of Engler and Prantl, Die
Nattirlichen Pflanzenfamilien, admits only "some hundreds." The agree-
ment is still less as to generic limits. Killermann recognizes 16 genera in
the family limits adopted in this work, but Murrill, in North American
Flora (1907-1908) recognizes 78 genera for North America alone. Bon-
darzew and Singer (1941) have made a very complete and radical revision
of the genera of this family. They exclude 11 genera from the old family
limits but still retain 53 genera. Their basis for segregation and classifi-
cation of the genera is largely anatomical, so that the 11 old Friesian
genera are broken up into many sharply defined and not so unwieldy
ones. Singer (1944) adds one genus and gives further information as to
the systematic arrangement within the family. William Bridge Cooke
(1940) recognizes 46 genera from North America (most of them also
occurring in Europe) and 19 more from the Tropics and the Southern
Hemisphere. The anatomical studies by Miss Ames (1913) contributed
considerably to the work leading to the further subdivision of the older
genera.

In the following discussion of the more important genera of the family
the attempt has been made to give the modern names of the genera and
species mentioned as well as the names that are to be found in the older
standard works.

The genus Poria was in its customary limits used for the completely
resupinate members of this family, regardless of the color and consistency
of the trama, color of the spores, etc. Studies by Baxter (1929-1949) and
others have shown that many species closely related to other genera may
develop in a resupinate manner and thus be assigned to the genus Poria,
which as a consequence became a catchall for unrelated forms which
agreed only in their resupinate habits. Nevertheless, it seems that there
remains a body of species that are more or less closely related and which
properly may be given this name. Poria produces resupinate spore fruits
which adhere to the substratum and consist mainly of a thin layer of

POLYPORACEAE

489

underlying mycelium and the pore layer. Some species produce their
fruiting bodies only on the under side of branches or logs so that the
pores point directly downward, but this is not the case in all species. The
pores are mostly rather small, angular or round, and not very deep.
The spore fruits vary in size and color as well as in color of the spores.
Some species ascribed to Poria have the structure of trama and hymenium
characteristic of some of the species of Corticium and other closely related
Thelephoraceae, differing only in their poroid habit. They probably belong
in that family. It may be that they are intermediate forms between the
two groups. Aside from these forms and those that are closely related to
nonresupinate genera the remaining resupinate species have been divided
further on the basis of texture, and color of trama and spores (Murrill,
1907; Donk, 1933; Cooke, 1940; Bondarzew and Singer, 1941).

The annual-fruited forms with effused-reflexed, or shelf-like or stipitate
structures and with membranous or leathery texture and with pileus and
pore trama similar and continuous were formerly included in the old
genus Polystictus with many hundred species. By the more recent stu-
dents of this group this genus has been broken up into six or eight or
more genera. Perhaps the commonest species of this group is Coriolus
versicolor (L. ex Fr.) Quel. It is very common on dead stumps, logs, etc.,
and forms great numbers of overlapping semicircular or kidney-shaped,
velvety-haired pilei, which are strongly marked by zones of various colors.
The individual pilei are 2 to 5 cm. in diameter and may grow together
at the margin to form broad sheets if they are emerging from the cut top
of a stump. On the under side of a log they may be resupinate or effused-
reflexed. Hirschioporus abietinus is grayish white and hairy above and
concentrically furrowed, the edge of the pileus and the pore surface being
violet colored in fresh specimens. They are formed on branches, logs, etc.,
being resupinate on the under side of the substratum but forming shelves
at the sides. Mostly on coniferous wood. The pores in age break up into
flattened teeth. Coltricia perennis (L. ex Fr.) Karst. grows on the ground
usually in coniferous forests and forms a funnel-shaped, centrally stipitate
spore fruit, brown and velvety above when young, glabrate with age,
more or less strongly concentrically marked.

The genus Polyporus in its older limits differed mainly from Polystictus
in being as a rule larger and thicker and more fleshy when young, and .
with the trama of the pileus usually different from that of the pore layer
so that the latter sometimes separates from the former in age. At maturity
the spore fruits become cheesy or leathery or corky, rarely woody or
membranous. Fifteen or more genera have been segregated by some of
the modern students of this genus. Donk (1933) and Singer (1944) sepa-
rated off some of the species forming the genera Boleiopsis Fayod and
Scutiger Murrill, and Grifola (S. F. Gray) and placed them near the

490

CLASS BASIDIOMYCETEAE

Thelephoraceae and Clavariaceae because of the similarity of the struc-
ture of the spore fruit and spores to those famihes, in spite of the occur-
rence of the hymenium in pores. The genus Polyporus in the Hmited sense
consists of tough, fleshy, centrally or laterally stipitate forms. P. iuherasier
(Jacq.) Fr., considered by Donk and Singer to be the type species of the
genus, develops, underground, large, hard, sclerotium-like structures of
intermingled hyphae and particles of soil, called pietra fungaia in Italy.
When placed in a warm, moist situation several sporophores develop from
each sclerotium. These are centrally stipitate, with pileus somewhat
funnel-shaped at maturity, scaly, yellowish, with rather large pores. This
is fleshy when young and is prized for food by the Italians who collect
the sclerotia and preserve them for some time and grow from them the
edible spore fruits. Closely related is P. squamosus (Huds.) Fr. (P. caudi-
cinus (Scop.) Murr.), which is very common and destructive to many
kinds of deciduous trees. Its stipe is usually lateral or eccentric and the
pileus may exceed a diameter of 50 cm. and a thickness of 3.5 cm. It
occurs in imbricated masses growing from the trunks of the infected
trees. It also is edible when young. Other fungi included in the old genus
Polyporus are the following: Laetiporus sulphureus (Bull, ex Fr.) Murr.,

%' 4- «^'

^Si*^"

Fig. 161. Polyporales, Family Polyporaceae. Grifola berkeleyi (Fr.) Murr. (Courtesy,

M. B. Walters.)

POLYPORACEAE 491

Fig. 162. Polyporales, Family Polyporaceae. Laetiporus sulphureus (Bull, ex Fr.) Murr.

(Courtesy, F. C. Strong.)

which forms a series of shelves up to 60 cm. broad, bright yellow to
orange in color, and rather fleshy and edible at first and dry and cheesy
at maturity. The dried sporophores when ground up and soaked in water
are edible upon cooking. It occurs at the bases of trunks of deciduous
trees, often oak, whose wood it destroys. Occurring very commonly on
standing trunks of birch {Betula) are the spore fruits of Piptoporus
hetulinus (Bull, ex Fr.) Karst. The more or less hoof-shaped, reniform or
globose sporophores are attached by a narrowed, almost stipe-like portion
to the side of the trunk. The gray or whitish surface consists of a thin
layer which flakes off with age from the white pilear trama. The layer of
3 to 8 mm. long pores separates easily from the thick layer of the pileus.
(Figs. 161, 162.)

The old genus Trametes was supposed to be characterized by the
uninterrupted continuation of the pilear trama into that of the pore
layers, with the further character that the pores were of different depths
in the same spore fruit. These are true of some species but also occur in
some of the genera formerly included in Polyporus, Polystictus, and Fomes.
As a result the genus Trametes has been much reduced and segregated
into six or eight genera. Among these is Pogonomyces hydnoides (Schw.)
Murr., a very common species of Florida and the Tropics. Its dimidiate,
sessile, sometimes imbricate spore fruits may be 5 to 10 cm. broad and
up to 1 cm. thick. The upper surface is covered by long, black, stiff,
branched fibers which resemble considerably the teeth of some species of
Hydnaceae. The pileus trama is dark brown, punky to corky and that
of the tubes light brown. Pycnoporus cinnabarinus (Jacq. ex Fr.) Karst.

492

CLASS BASIDIOMYCETEAE

Fig. 163. Polyporales, Family Polyporaceae. Ganodernia applanatum (Pers. ex Fr.)
Pat. Habit view on dead trunk of maple (Acer). (Courtesy, F. C. Strong.)

is also dimidiate, 4 to 10 cm. in diameter and up to 1 cm. thick. It is
bright orange to cinnabar-red and the trama also is red, as are the
pores. It occurs on dead wood of various deciduous trees in America,
Europe, and Asia. At maturity it is corky to punky.

The old genus Fomes is now mostly broken up into several genera. Its
chief characters were the corky to woody texture of the pileus and the
perennial nature of the spore fruit, so that in successive growing seasons
new layers of pores are formed below the ones last formed. The genus
Fomes in the restricted sense has a corky to punky pilear trama which is
more or less rusty brown in color. The spore fruits are somewhat hoof-
shaped. F. fomentarius (L.) Gill, forms its sporophores on standing trees.
They are provided with a hard crust, black and shining with age, and
are 8 to 10 cm. in width laterally, 7 to 9 cm. from front to back, and
3 to 10 cm. tall, depending upon age. These were formerly used as a
source of tinder for kindling fires and are often still called "punks."
Fomitopsis differs in having its pilear trama whitish or light-colored
(not rust brown). F. officinalis (Vill.) B. & S. grows on larch (Larix) in
the Northern Hemisphere. Its chalky white, intensely bitter, friable trama
has been used for medicine for many centuries. The large spore fruits
may become almost cylindrical, about 15 cm. in diameter and up to

POLTPORACEAE

493

Fig. 164. Polyporales, Family Polyporaceae. Ganoderma applanatum (Pers. ex Fr.)
Pat. Vertical section through three-year-old sporophore. (Courtesy, Buller: Researches
on Fungi, London, Longmans, Green and Co.)

30 or more cm. in height. Phellinus igniarius (L. ex Fr.) Quel, resembles
somewhat Fomes fomentarius but lacks the horny crust and is pubescent
when young. The hymenium of the pores has numerous sharp brown
spines. Ganoderma differs from the preceding genera in possessing spores
truncated at one end and two-layered, the brown endospore being spiny,
the spines projecting up into the hyaline exospore. The surface of the
spore fruit has a hard crust formed by a palisade of thick-walled elongated
cells. A stipe may be present in some species. A varnish-like coating may
be present over the whole surface or only on the stipe or may be entirely
lacking. In the narrow use of this name only those with the varnished
layer would properly belong to the genus. G. lucidum (Leyss. ex Fr.)
Karst. occurs on coniferous and deciduous trees and is annual. It is
varnished over the whole top surface as well as the stipe. G. curtisii
(Berk.) Murr. is perennial and may produce several layers of pores.
It loses its laccate covering early. G. applanatum (Pers. ex Fr.) Pat. lacks
the laccate surface entirely but has a whitish to gray crust. It is one of
the commonest species in North America, being found on fallen trees
and old stumps of deciduous species almost everywhere. Its spore fruits
may attain a diameter of 75 cm. White (1920) estimated that a large
spore fruit of this species may liberate 30 billion spores a day for several
months attaining a total of 5500 billion spores for the season. Yet of all
this vast number of spores carried far and wide by the wind all but a very
few must perish. With so many spores in the air it is not to be wondered
at that the fungus is very common wherever a deciduous tree has died
or its trunk has fallen. This fungus does not attack healthy uninjured
trees. (Figs. 163, 164.)

Probably closely related to the genera grouped about the genus

494 CLASS BASIDIOMYCETEAE

Trametes are those forming the Tribe Daedaleae (of Bondarzew and
Singer, 1941). In these the pores are elongated radially or are labyrinthi-
form. Daedalea is distinguished by having the pores elongated or laby-
rinthiform. Its spore fruits are shelf-like and corky. D. confragosa (Bolt.)
Fr. is very common in North America and Europe. In Lenzites the pores
are elongated radially from the point of attachment so as to resemble
gills, with occasional cross connections which may disappear with age.
The spore fruit is more or less corky. L. hetulinus (L.) Fr. is common on
birch and other trees in North America and northern Eurasia. Its fruiting
bodies are 3 to 7 cm. broad, 3 to 10 mm. thick, velvety and zonate above.
But for the cross connections in the young specimens this might well be
placed in the Family Agaricaceae. Donk (1933) and some others follow
Schroeter in dividing the genus Daedalea by separating off D. confragosa
to form the genus Daedaleopsis. Karsten (1882) separated off from
Lenzites the species with rust-brown trama as the genus Gloeophyllum.
This would include the common Lenzites saepiaria (Wulf.) Fr., which
causes the decay mainly of coniferous wood. The species of this tribe are
very variable as to pore form. In the same species some specimens may
have poroid, labyrinthiform, lamelloid, or even irpiciform hymenophores.
The distinctions between some species of Trametes, Daedalea, and Lenzites
are therefore rather arbitrary.

Undoubtedly belonging to the Polyporaceae are the species of Irpex
(Irpiciporus of Murrill) and probably the genus Echinodontium. Irpex
may be resupinate, effused-refiexed, or shelf-like. The younger parts of
the hymenophore are poroid but with increasing age the walls of the
pores grow unevenly so as to produce flattened teeth. Thus the specimen
comes to resemble closely some of the species of the old genus Polystictus.
Echinodontium has woody, shelf-like, or unguliform fruiting bodies re-
sembling some of the forms of the genus Fomes but with the hymenophore
composed of irpiciform plates which have small lateral teeth along their
edges. The spore fruits are brightly colored and were formerly used by
the Indians of Northwestern United States as a source of a red dye. It
causes decay of the hemlock {Tsuga) and of Fir {Abies) in Alaska and
northwestern United States. Both Irpex and Echinodontium were formerly
placed in the Hydnaceae.

The course of evolution in this family is very uncertain. Some of the
Pon'a-like forms may be primitive but as very many of the normally
pileato or shelf-like genera may become resupinate this latter habit
cannot always be considered to be a proof of primitiveness. Some of the
Poria group may have arisen from Thck^phoraceae that were more or
less Corticium-like. It may be possible that from the Hydnaceae, by
union of the teeth into pores some Polyporaceae may have developed.
Probably the stratose species like Fomes have developed from annual

ORDER AGARICALES 495

species. The labyrinthiform or lamelloid species like Daedalea and Lenzites
may show relationship to the Agaricaceae but they may represent merely
a parallel course of development. Some of the genera close to Polyporus
have gills radially elongated.

Order Agaricales. Largely fleshy, but some leathery or even corky or
woody at maturity. Basidia usually chiastic (after removal of Cantharellus
to the forgoing order). Hymenium on lamellae (gills) or pores (Boleta-
ceae), gymnocarpic or pseudoangiocarpic or angiocarpic. Spore fruits
mostly stipitate, more often centrally, but sometimes attached laterally
or even resupinately, without stipe. Hymenium usually not formed on the
edges of the pores or lamellae. The distinction between Polyporales and
Agaricales is not always sharp and it is possible that these intergrade so
completely that the two groups cannot justifiably remain as distinct
orders.

Family Boletaceae. This family consists of fungi growing on the
ground almost exclusively. The spore fruits are fleshy and stipitate, cen-
trally so in most species. The pileus is thick and convex and the layer of
pores is mostly easily separable from it. The pores are usually easily
broken apart from each other. In the genus Ixechinus, described by Heim
(1939) from Madagascar, the trama separating the young pores splits so
that at maturity they are separate and become divergent with the up-
rolling of the pileus. Thus they have a superficial resemblance to Fistulina,
but the origin of the separate pores is entirely different. In some genera a
veil covers the layer of pores in the young spore fruits, extending from the
edge of the pileus to the stipe. The pores in some genera are elongated
somewhat in a radial direction, suggesting a transition to or from the
Agaricaceae.

Some species form large spore fruits. Heim (1936) described Boletus
(Phlehopus) colossus from Madagascar with a pileus up to 60 cm. broad
and 4 to 6 cm. thick, and with a stipe up to 25 cm. tall and 22 cm. thick in
the lower swollen basal portion. The whole fungus weighed 6 kg. The flesh
of the various species in the family may be mainly white or pink or yellow.
In many species it becomes blue or blue-green when bruised, in others re-
maining unchanged. Treatment with KOH, NH4OH, and other chemicals
brings about various color changes which are of value in the distinction of
species and genera. The spores vary from pale to yellowish to purplish or
yellow-brown and may be small or above 20/x in length. They may be thin-
walled and smooth or may have external ridges, warts or reticulations. In
some cases the endospore is covered with prickles which reach into or
through the exospore. Germ pores are produced in a few species. Some
species are edible and some are known to be poisonous. Boletus edulis Bull,
ex Fr., according to Mez (in a verbal communication to the author), con-
tains a toxalbumin that when injected into the blood stream is extremely

496 CLASS BASIDIOMYCETEAE

poisonous but whose toxic character is destroyed in the process of diges-
tion when the fungus is eaten.

The spore fruits of various members of the family have been studied as
to their ontogeny by Kiihner (1927), Elrod and Snell (1940), and others.
In some cases they are entirely gymnocarpic, i.e., as they develop they are
more or less cylindrical and then at the top the hyphae spread outward to
form the pileus. The basidia begin to appear and to bear spores along the
stipe, especially the upper portion, and the under side of the pileus, before
the pores begin to develop. They may even be formed on the upper surface
of the pileus. The enlarging pileus curves downward at the margin as the
pores develop but in the truly gymnocarpic forms never curves inward so
far as to come into contact with the stipe. In the pseudoangiocarpic
species the development is like the foregoing except that the edge of the
pileus eventually comes into contact with the stipe, forming a circular en-
closed chamber lined above by the developing pores and centrally by the
stipe. The marginal tissues of the pileus and the portion of the stipe with
which they join may enlarge as the pileus grows in diameter and flattens
out, so as to produce an annulus which in Paragyrodon sphaerosporus (Pk.)
Sing. {Boletus sphaerosporus Pk.) spreads from near the base of the stipe
to the margin of the mature pileus as a grayish-white sheet up to 6 or more
cm. broad, leaving a chamber between it and the pore layer. Whether an
annulus is formed or not the enlargement and flattening out of the pileus
eventually exposes the pores to the air so that the spores may be carried
away by air currents. It is doubtful whether true angiocarpy occurs in this
family, i.e., development of the pores in a cavity formed internally in the
spore fruit and not by the curving downward and inward of the pileus
margin. In the genus Gastroboletus the surface layer of the pileus and
stipe remain connected and include the young hymenophore like a perid-
ium. Here, as in Paragyrodon sphaerosporus, the development is probably
pseudoangiocarpic.

The several hundred species here included in one family were divided
by Singer (1936) into two famihes, Boletaceae and Strobilomycetaceae
with a total of 21 genera, to which (1945-1947) he added three other
families, Gomphidiaceae, Paxillaceae and Jugasporaceae, usually placed
in the old family Agaricaceae. These five families then form his Suborder
Boletineae. Coker and Beers (1943), on the other hand, recognize only
three genera of Boletaceae in the usual sense, in North Carolina. Murrill
(1910) recognizes 11 genera in North America.

It has long been recognized that the genus Paxillus of the old family
Agaricaceae has many points of similarity to the Boletaceae: frequent oc-
currence of ti-ansverse ridges between the lamellae, ease of separation of
the lamellae from the pilear trama, gymnocarpic development of the spore
fruit and certain cytological and chemical similarities. Whether these

ORDER AGARICALES 497

justify transferring Paxillus to the Boletineae, as Singer does, or indicate
the close relationship of the latter group to the Agaricaceae is more a
matter of opinion. In this hookPaxillus will be retained in the Agaricaceae.

The Boletaceae are found in the temperate, subtropical and tropical
regions of both hemispheres, but especially where the rainfall is fairly
abundant or in the season of the year when considerable rain falls. They
do not occur in arid regions. Many, perhaps most, of them occur in con-
nection with mycorrhizal development on roots of mostly woody plants.
Some are confined to the roots of conifers and some of this group are
limited to certain genera (e.g., Larix, Pinus, etc.). Gyrodon merulioides
(Schw.) Sing. {Boletinus porosus (Berk.) Pk.) is known only in proximity
to trees of ash {Fraxinus) and Paragyrodon sphaerosporus (Pk.) Sing.
{Boletus sphaerosporus Pk.) only near species of oak (Quercus), etc.

Among the larger and commoner species several may be mentioned.
Boletus edulis Bull, ex Fr. has a reddish brown pileus, white or yellowish
within, the flesh not becoming blue upon wounding. The pores are yellow-
ish and become greenish with age. The stipe is reticulately marked. The
pileus is 6 to 20 cm. broad and 2 to 4 cm. thick and the stipe cylindrical or
enlarged below and 5 to 10 cm. tall and 3 to 4 cm. thick. The spores are
yellowish to ochraceous brown. The flesh has a pleasant nutty taste. This
highly prized edible species occurs in frondose woods. The species related
to Boletus luridus Schaeff. ex Fr. (Suillellus luridus (Schaeff.) Murr.) are
often about the same size as the foregoing and the pileus has much the same
appearance above. The flesh is whitish or yellowish but becomes blue very
rapidly when exposed to the air. The pores are yellowish with red mouths.
The stipes are 5 to 10 cm. tall and 1 to 2 cm. thick, reddish below, yellow
above, reticulated near the top. This species is reputed to be poisonous.
Tylopilus felleus (Bull, ex Fr.) Karst. (Boletus felleus), also may be con-
fused by a beginner with both the foregoing. It differs in its pink spores,
its white flesh, which may turn pink on wounding, and its intensely bitter
taste. The pores, which are white, become flesh colored as the spores are
produced in large numbers. The stipe may be reticulate above or com-
pletely reticulate. All three species may be found in the same woods.
Suillus luteus (L. ex Fr.) S. F. Gray grows in the vicinity of species of pine.
The yellowish to reddish brown pileus is very viscid. The flesh is pale yellow-
ish, not changing color when wounded. The stipe is pale yellow to reddish
brown and glandular dotted, and has a large persistent annulus. It is edible.
Strobilomyces floccopus (Vahl ex Fr.) Karst. (S. strobilaceus (Scop, ex Fr.)
Berk.) has dark spores, completely covered by a network and with a dis-
tinct germ pore. The pileus and stipe are gray when young but the numer-
ous shaggy scales quickly become dark. The gray to white pores become
reddish or black on wounding or bruising, and are more or less lamellar
near the stipe, which has an annulus. This edible species grows in frondose

498

CLASS BASIDIOMYCETEAE

,f*-

or mixed frondose and coniferous woods. Gyrodon merulioides has an ec-
centric or lateral stem and the pores are formed by radiating lamellae
connected by numerous cross veins not quite so high as the main lamellae.
The pileus is reddish brown, with yellow flesh slowly turning bluish green
when wounded. The hymenial surfaces are yellow becoming slightly blue
on wounding. The spores are yellowish brown. The pores are decurrent

somewhat on the hollow stipe. This
approaches closely some of the species
of Paxillus. (Fig. 165.)

Family Agaricaceae. This
family in its broader and more cus-
tomary usage included those fungi
whose fruit bodies increased the
hymenial surface by the production
of radiating lamellae which are
entirely covered, or all but the edge,
by the hymenium. The latter may or
may not extend from gill to gill on
the interlamellar surface of the
pileus. The interior tissue of the
lamella (the trama) may continue
unchanged up into the pileus or the
pilear trama may be distinct in
structure, €olor, etc., from the
lamellar trama, paralleling the con-
ditions in the Polyporaceae.
In contrast with the Polyporaceae where the spore fruits are prevail-
ingly rather dry at maturity those of the Agaricaceae are mostly fleshy,
although some dry forms occur. In the vast majority of cases they are
centrally stipitate, rarely laterally so, occasionally attached laterally with-
out a stipe, or even partially resupinate. In size the pileus may vary from
a very few millimeters in diameter in some species of Marasmius to 40 cm.
in specimens of an exannulate form of Agaricus arvensis Schaeff. ex Fr.,
collected by the author. A specimen of this size must be capable of pro-
ducing an enormous number of spores since Buller (1909) has shown that
a not unusually large specimen of A. campeslris L. ex Fr. can produce 1800
million spores.

In general the basidia are club-shaped, varying to ovoid or cylindrical.
Usually four l)asidiospores are produced although species or races fre-
quently occur in which the number is two. In the latter case this may
result from the development of spore fruits on monocaryon mycelium and
the consequent lack of nuclear fusion and meiotic divisions in the basid-
ium, there being only one division and that mitotic. The spore fruit may

Fig. 165. Agaricales, Family Bole-
taceae. Tylopilus felleus (Bull, ex Fr.)
Karst. (Courtesy, Atkinson: Studies of
American Fungi, Ithaca, N. Y., Andrus
and Church.)

ORDER AGARICALES 499

arise from a dicaryon mycelium with normal nuclear phenomena in the
basidium and yet produce only two spores on the basidium with two
nuclei passing into each. In some members of the family, as occurs fre-
quently on other Eubasidiae a mitotic division subsequent to the second
meiotic division results in the production of eight nuclei in the basidium
of which four may enter into the basidiospores and four remain behind, or
two proceed into each spore. In a stained specimen of Coprinus sp. studied
by the author each basidium possessed eight nuclei but after spore dis-
charge four still remained in the basidium. With the exclusion of Cantha-
rellus all Agaricaceae are chiastobasidial.

The hymenium may consist entirely of basidia or there may be cystidia
of various types. In Coprinus the basidia are separated by large sterile
cells called paraphyses, not so tall as the basidia but much broader so that
the latter are arranged in squares, one at each corner where four para-
physes meet. Such paraphyses are not produced in most of the genera of
the family. The cystidia may differ in appearance from the basidia only in
the absence of sterigmata or they may be elongated, pointed, or forked, or
knobbed so as to resemble a tenpin. The upper portion of the cystidium is
often covered with crystals. Some cystidia are the terminal cells of laticif-
erous hyphae or of hyphae containing mucilaginous substances. The
cystidia occurring at the edge of the lamellae are called cheilocystidia and
may resemble or differ from those borne on the faces of the gills, the
pleurocystidia. Where, as in some species of Coprinus, the large, stout
pleurocystidia extend across the space between two gills and assist in
holding them apart they are sometimes called trabecular cystidia.
(Fig. 166.)

Fayod (1889) distinguished several types of gills by their tramal struc-
tures. Other mycologists since then have recognized the value of these
distinctions in the classification of the Agaricaceae. The outer surface
always consists of the vertically standing basidia (and cystidia if present)
arising from a subhymenium, a thin or thick layer immediately below the
basidia. This may be indistinguishable. In the mixed or irregular trama
the hyphae are arranged without apparent order, being sinuous or inter-
laced. The regular trama consists of elements clearly parallel. In the
bilateral trama the trama proper is reduced to a thin median plane from
which the hyphae diverge obliquely in a curved line toward the strongly
developed subhymenium. In the inverse trama the young lamella has the
"regular" structure with a distinct subhymenium. As maturity ap-
proaches hyphae grow from the subhymenium obliquely inward, filling
the space formerly occupied by the vanished median portion of the trama.
It must be recognized that these four types grade into one another and
that at times it is difficult to decide which type is present, especially if
the gill that is being studied is a little too young or too old.

500

CLASS BASIDIOMTCETEAE

Fig. 166. Agaricales, Family Agaricaceae. Mechanism in Coprinus for holding gills
apart to permit falling of the spores. (A) Coprinus atramentarius Fr. with gills held
apart by trabecular cystidia; autolysis beginning at edge. (B) C. slerquilinus Fr. with
thickened edges that hold the gills apart. Note the long and short basidia. (Courtesy,
Buller: Researches on Fungi, London, Longmans, Green and Co.)

ORDER AGARICALKS 501

Many Agaricaceae show clamp connections at the septa of almost all
the hyphae of the lamellar and pilear trama, the cortex of the pileus, and
the tissues of the stipe as well as the mycelium from which the spore fruit
arose. In many cases the clamp connections occur only at occasional septa
and only in special portions of the pileus or stipe, often only in the cortical
regions. In Russula and Lactarius groups of spherical cells (sphaerocysts)
are found in the pileus and even in the lamellae in addition to the usual
slender hyphae. The pilear trama of other genera may show enlargements
of the hyphae but not so characteristic as those of the genera mentioned.
Slender hyphae bearing clamp connections frequently do not develop
these structures where cells much larger in diameter are developed as they
grow. In the genus Lactarius the tissues of the pileus, stipe and gills
possess branching hyphal tubes filled with latex which flows out of breaks
in the tissue and coagulates. The fresh latex may be white, pink, yellow,
green, blue, or even colorless. It may change color on exposure to the air
or remain unchanged. Aside from the protection to wounds afforded by
the coagulated latex it may be that the laticiferous tubes serve for food
transportation. Besides this genus laticiferous vessels are reported by
Heim (1936b) in three other genera of Agaricaceae from Madagascar,
Bertrandia, Mycena, and Rhodophyllus, as well as in some species of
Gasteromycetes.

The species of Agaricaceae may be " homothallic " or " heterothallic "
and bipolar or quadripolar. Most genera of the Agaricaceae have not been
grown in culture so that it is not known how extensively the appearance of
two or more sexual phases is to be found. The phenomena of sexuality
have been discussed rather fully in the preceding chapters. Oidia are pro-
duced in abundance in monocaryon cultures but are usually absent from
dicaryon mycelium. Where they do occur, as Vandendries and Martens
(1932) described for Pholiota aurivella, they may be binucleate and give
rise to dicaryon mycelium or divide into two uninucleate cells which give
rise to monocaryon mycelium.

The Agaricaceae are mainly saprophytic, living on the ground, decay-
ing leaves, bark, wood, manure, etc. Some species of Nyctalis and Volvaria
are parasitic on other Agaricaceae. Armillariella mellea (Vahl) Karst.
{Armillaria mellea) attacks the roots of trees and kills their cortical tissue,
growing up in the living portions of the bark of the tree trunk and causing
the death of the tree. It is especially destructive to apple and cherry trees
planted where previously there were oaks or other trees on whose roots
this fungus once grew parasitically, continuing to live as a saprophyte until
the apple or cherry roots became available. Agaricus campestris L. ex Fr.
and one or two other species are cultivated for food and many wild species
are collected for this purpose. Buller (1922) pointed out that Marasmius

502

CLASS BASIDIOMYCETEAE

Fig. 167. Agaricales, Family Agaricaceae. Armillariella mellea (Vahl) Karst. (Courtesy,

Dow V. Baxter.)

oreades (Bolt, ex Fr.) Fr. when grown in manure produces large amounts
of edible fungi much larger than the wild form. (Fig. 167.)

Among the edible wild species are various kinds of Lepiota, e.g., L.
naucina (Fr.) Quel., L. procera (Scop, ex Fr.) Quel. {Leucocoprinus pro-
cerus (Scop, ex Fr.) Pat.). It must be noted that some species of this genus
are poisonous. Most of the larger species of Agaricus (PsaUiota of some
authors) are edible, e.g., A. campestris L. ex Fr., A. rodmani Pk., A.
arvensis Schaeff. ex Fr., etc. Pluteus cervinus (Schaeff. ex Fr.) Qu61.,
Coprinus comatus Fr., C. micaceus Fr., C. atramentarius Fr. (but see
under poisonous species below), Marasmius oreades, Ladarius volemus
Fr., L. deliciosus (L.) Fr., Pleurotus ostreatus (Jacq. ex Fr.) Quel.,
P. ulmarius (Bull, ex Fr.) Qu61., CollyUa radicata (Rehl.) Berk., and
various other species of CoUyhia, Tricholoma personatum (Fr.) Qu^L,
Armillariella mellea, and many others are edible. Of the genus Amanita,
A. rubcscens (Pers. ex Fr.) Fr. and A. caesarea Schaeff. ex Fr. are highly
prized, but see note concerning this genus among the poisonous species
below. Aside from the hundreds of edible species of mushrooms there are
many that are poisonous, some mildly so, some dangerously. Besides these
edible and poisonous sorts the larger number of species are too small or
tough or shmy in consistency, or of disagreeable odor or taste even though

ORDER AGARICALES

503

not poisonous, or occurring only occasionally here and there so as to be
too scattered for collection in sufficient numbers for a meal, so that the
majority are not usually subjects of concern as possible edible products.
Among the poisonous mushrooms several species of Amanita are ex-
ceedingly dangerous. A. phalloides (Bull.) Fr. and some of its close rela-

FiG. 168. Agaricales, Family Agaricaceae.
Amanita verna (Fr.) Quel. (Courtesy, M. B.
Walters.)

tives (e.g. A. verna (Fr.) Quel.) are so poisonous that a piece one cubic
centimeter in size will cause severe illness or even death, there being no
known antidote for the poisonous principle. A. muscaria (L.) Fr. was
formerly used for the purpose of destroying flies as the sticky cuticle is
very poisonous. In some parts of Europe and Siberia after careful removal
of the cuticle the remainder of the fungus is eaten with apparent safety.
Some tribes in Siberia make an intoxicating beverage out of the fungus.

504

CLASS BASIDIOMYCETEAE

5?i>.

.^:£W

-'Mi

/ ;'

«W

^•*

Fig. 169. Agaricales, Family Agaricaceae. Lepiota rachodes (Vitt.) Quel. (Courtesy,

M. B. Walters.)

It is wiser to avoid this species as numerous cases of poisoning and some
of death have been recorded after its ingestion. Atropin is a partial anti-
dote. Lepiota morgani Pk. {Chlorophyllumesculentum Mass., Leucocoprinus
molyhdites (Meyer) Heim) is abundant in parts of North America as well
as in the Tropics. It has considerable similarity to the edible L. procera
but is stouter and has green spores, the gills taking on a greenish tinge as
the spores develop. This species is quite poisonous for some persons but
may be eaten with apparent impunity by others. L. helveola Bres. is also
poisonous, sometimes very dangerously so (Josserand, 1931). ^lany of the
pink-spored fungi {Entoloma or Rhodophyllus of some authors) are poison-
ous although the edible Pluteus cervinus has pink spores and is safe.
Several species of Tricholoma are unsafe. Recently a case of slight poison-
ing occurred with fresh specimens of a species of Agaricus where other
persons eating specimens of the same lot were not affected. (Figs. 168,

169.)

The general conclusion to be drawn from the foregoing is that because
some species of a genus are safe this does not necessarily give a clean bill
of health to other species of the same genus. Besides that there are per-
sonal susceptibilities to the poisonous characteristics of some fungi. It has
been shown (by Suss, 193(), and Hugon, 1938) that Coprinus comatus and

ORDER AGARICALES

505

C. atramentarius may cause quite severe poisoning if alcoholic beverages
(beer or wine, as well as distilled liquors) have been drunk shortly before
or within several hours after eating the fungi. The method of preparation
may have considerable effect upon the poisonous nature of some species.
Parboiling and pouring off the water will often remove the poison but not
in all cases. Of course almost all fungi which have reached the state of
incipient decay are dangerous. This is the basis of the "silver spoon" test,
for the blackening of the silver surface is not due to the poison naturally
resident in the fungus but to the products of incipient decay. It is unsafe
to eat a fungus unless its identity is certain and then only perfectly fresh
specimens or dried unspoiled specimens.

Botanically it is insisted that the words mushroom and toadstool
are practically synonymous and may be used indiscriminately for both
edible and poisonous species. It must be recognized, however, that orig-
inally the word toadstool was derived from a word meaning death's chair
(Todesstuhl).

The poisonous principles in the Agaricaceae appear to belong to the
chemical groups of "toxalbumins" in some cases and alkaloids in others.

Studies of the development of the spore fruits of centrally stipitate
members of this family show that with reference to the hymenial origin
they may be gymnocarpic, pseudoangiocarpic or truly angiocarpic. In the
first the hymenium is external from the beginning and never enclosed in a
cavity. At the upper part of the young columnar spore fruit the tissues
spread out laterally, to form the young pileus. On the under side of this
and often on the upper part of the stipe the hymenium begins to develop,
gradually producing radial folds, the gills. At maturity these bear the
mature basidia and basidiospores and are at no time cut off from the out-
side air by tissue of any kind. (Fig. 170 A-C.) In the pseudoangiocarpic
species development is as above at first, but the broadening pileus curves
downward at the edge and finally curves back to the stipe with which it
comes into loose contact or to which it becomes united by the intermin-
ghng of hyphae from the stipe and edge of the pileus. Thus a closed circular
cavity develops on whose roof the lamellae are produced and become
covered with the hymenium. At the approach of maturity the pileus
flattens out and its edge breaks free from the stipe, so that now the
hymenium with its ripening spores is exposed to the air. There may be
left on the stipe a collar (annulus) or the sheet of connecting tissues may
break away from the stipe and remain hanging at the edge of the pileus as
a cortina. This may be in broken sheets or like a spider web. (Fig. 170
D-G.)

In the angiocarpic forms there develops within the tissues of the pileus
a circular layer of palisade cells, the hymenium primordium, on the lower
or inner side of which a circular cavity is formed into which the radiating

506

CLASS BASIDIOMYCETEAE

p5»!55«?-«.-j!.a%-v:«s;»!^«(^agp M^^

Fig. 170. Family Agaricaceae. Methods of development of tlie hymenophore.
(A-C) Gymnocarpic type, in Omphalia chrysophylla Fr. (D-G) Pseudoangiocarpic
type, in Lentinus tigrinus (Bull.) Fr. (H, I) Angiocarpic type, in Agaricus campestris L.
ex Fr. (A-C, courtesy. Blizzard: Am. J. Botany, 4(4): 221-240. D-G, courtesy,
Kiihner: Compt. rend., 181(3): 137-139; H-I, courtesy, Atkinson: Botan. Gaz., 42(4):
241-264, Univ. Chicago Press.)

ORDER AGARICALES 507

gills protrude as they develop. According to Levine (1922) the early ap-
pearance of the annular cavity is an artefact and the openings arise first
between the developing gills and only later below their edges if at all. In
Coprinus the gills remain in contact with the upper part of the stipe until
the spore fruit is nearly mature and the pileus begins to expand. Eventu-
ally this circular cavity is bounded on the inner side by the portion of the
spore fruit that becomes the stipe, and by the flattening of the pileus a
circular break next to the stipe permits the pilear expansion so that the
gills now become exposed to the air and shed their spores. (Fig. 170 H-I.)
In these forms the surfaces destined to bear the hymenium arise within
the pileus while in the other two types it was at first exposed to the air
and became ultimately enclosed only in the pseudoangiocarpic type of
development. For details as to gymnocarpic and pseudoangiocarpic de-
velopment consult Kiihner (1925b, 1926), Blizzard (1917), Douglas
(1918), Walker (1919), Reijnders (1933), Heim (1936b, 1937), and others,
and for the angiocarpic type Atkinson (1906, 1914, 1915, 1916), Levine
(1922), Douglas (1916, 1920), and some of the foregoing hst.

The 5000 to 8000 species of the Agaricaceae are divided into 50 to 112
genera, according to the ideas of the various students of the group. Fries
(1821) recognized only two genera in the limits of the family as here
treated, Agaricus and Schizophyllum. Killermann (1928) recognized 66 in
the second edition of Engler and Prantl. Singer (1936) included about 112
genera. More often these are all included in only one family but more
recently Heim, Singer, and others have divided them into 10 or 11 fam-
ilies. It will probably be desirable to make a segregation, but until there is
greater agreement as to just what the generic distinctions must be and
how the families should be delimited the author will be conservative and
include all in one family.

Fries divided his genus Agaricus into 38 tribes, most of which were
later recognized as genera by him and by other mycologists. The primary
basis of the division into two genera was a sphtting of the lamellae in
Schizophyllum and the entire lamellae in Agaricus. The latter was divided
into series on the basis of spore color : white, rose, ochraceous, rusty, purple-
brown, and black. In the "Sylloge Fungorum" (Saccardo, 1887) these
primary subdivisions were called respectively Leucosporae, Rhodosporae,
Ochrosporae (including both ochraceous and rust-colored spores), Melan-
osporae (including both purple-brown and black spores). The location of
the stipe (central, eccentric, or lateral), the presence or absence of annulus
and volva, the shape of the pileus, the character of the stipe, the relation
of lamellae to stipe (i.e., free, attached, decurrent, etc.), the color of the
gills and of the pileus were all characters used in dividing these spore-color
groups into the lesser groups (tribes or genera) . All these characters were
largely external and did not take into consideration the internal structures

508 CLASS BASIDIOMYCETEAE

such as have been found of so great importance in the Polyporaceae.
Fayod (1889) and others (Karsten, Heim, Singer, etc.) have emphasized
more and more the necessity of basing the generic distinctions upon these
anatomical and chemical characters. The type of gill structure as empha-
sized by Fayod, the amyloid or nonamyloid character of the spore wall,
the structure of the cuticle of the pileus, the presence or absence of clamp
connections, all these have proved to be of great importance. Just how far
these characters should be used in distinguishing genera or whether they
should only be used for lesser units of classification is not yet agreed upon,
hence the above-noted great differences in the number of recognized
genera or even families.

The characters of size — of spores, cystidia, pilear hyphae, etc., as well
as of the whole spore fruit — are not always reliable. Thus Togashi and
Oda (1934) showed that the age of the pileus has considerable effect upon
the spore size. In Armillariella mellea and in Pholiota adiposa (Fr.) Quel,
the length and width of the spores shed on the fourth day were 12 to 15
per cent less than of the spores shed on the first day from the same sporo-
phores. On the other hand Collyhia velutipes (Curt.) Fr. {Myxocollyhia)
does not exhibit any such change in spore size with the age of the pileus.
The size and even the shape and color of the pileus, and sometimes of the
gills, depends considerably upon the temperature, humidity, and illumina-
tion, and to a large degree upon the amount of food available in the sub-
stratum. The position of the stipe, whether central, eccentric, or lateral,
may vary in the different spore fruits growing from the same tree or log,
depending upon the position of the point of exit from the host. If this is
from the top it may be centrally stipitate and from the side the stipe may
be eccentric or lateral.

In general the spores may represent four rather distinct types: (a)
With an endospore and epispore, often with a distinct germ pore. Such
spores are mostly ellipsoidal or short cyhndrical, or oval, and are often
somewhat truncate at one end. (b) With only one spore wall layer visible,
usually subspherical, oval, elhpsoidal or cyhndrical. (c) Very strongly
angular or with large knobs. Spores of this type are mostly rose-colored to
pink, (d) More or less ellipsoidal or oval or cylindrical with mostly six or
eight longitudinal grooves so that in cross section they are not round but
angular or lobed. These spores also are more often rose or pink. The outer
surface of the spores of all types may be smooth (more often the case) or
somewhat roughened or tuberculate or reticulately marked by ridges.
These tubercles or ridges are stained blue by iodine reagents in amyloid
spores such as occur in species of Russula and Ladarius.

The angiocarpic spore fruits such as those of Amanita show an external
universal veil or cortex before they open. Upon completion of the expan-
sion the portion of this veil on the upper surface of the pileus may be

ORDER AGARICALES 509

visible as patches of tissue, as in A. muscaria, where the veil does not keep
pace with the remainder of the expanding pileiis and so is torn to pieces.
On the other hand if it does keep pace in its growth it is detectable as a
thin fibrous layer or cuticle of some other type. Around the foot of the
stipe the universal veil is left as a loosely or closely adhering cup, the
volva. In angiocarpic forms like Agaricus there is no volva, as the rupture
is so low doAvn that no free basal portion of the veil is distinguishable. In
angiocarpic forms with an annulus or secondary veil this arises as a sheet
of tissue below the circular hymenial cavity, either separating this cavity
from the outside air below or, if attached high on the stipe, sheathing the
upper part of it. The annulus may be simple (as in Agaricus campestris) or
of two distinct layers (as in A. arvensis). It may break free from the edge
of the pileus and also from the stipe to form a movable ring (as in Lepiota
procera) or at one or the other place, leaving a ring attached to the stipe
or broken sheets or threads attached to the edge of the pileus (a cortina).
In pseudoangiocarpic forms the annulus, as in the Boletaceae, may be
composed of tissues from the stipe, that gi-ew out and attached themselves
to the incurving pileus margin or of tissues from the edge of the pileus that
became attached to the surface of the stipe. The truly gymnocarpic forms
show neither annulus nor volva nor patches or cuticular structure on the
pileus that represent in any manner the remnants of a universal veil.

Reijnders (1933) assembled from literature and from his own investi-
gations a list of 79 species of Agaricaceae of which the mode of develop-
ment of one is doubtful, 21 gymnocarpic, 2 pseudoangiocarpic and 55
angiocarpic. Heim (1937) added two pseudoangiocarpic species to this
list. It must be noted that some writers list the latter type of development
as gymnocarpic so that probably there should be in Reijnders' hst a
higher proportion of species with pseudoangiocarpic type of development.
All of the Ochrosporae and Melanosporae among the 81 species concerned
are angiocarpic ; of the four Rhodosporae listed one is pseudoangiocarpic
and three gymnocarpic. Of the 41 Leucosporae 19 are angiocarpic, 19
gymnocarpic, 2 pseudoangiocarpic and 1 doubtful. If the angiocarpic
species, following Singer (1936) and others, are the more primitive, arising
from the Gasteromycetes, it is apparent that the Agaricaceae with colored
spores are the more primitive and those with white or pink spores further
from the ancestral forms. If the light-spored forms are considered the
more primitive then the angiocarpic genera in general are further ad-
vanced in evolution. The peculiar type of hymenium of Coprinus and of
some species of Pscudocoprinus, with broad paraphyses and the more
slender basidia arranged in squares at the corners is not found except in
these and perhaps a few other black-spored species. Aside from this ar-
rangement of the basidia and paraphyses these two genera have certain
other specialized structures that appear far from primitive. In the stout.

510 CLASS BASIDIOMYCETEAE

more fleshy Coprini the broad gills are close together but not in contact
and the original downward curve of the pileus keeps them in contact with
the stipe. Thus the outer portion of the gills furthest from the center of
the pileus is situated at the bottom of the gill mass. The maturing of the
basidia and discharge of the basidiospores occur first in a narrow band at
this outer edge of the gill. The spores have only a short distance to fall to
escape into the air. Then this band from which the spores have been dis-
lodged undergoes autodigestion into a dark-colored fluid which either
dries up or drops off and the next strip of the hymenium produces its
spores. Thus the spores never have to fall far between the crowded gills.
The black inky drops are not colored by the spores and are not the means
by which the latter are distributed. In the Coprini with narrow gills, quite
widely spaced, also in Pseudocoprinus, the tissue of the pileus is very thin.
It spHts radially over the median plane of each gill which spreads out in a
V-shaped cross section, thus making the fall of the spores more efficient.
Buller (1909) gave a thorough discussion of these different types of pilei.
In most of the remainder of the Agaricaceae the basidia reach maturity
not in a band but here and there all over the surface of the gills. In Pan-
aeolus, another dark spored genus, the spores mature in patches on the
gifls, giving them a more or less variegated appearance. Possibly this in-
dicates a tendency toward the habit in the fleshy Coprini.

It is worthy of note that Pietro Antonio Micheli published in 1729
what is probably the first key to the species of this family, which he
recognized merely as a single genus Fungus. Among the characters used
by him in his key were the clustered or separate growth of the spore
fruits, their branching or nonbranching, presence or absence of volva,
presence or absence of annulus, nature of annulus, i.e., whether free or
attached to the stipe, presence or absence of striations on the pileus, loca-
tions of striations if present, whether pileus and gills were of the same or
different colors, presence or absence of latex, etc. Although his system,
too, was largely artificial it served to distinguish the species known to
Micheli and in some particulars was no more artificial than the one more
recently employed.

In spite of the various systems of classification proposed in the last few
decades the relationships within the Hymenomycetes are still very un-
certain. The simplest forms appear to be those related to Corticium in the
Thelephoraceae. In many respects this genus is morphologically very
much like Ascocorticium of the Taphrinales. It may be that by the extru-
sion of the spores into external pockets the basidium has arisen from the
ascus and thus the gap between the Ascomyceteae and Basidiomyceteae
was bridged. On the other hand the basidium of a form like Auricularia
may represent a four-spored ascus in which the ascospores instead of
escaping germinate in situ and produce secondary spores. From this struc-

KEY TO THE MORE IMPORTANT GENERA OF FAMILY THELEPHORACEAE ' 511

ture may have arisen finally the holobasidium of Corticium. From the
latter as a starting point the greater complication of the hymenium lead-
ing to increased spore-bearing surface brings us to such famihes as Clava-
riaceae, Hydnaceae, and Polyporaceae. Within the latter there is a tend-
ency toward radially elongated pores or even lamellae. From some of the
fleshy Polypori with central stems may have arisen forms like the Bole-
taceae. Some of the latter have pores that are semilamellate. So there is a
possibility of the origin of the Agaricaceae from either of these families.
On the other hand there is an undeniable close relationship between a
number of the Agaricaceae and several genera of the Gasteromycetes.
Singer (1936) held that this indicates a descent of the former from the
latter. In this case the universal veil would represent a reduced peridium
and evolution within the Agaricaceae would have to be considered as
progressing from the angiocarpic through the pseudoangiocarpic to the
gymnocarpic forms. The chief objection to this theory is that it would
involve the transformation of the spore attachment from a symmetrical
one to the obliquely perched spore of the Agaricaceae. Inasmuch as the
theory further presupposes that the Gasteromycetes were evolved from
simple forms with obliquely attached spores this would involve the muta-
tion back again to the latter condition with an ancestral interphase of
symmetrical spore attachment. The further phylogenetic considerations
of these groups will be given more in detail in Chapter 17.

Key to the Genera of Family Exobasidiaceae

Mycelium intracellular; clusters of basidia, often among elongated paraphyses,
emerging from the stomata; basidia with two sterigmata. Kordyana

Mycelium intercellular; basidia emerging from between the epidermal cells, usu-
ally four- to six-spored ; no paraphyses. Exohasidium

Key to the More Important Genera of Family Thelephoraceae

Hymenophore resupinate, floccose or continuous, of only one layer.

Hymenophore floccose or felty or pellicular; basidia in scattered clusters, mostly
not forming a continuous hymenial layer.
Fructification tenuous with scanty subiculum, the hyphae distinct, loose;
basidia subglobose or short-claviform, with two to four stout, elongate,
more or less cornute or flexuous "epibasidia"; spores smooth, germi-
nating by repetition. See Ceratobasidium

in Chapter 13.
Fructification mucedinoid, reticulate-pellicular, finely granulose; hyphae
short-celled, branching at right angles, often with formation of cruciform
cells; basidia in cymose, often candelabrum-like clusters, short cylindric
with four or six to eight sterigmata; spores mostly smooth but in a few
species spinulose; in some species spores germinating by repetition.

Pellicularia

(Botryobasidium)
Fructification felt-like or "hypochnoid," composed of loosely interwoven
hyphae; basidia in scattered clusters or sometimes in a compact, smooth

512 CLASS BASIDIOMYCETEAE

or papillar hymenium; basidiospores usually colored, rough- walled to
echinulate. Tomentella

(Hypochnus of some authors)
Hymenophore membranous or coriaceous, not reflexed or saucer-shaped:
basidia mostly forming a continuous smooth or papillose hymenium.
Basidiospores white or rarely bright-colored.

Cystidia lacking. Corticium

Cystidia present in hymenium or in subhy menial tissues or both.

Peniophora
Brown stellate organs pi-esent in subhymenial tissue.

Asterostroma
Hymenium interrupted by sterile pegs or projections.

EpitJiele
Basidiospores ochraceous, ferruginous, or fuscous, smooth.
Cystidia lacking. Coniophora

With cystidia. Coniophorella

Hymenophore leathery or corky, saucer-shaped, or at least upturned at the
edges; basidiospores pale-colored, usually large; cystidia of various
types. Aleurodiscits

Resembling the foregoing but with groups of dendrophyses (feather-like
cystidia) forming projections above the hymenium.

Dendrothele
Saucer or cup-shaped, more or less gelatinous-fleshy. Cytidia
Hymenophore adhering closely to the substratum; antler-like cystidia forming
a felty layer above the basidia. Vararia

(Asterostromella)
Hymenophore of three layers, resupinate or more often reflexed-effuse or attached
laterally.
Leathery, without cystidia. Stereum

Leathery, sometimes almost woody or corky, variable in shape; with stiff,
brown, pointed setae extending from the hymenium.

Hymenochaete
Hymenophore upright, often stalked, simple or branched or funnel-shaped, or
pendent cup-shaped or separate tubes.
Hymenium Hning the inner surface of pendent cups or tubes.
Sessile or stalked cups; separate. CypheUa

Tube-like arising together, but not united, from a common subiculum.

Solenia
Hymenium lining the outer or lower surface of funnels or flattened lobes.
Leathery; spores mostly l^i'own and roughened.

Hymenium almost smooth or warty. Thelephora

Hymenium with woody ribs; tropical, Cladoderris

Other uncommon genera. Skepperia, Hypolyssus

Fleshy, much branched, forming a round, cabbage-like structure; spores
hyaline, smooth. Spnrassis

Key to the Genera of Family Cantharellaceae

(After Smith and Morse, 1947)

Fruit body typically fleshy and (centrally stipitate or stipe eccentric, but well-
developed (or whole fruit body funnel-shaped and stipe not distinct).
Hymenium smooth or nearly so. Craterellus

KEY TO THE GENERA OF FAMILY CLAVARIACEAE 513

Hymenium in the form of radiating, sometimes reticulate, ridges or folds or as
obtuse, forked lamellae. Cantharellus (Gomphus)

Fruit body very thin and delicate, stipe small or rudimentary.
Spores hyaUne. Leptotus (Dichjolus)

Spores tawny to incarnate. Arrhenia

Key to the Genera of Family Clavariaceae

(Based on Doty, 1948)

Spores hyaline (rarely tinted), mostly smooth and thin- walled; basidia two- or
four-spored.
Fructification with broad, flattened terminal portions of the branches (probably

better in Family Thelephoraceae). Sparassis

Fructification not as above.

Spores large (7 fx or over), globose; basidia two-spored; hymenium white or
gray; not staining green with Fe2S04 solution.

Clavulina
Spores distinctly smaller or ellipsoid; basidia typically four-spored if spores
spherical; hymenium often colored otherwise or staining
green with Fe?S04 solution.
With stout hyphae in the trama producing setae or gloeocystidia in the
hymenium or subhymenium, or the apices of the branches
truncate or cup-shaped; spores under 8 ju long; not staining
green with Fe2S04 solution.
Apices of some branches truncate to cup-shaped, with gloeocystidia in

the hymenium layer. Clavicorona

Apices of branches acute; with strong setae in the hymenium or sub-
hymenium.
Setae loosely dichotomously branched (dichophyses).

Eriodadus
(Lachnocladiu m )
Setae closely dichotomously branched (asterophyses).

Stelligera
Without such hyphae or apices; spores various; or staining green with
Fe2S04 solution.
Fructifications branched.

Very finely branched (under 1 mm.) ; toughish. Pterula

Branches larger; flesh fragile or putrescent.

Hymenium staining green with Fe2'^04 solution. Clavariella
Hymenium not staining green with Fe2S04 solution.

Clavaria
Fructifications simple or rarely branched above.

Fructifications enlarged above or over 1 cm. in diameter above; spores
ellipsoid.
Minute (not over 5 mm. tall) ; or with an abruptly inflated head ; not
staining green with Fe2S04 solution.
With an inflated, down-turned head. Physalacria

With the apex merely enlarged.

Hymenium on the expanded blunt apex. Pistillina

Hymenium on the sides of the club. Pistillaria

Large (over 2 cm. tall); or turning green with Fe2S04 solution.

Clavariadelphus

514 CLASS BASIDIOMYCETEAE

Fructifications not enlarged above (i.e., slenderly clavate to filiform
clubs) ; spores various.
Fructifications over 2 cm. tall; not obviously restricted to specific
hosts or substrata; or the fructifications fascicled fleshy
forms; often globose spores.
Tramal hyphae with many secondary cross walls; clamp connec-
tions rare. Clavaria
Tramal hyphae with secondary cross walls only rarely; clamp
connections on most cross walls. Clavulinopsis
Fructifications smaller, restricted to specific hosts or substrata
which may be sclerotia or living plants; spores eUipsoid or
flattened on one side.
With a sclerotial base; stipe slender and distinct; mostly over 5

mm. tall. Typhula

Without a sclerotial base; stipe not distinct; mostly less than 2
mm. tall. PistiUaria

Spores typically ochraceous, mostly roughened or obdurate walls; basidia four-

spored.
Toughish to woody; spores echinate or sharply warty; hymenium sometimes
unilateral; not becoming green with Fe?S04 solution.'
Coarse, leathery to woody fungi; hymenium often unilateral or branches
flattened. Thelephora

(some species)
Delicate toughish to woody fungi; hymenium covering all surfaces of the
rounded branches. Scytinopogon

Fleshy; spores smooth, verrucose to echinulate; hymenium on all lateral sur-
faces of the branches; becoming green with Fe2S04 solution.
Simple, unbranched fungi with broadened sterile apices; sometimes "mush-
room-like" in form. Gomphus

{Cantharellus in part)
Branched fungi; coralloid in form. Clavariella

Key to the More Important Genera of Family Hydnaceae

(Based on Miller, 1983)

Fructification with a porose-reticulate hymenial surface covered over with minute
warts over which the hymenium continues. Grammothele

Fructification with distinct warts or teeth, never poroid.

Trama dark; spores roughened, subhyaline to dark, usually brown.

Resupinate, soft, floccose, growing on wood. Caldesiella

Stipitate, fleshy or coriaceous; growing on the ground.

Fleshy. Hydnum

Fibrous tough. Calodon

Trama pale; spores smooth or sometimes echinulate, hyaline or slightly colored.

Teeth arising directly from the woody substratum. Mucronella

Teeth developed on a distinct hymenophore.-

' Some of the following genera may belong in other families but are clavarioid in
form.

2 The genera Hydnochaele, Irpex, and Echinodontium, in which the flattened teeth
develop by the breaking up of pores should be sought in Family Polyporaceae.

KEY TO THE GENERA OF FAMILY FISTULINACEAE 515

Resupinate or reflexed, spines borne on tough branching processes which
are partially submerged in a brownish tomentum.

Gloiodon
Resupinate, reflexed or stipitate; teeth or spines not borne on such proc-
esses.
Resupinate, thin, floccose, crustaceous, ceraceous, or subcoriaceous.
Ceraceous; teeth thick, occasionally slender, obtuse, deformed or ir-
regularly scattered. Radulum
Texture otherwise; teeth varying from short fragile warts to long
conspicuous teeth or spines.
Stellate setae present. Asterodon
Cystidia present. Odontia
Cystidia or setae absent.

Warts short, hemispheric, cylindrical, or subulate and fragile.

Grandinia
Teeth or spines conspicuous, long, slender, usually terete.

Oxyodontia
Reflexed to stipitate, rarely resupinate; fleshy to coriaceous.
Fleshy, growing on the ground.

With central stipes; spores smooth. Dentinum

Pileus irregular, with deformed stipe, spores minutely echinulate.

Hydnodon
Fleshy or coriaceous; growing on a woody substratum.

Richly branched or pulvinate; soft; fleshy. Heridum

Cap coriaceous with long, laterally attached stipe, spores slightly
roughened. Auriscalpium

Reflexed to obscurely laterally stipitate, occasionally resupinate;
subfleshy to coriaceous; spores smooth. Steccherinum

Key to the Genera of Family Meruliaceae^

(Based in Part on Bourdot and Galzin, 1927)
Spores hyaline.

Hymenium with irregular tubercules or radiating (not anastomosing) folds;
waxy, then indurated ; mostly resupinate (sometimes placed in the Hydnaceae) .

Phlehia
Hymenium with lamelliform folds, crisped; membranous; sessile centrally or

laterally. Plicatura (Trogia)

Hymenium with folds anastomosed into alveoli or pores; sessile or resupinate;
more or less gelatinous. Merulius

Spores rust-colored or ochraceous; resupinate, rarely reflexed.

Gyrophana (Serpula)

Key to the Genera of Family Fistixlinaceae*

Fleshy, mostly forming a laterally attached pileus, sometimes resupinate. Pores
remaining separate although in close contact. Fistulina

Membranous or crusty; resupinate; with scattered warts which elongate to
become pores. Porothelium

3 Some authors (Bourdot and Galzin, 1927; Rea, 1922, etc.) include in this family
the following genera with smooth hymenium: Coniophora, Coniophorella, Jaapia.
^ Cyphella and Solenia possibly belong here also, instead of in the Thelephoraceae.

516 CLASS BASIDIOMYCETEAE

Key to the More Important Genera of Family Polyporaceae

Always resupinate; annual (except Fomitoporia); with thin- walled, not truncate

spores; without setae, but sometimes with cystidia (see also
some resupinate species or specimens of Gloeoporus, Hap-
alopilus, Tyromyces, Coriolus, Coriolellus, etc. which are
mainly more or less pileate).^
Context white or light-colored, not becoming brown.

Not becoming brighter on being bruised or with age. Poria

Becoming some bright color on being bruised or with age. Podoporia
Context brown or black.

Strongly irpiciform or hydnoid; at the margin with remains of tubes.

Hydnochaete
Definitely porose.

Fructification black. Melanoporia

Fructification brown, spores hyaline; perennial. Fomitoporia

Fructifications brown, spores brown. Physiporus

Usually pileate; stalked or sessile; reflexed-effuse; under some conditions some-
times resupinate.
Spores truncate at the apical end, two-layered, the epispore smooth and the
endospore with spines or other types of projections into the
epispore; sessile or stipitate; upper surface with a hard, often
laccate, crust. Ganoderma

Spores not as above.

Volva-like structure present. Cryptoporus

Volva-like structure wanting.

Hymenophore definitely porose, in a few species the pores breaking up into
flattened tooth-like plates; annual.
Context white or light-colored, not pronounced brown.
Fleshy or tough; stipitate; context homogeneous.

Stipe much branched, at the bases or near trees. Grifola

{Polypilvs)
Stipe simple; pileus fleshy; terrestrial.

Spores strongly warted, light brown. Boletopsis

Spores smooth, hyaline. Scutiger

(Albatrellus)
Stipe simple; pileus fleshy to tough; large to medium size; growing
on wood or from a sclerotium; stipe central, eccentric or
lateral; pores round or radially somewhat elongated.

Polyporus
(including Melanopus and Leucoporus)
Stipe curved, attached at the top of the minute turbinate hymeno-
phore; emerging from the lenticels of dead twigs; pores
round, small. Porodisculus

Stipe lateral, very short, attached at the top of a large hoof-shaped
to bell-shaped hymenophore which has a thin, separable
pelHcle; on dead trunks of Betula, pores round, small.

Piptoporus

^ Based on Cooke, 1940. In addition, Bondarzew and Singer (1941) recognize |

several other genera of resupinate Polyporaceae, distinguished by entirely different '

characters than shown here. Murrill's (1907) genera differ also in some ways.

KEY TO THE MORE IMPORTANT GENERA OF FAMILY POLYPORACEAE 517

Stipe central or lateral; tubes large, alveolar, often radially elon-
gated; growing on wood. Favolus
Context duplex; spongy above, woody below; centrally stipitate;
tomentose. Abortiporus

(Heteroporus)
Without stipe; sessile to effused-reflexed (rarely resupinate).
Pileus more or less firm, flexible or rigid.

Context duplex, spongy above, firm below. Spongipellis
Context uniform; hymenium at maturity more or less smoke-
colored. Bjerkandera
Context uniform, hymenium white or pallid.

Fleshy to fleshy-tough; friable when dry. Tyromyces

Punky to corky; not friable when dry. Trametes

Pileus thin, leathery and more or less flexible; surface usually zonate.
Hymenophore preceded by a cup-shaped sterile body.

Poronidulus
Hymenophore normally pileate; tubes small and mostly regular;
pore layer not violet-colored; true cystidia lacking.

Coriolus
Hymenophore semi-resupinate; tubes large and irregular; dentate

but not irpiciform. Coriolellus

Hymenophore pileate; tubes early breaking up into flat, tooth-
like plates; margin and pore surface when young violet-
colored; true cystidia present. Hirschioporus
Effused-reflexed or resupinate; poroid remains visible only at
margin. Irpex

(Irpiciporus)
Context bright-colored; yellow to red; not stipitate.

Tubes broad and long; trama orange-red. Aurantioporellus

Tubes small; trama orange to red; of moderate size.

Pycnoporus
Tubes small; trama and tubes yellow; large; imbricate.

Laetiporus
Context brown; spores hyaline.

Duplex; upper layer forming upright tufts of stiff bristles; tubes small.

Pogonomyces
Duplex; upper layer spongy; tubes large. Funalia

Duplex; upper layer spongy; tubes small, sometimes lamellate.

Coriolopsis
Context simple; light brown.

At first fleshy, becoming slightly corky. Ischnoderma

Tough from the first; surface encrusted. Antrodia

Tough from the first; surface not encrusted, glabrous.

Hapalopilus
Context simple; dark brown; context friable; often stipitate.

Phaeolus
Context brown; spores brown.

Sessile. Inonotus

With central stipe; hymenium with setae. Polystidus

With central stipe; hymenium lacking setae. Coltricia

Hymenophore large, woody, perennial; mostly sessile but sometimes
stipitate.

518 CLASS BASIDIOMYCETEAE

Pores breaking down to form flat teeth with denticulate margin.

Echinodontium
Not forming denticulate teeth.

Context wood-colored or pale brown. Fomes

Context white or light-colored. Fomitopsis

Context brown; surface not encrusted. Phellimis

(Pyropolyporus)
Context brown; surface encrusted; several pilei massed together.

Globifomes
Walls of pores breaking down to form concentric lamellae.

Cyclomyces
Pore walls breaking down to form radiating lamellae or labyrinthiform
passages.
Context brown. Gloeophyllum

Context light-colored.

Hymenophore lamellate; woolly and zonate above. Lenzites
Hymenophore typically labyrinthiform, but sometimes broadly lamel-
late; glabrous but rough above. Daedalea
Like Daedalea but the labyrinthiform tubes small, with thin walls,
becoming lamellate with age.^ Daedaleopsis

In the more conservative classifications the genera included in this Key to the
Polyporaceae are left united into a much smaller number of large, polymorphous
genera, as follows:

Poria includes also Podoporia, Hydnochaete, Melanoporia, Fomitoporia, Physi-

porus.
Polyporus includes also Abortiporus, Bjerkandera, Boletopsis, Cryptoporus,

Grifola, Hapalopilus, Inonotus, Ischnoderma, Laetiporus, Phaeolus, Pipto-

porus, Porodisculus, Pycnoporus, Scutiger, Spongipellis.
Polystictus is sometimes included in Polyporus; if maintained as a distinct genus

it may include also Coriolus, Coriolellus, Coriolopsis, Coltricia, Funalia,

Hirschioporus, Poronidulus.
Trametes includes also Antrodia, Pogonomyces, and species out of some of the

foregoing genera grouped in Polyporus and Polystictus.
Daedalea includes also Daedaleopsis.
Lenzites includes also Gloeophyllum.
Irpex and Echinodontium are included in Family Hydnaceae.

Key to the Subfamilies and More Important Genera of Family Boletaceae^

(Based partly on Snell, 194-1 and 1942, and Singer, 1945-194?)

Tubes adhering to one another even to maturity although usually easily separable
mechanically.
Spores mostly dark and ornamented with warts, spines, reticulations, or longi-
tudinal ridges, sometimes smooth spores present on the same hyme-

^ See also Coriolopsis trabea (Pers.) B. & S., whose pores are often lamellate.

7 Gilbert (1931) and Singer (1945-47) divide these genera among two families:
Strobilomycetaceae, including Strobilomyces, Boletellus, and Porphyrellus; and Bole-
taceae, including all the other genera. Coker and Beers (1943) recognize only one
family and three genera: Boletus, Boletinus, and Strobilomyces.

KEY TO THE SUBFAMILIES AND GENERA OF FAMILY BOLETACEAE 519

nium, spores with a germ pore and often truncate at one end, often
over 20 n in length; no clamp connections observed.

Subfamily Strobilomyceteae
Spores globose to short ellipsoid; spore print black; tubes white or gray at
first, becoming darker; pileus and stipe warty, woolly or spinose-
squarrose. Strobilomyces

Spores more elongated; spore print reddish-brown; tubes white to cream
color, becoming pink vinaceous to sordid gray to porphyry brown;
spores smooth or with short warts or spines perforating the exospore.

Porphyrellus.
Spores elongated (or short elliptical in species with reticulate markings);
spore print olivaceous brown to blackish but not reddish; spores
smooth or with longitudinal wing-like ridges or ribs, or with warts, or
reticulately marked. Boletellus

Spores pale pink to yellowish or ferruginous or olive brown; germ pores not
visible; spores mostly less than 20 n in length.

Subfamily Boleteae
Tubes short (see also Phlebopus), radially elongated, often with the radial
walls higher than the cross connections; clamp connections regularly
present; stipe sometimes eccentric or lateral.
Hymeiiophore depressed or subfree around the apex of the stipe.
Spore print yellow. Gyroporus

Spore print olive brown. Phaeogyroporus

Hymenophore more or less arcuate-decurrent.

Veil present. Paragyrodon

Veil absent. Gyrodon

Tubes long (see also Phlebopus, below) ; clamp connections entirely lacking or
not numerous.
Pores radially elongated or lamellate (see also Suillus under Pores round) .
Pores distinctly lamellate, but always with low anastomoses; veil lack-
ing; spore print olive brown. Phylloporus
Pores radially elongated but not lamellate; veil present; pileus rarely
viscid; stipe without glandulae; spore print olive brown.

Boletinus
Pores radially elongated but not lamellate; pileus always viscid; without
veil, or if with veil then pores round; stipe frequently with glandulae;
spore print olive brown. Suillus

Pores round; spores olive brown (see also Suillus under Pores radially
elongated).
Pileus always viscid; veil present; stipe frequently with glandulae.

Suillus
Pileus tomentose; rarely viscid; stipe without glandulae.

Xerocomus
Surface of pileus and stipe more or less pulverulent; spores small, ellip-
soidal to ovoid; tubes long, adnate or depressed around stipe; veil
sometimes present; stipes mostly cylindrical or only slightly thickened
downward, often viscid. Pulveroboletus

Surface of pileus not noticeably pulverulent, sometimes slightly viscid;
tubes short or of medium length; hymenophore arcuate, at least when
young, and somewhat decurrent; veil absent; spores small to medium
size; stipes frequently swollen- ventricose; not viscid.

Phlebopus

520 CLASS BASIDIOMYCETEAE

Veil none; stipe not viscid; pileus sometimes sub-tomentose, not pulver-
ulent; cuticle often consisting of a trichodermium (i.e., a palisade of
parallel, vertical hyphae) ; stipe usually swollen below but sometimes
cylindrical, often reticulately marked but not provided with glan-
dulae; stipe not with furfuraceous or squamulose scabrosities; pores
small; tubes usually long. Boletus

Veil none; stipe not viscid, slender and tapering upward, scabrous;
spores naviculate; pores very small; tubes long, depressed around
stipe. Leccinum

Pores round; spore print rusty yellow; spores under microscope golden
yellow; veil absent; stipe entirely smooth, equal or ventricose; stipe
never reticulate. Xanthoconium

Pores round; spore print flesh color or vinaceous; veil absent; tubes more
or less flesh-colored; stipe often reticulate. Tylopilus

Tubes divergent from one another at maturity as the pileus spreads; only known
from Madagascar. Ixechinus

Keys to the More Important Genera of Family Agaricaceae^

(Modified from Key by A. H. Smith, 1938)

Trama of pileus and usually of the gills composed of nests of sphaerocysts sur-
rounded by connective tissue and with lactifers irregularly dis-
persed throughout. (By many recent students considered as
Family Russulaceae.)
Cut or broken parts of the fruiting body exuding a watery to milk-like or

colored latex. Lactarius

No latex present; fruiting body often very fragile. Ritssula
Trama of pileus not with sphaerocysts.

Parasitic upon other agarics; flesh of cap breaking down into a mass of chlamy-
dospores. Asterophora

(Nyctalis)
If parasitic on agarics, flesh not breaking down into chlamydospores.
Hymenium typically waxy; spores smooth.

Spore print white. Hygrophorus

(including Hygrocybe, Camarophtjllus, Lim.acium)
Spore print smoky gray to blackish. Goynphidius
Hymenium not waxy, or if appearing so, the spores echinulate.

Fruiting body typically rather tough, if fleshy, or membranous and very
pliant; reviving when remoistened.
Gills with distinctly dentate to serrate edges. Lentinus

(including Panus)
Gills with edges even, or merely slightly fimbriate.

Stipe eccentric, lateral, or wanting (usually not reviving when re-
moistened) . Pleurotus

* Used in the wider sense of the term. It should be noted that these keys of the
Agaricaceae are merely artificial keys by which to determine the genera and do not
represent a system of classification based upon phylogenetic considerations. Singer
(1936, 1949) has divided the gill fungi (Agaricaceae in the older sense) into many
families and split up many more of the genera; the bases of segregation being largely
the structure of the gills and type; of surface structure (cuticle, etc.) and the structure
of the spores, as well as the chemical character of the spore and hyphal walls. His
system is, so far as possible, based upon supposed true phylogenetic relationships.

KEY TO THE MORE IMPORTANT GENERA OF FAMILY AGARICACEAE 521

Stipe central, typically 0.5-7.0 mm. in diameter; spores never amyloid.

Marasmius
Fruiting body typically woody or semiwoody.

Lamellae splitting along the fedges. Schizophyllutn

Lamellae arranged concentrically around the stipe.

Cydomyces
Lamellae more or less poroid. Daedalea and Lenzites^

Fruiting body typically soft, or if membranous, rather fragile ; not usually
reviving when remoistened.
Stipe eccentric; lateral, or wanting.

Spore print white to tinged lilac or creamy-vinaceous.

Pleurotus
Spore print pinkish.

Spores longitudinally striate; stipe lacking or rudimentary.

Octojuga

{Clitopilus in part)
Spores as above; stipe well developed. Clitopilus

Spores angular. Rhodophyllus^^

Spore print yellow to rusty brown; lamellae separating easily from the
pileus; the decurrent lamellae sometimes anastomosing to become
poroid near the stipe. Paxillus

Spore print as above; lamellae not separating easily from pileus.

Crepidotus
Stipe typically central.

Spore print white to creamy or pale creamy vinaceous.

See below Key to Leucosporae
Spore print pink to flesh color. See below Key to Rhodosporae

Si^ore print yellow to rusty brown or earth brown.

See below Key to Ochrosporae
Spore print cocoa color, chocolate color or purplish to black.

See below Key to Melanosporae

Key to the Centrally Stipitate, Soft, Putrescent Genera of Leucosporae

Stipe slender, if more than 5 mm. thick then with a distinct cartilaginous cortex.
Stipe somewhat horny in consistency; gills decurrent to adnate; fruiting bodies
marasmoid in appearance but spores amyloid.

Xeromphalina
Not as above.

Cap margin typically straight, or if incurved then gills not truly decurrent;
if margin of pileus incurved and pileus brown to gray to blackish,
then pileus with a differentiated hypoderm.

Mycena
Cap margin strongly incurved or inroUed and gills typically decurrent.

Omphalina
(Omphalia of many authors)
Cap margin strongly incurved and gills typically adnate to adnexed; pileus
lacking a differentiated hypoderm.

^ These three genera, Cydomyces, Daedalea, and Lenzites, are usually included in
Family Polyporaceae.

" Includes several other genera with similar spores.

522 CLASS BASIDIOMYCETEAE

Basidia with darkly staining granules with aceto-carmine.

Lyophyllum

Basidia not as above. Collybia

(including Myxocollybia)

Stipe usually over 3-5 mm. in thickness, typically fleshy.
Stipe readily separable, with a clean break, from the pileus.
Volva present and annulus absent, Vaginata

(Amanitopsis)

Volva and annulus present. Amanita

{Venenarius)

Volva absent; annulus present.

Gill trama divergent; pileus viscid; annulus large to small or obsolete, not
movable. Limacella

{Lepiota in part)

Gill trama interwoven to parallel.

Spores thick- walled, with germ pore; annulus movable.

Leucocoprinus

Spores thin-walled, without germ pore; annulus fixed or movable.

Lepiota
Stipe and pileus confluent (not readily separable) ; annulus present or absent.
Universal veil (or cuticle of cap and the outer layers of sheath on stipe)

powdery to granulose. Cystoderma

Veil if present not granulose on outer surface (hyphal cells not readily sep-
arable from each other).
Partial veil present.

Veil typically leaving a membranous annulus.

Lamellae clearly decurrent. Armillariella

Lamellae emarginate or sinuate, with a tooth.

Armillaria
Veil fibrillose, merely leaving a fibrillose zone on the stipe.

Tricholoma
Partial veil absent or rudimentary.

Spores amyloid; fruiting body typically large and fleshy; pileus ap-
pressed fibrillose to dull and unpolished, rarely hygrophanous and
glabrous ; conspicuous mycelium surrounding base of stipe and ex-
tending through the surrounding debris (if hygrophanous with
rough spores, see Melanoleuca); pigment, if present, intracellular.

Leucopaxillus
Spores not amyloid or if so fruiting body not as above.

Gills broadly emarginate, with a waxy luster and some shade of flesh
color; spores typically echinulate and not amyloid.

Laccaria
Gills not waxy-appearing as above; adnexed to emarginate; spores
typically roughened but amyloid; harpoon-like cheilocystidia pres-
ent; pigment, if present, epicellular. Melanoleuca
Gills decurrent to adnate; fruiting bodies often blackening when
bruised; typically hygrophanous with somber colors; growing in
tufts; basidia with dark granules when stained with aceto-carmine.

Lyophyllum
Gills adnate to sinuate; not hygrophanous; never turning black;

spores typically not amyloid. Tricholoma

Gills decurrent; spores not amyloid, Clitocybe

KEY TO THE MORE IMPORTANT GENERA OF FAMILY AGARICACEAE 523

Key to the Mostly Centrally Stipitate, Soft, Putrescent Genera of Rhodosporae

Spores angular. Rhodophyllus'^

Spores not angular.

Spores longitudinally striate.

Stipe eccentric to central; terrestrial. Clitopilus

Stipe lateral or lacking; not terrestrial. Octojuga

Spores smooth, globose to ellipsoid.

Volva well developed; annulus lacking. Volvaria

Volva typically wanting.

Annulus present. Annularia

Annulus absent.

Gills free; cap and stipe readily separable. Pluteus

Gills attached to stipe; pileus and stipe not separable.

Psathyrella

Key to the Centrally Stipitate, Soft, Putrescent Genera of Ochrosporae

Partial veil cobweb-like; spores typically with a slightly wrinkled to warty exo-
spore and never truncate; typically terrestrial.

Cortinarius

Not as above.

Cuticle of pileus in the form of a viscid pellicle, or radially arranged as inter-
woven filamentous hyphae typically 1.5-5 n in diameter.
Stipe typically fleshy, about 4-20 (up to 40) mm. thick.
Typically terrestrial.

Annulus membranous; rudimentary universal veil often evident.

Rozites
Annulus not membranous, but a fibrillose zone may be present.
Pileus viscid. Hebeloma

Pileus typically dry and fibrillose. Inocyhe

Typically lignicolous.

Annulus present or stipe conspicuously scaly or both.

Pholiota
Annulus lacking; scales if present on the stipe easily obliterated.

Flammula
Stipe typically cartilaginous or slender and fragile.

Spores very pale and thin- walled ; gills typically subdecurrent to decurrent

and pale to bright cinnamon. Tubaria

Spores typically thick- walled and yellowish brown; gills typically adnate
or notched.
Margin of pileus straight at first. Galerina

(Galera in part)
Margin of pileus at first incurved or inroUed. Naucoria
Cuticle of pileus formed by a palisade of pyriform cells, or if the cells are irregu-
larly arranged, vesiculose and nearly isodiametric (best determined
in young pilei) ; spores typically truncate.
Pileus plicate-striate; paraphyses coprinoid; more or less deliquescent.

Bolhitius

11 Thi.s includes the genera known as Enlnloma. Nolanea, Leptonia, and Eccilia.

524 CLASS BASIDIOMYCETEAE

Not as above.

Stipe typically fleshy; spore print typically dull earthy brown to deep
rusty brown. Agrocybe

{Pholiota in part)
Stipe typically cartilaginous; spore print yellowish brown to rusty brown.

Conocyhe

{Galera in part)

Key to the More Important Centrally Stipitate, Soft, Putrescent Genera

of Melanosporae

Gills deliquescing at maturity; basidia separated by broad paraphyses.

Coprinus

Gills not deliquescing.

Pileus with cuticle formed of pyriform or vesiculose cells arranged in a palisade,
or in a compact layer one or more cells thick; fruiting body typically
fragile.
Gills spotted by the maturing spores; spores not discoloring rapidly in con-
centrated sulfuric acid. Panaeolus
Gills not conspicuously spotted by the maturing spores.

Pileus plicate-striate; paraphyses coprinoid. Pseudocoprinus

Pileus not plicate-striate, or, if so, paraphyses not coprinoid.

Psathyrella
(including Hypholoma in part)
Pileus with cuticle formed of slender filamentous hyphae (cells vesiculose in a
few, but then formed by the breaking up of chains, and surface of the
cap appears powdery).
Annulus typically present.

Gills free; stipe readily separating from pileus. Agaricus (Psaliota)
Gills attached; stipe not readily separable. Stropharia

Annulus not present; veil if present adhering to cap or along its margin.
Gloeocystidia present in hymenium; typically lignicolous, or on peat or
muck (if on dung or humus, see Stropharia). Naematoloma

{Hypholoma in part)
Gloeocystidia not present though other types of pleurocystidia may be
present and abundant. Psilocybe

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528 CLASS JASIDIOMYCETEAE

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CLASS BASIDIOMYCETEAE: SUBCLASS
EUBASIDIAE, "GASTEROMYCETEAE"

Gasteromyceteae

THE name Gasteromyceteae (often spelled Gastromyceteae) is given
to a group including those orders of Basidiomyceteae which have one-
celled, two- to four-spored basidia (rarely with more spores) produced
within closed spore fruits. These may have definite means of dehiscence
by which the spores reach the exterior or they may depend upon animals
which feed upon the spore fruits and thus carry the spores to various
places, or the spores escape only upon the decay of the spore fruits.
Mostly the basidia are produced in hymenia which line closed cavities,
but the latter instead of being exposed to the air at the time of spore
maturation, as in the Hymenomyceteae, remain closed until after the
spores have been produced. Frequently the spore fruits are produced
underground, emerging as they approach maturity or remaining sub-
terranean. Other forms grow on stumps, logs, etc. The mycelium usually
shows the presence of clamp connections. Conidia are known in a few
forms. Sexual reproduction is practically unknown, except for the union
of the two nuclei in the young basidium. Few species of this group have
been grown successfully in culture. Even the germination of the spores
has been found impossible of accomplishment in many species. The
basidia may be clavate or cylindrical, as in the majority of the Hymeno-
myceteae, but in the main are more inclined to be ovoid or globose with
sterigmata varying from short to long or even lacking. The spores are
perched symmetrically at the tips of the sterigmata and are not dis-
charged with violence as in the preceding orders. Sometimes the long
sterigma or a portion of it remains attached to the spore, like the handle
of a drumstick.

The young basidia are binucleate and the two nuclei unite, this union
being followed by meiotic divisions leading to the production of four
haploid nuclei. The first meiotic spindle is transverse in the majority of

530

GASTEROMTCETEAE 531

cases (chiastobasidial) but is sometimes longitudinal (stichobasidial), e.g.,
in Tulostoma. In some cases subsequent divisions produce eight nuclei or
even more. Usually the number of basidiospores is four, but not rarely six
to eight and in rare cases up to 12 spores are produced. These are almost
always at first uninucleate but in most of the few cases studied they early
become binucleate by the division of the original nucleus. Whether such
binucleate spores give rise to the dicaryon phase of mycelium is not
proved, but seems probable. Lorenz (1933) has shown that in Sphaeroholus
grandis Lorenz the uninucleate basidiospores give rise to monocaryon
mycelia. These mycelia show two sexual phases, i.e., this species is sexu-
ally bipolar, and only when the appropriate mycelia are mated does a
dicaryon mycelium with clamp connections arise. In this species the
basidiospores are uninucleate, although some species of this genus have
been described as producing binucleate spores. Crucihulum vulgare Tul,
and Cyathus striatus (Willd.) Pers. are both, according to Nils Fries
(1936), quadripolar.

In general the structure of the spore fruit in the Gasteromyceteae is as
follows: Externally there is a peridium consisting of one or more layers. It
may be firm and hard {Scleroderma) or soft and papery or may even dis-
appear during the development of the spore fruit, in Gautieria graveolens
Vittad., being present, according to Fitzpatrick (1913), only in the very
young stages. Within the peridium the tissue may consist simply of the
gleba or of the gleba traversed by "veins" or by a "columella" or by both.
The gleba consists of a more or less fleshy mycelial growth containing
usually numerous hymenium-lined cavities (hymemal cavities) but only
one in a few cases. The columella and veins are slender or stout strands of
hyphae having several functions, food conduction, support, and in some
cases the dehiscence of the spore fruit at maturity. The gleba in many
genera undergoes autodigestion after the basidiospores have been formed.
The tissues involved are the basidia and the fungous tissue lying between
the hymenial cavities, the trama. As a result nothing may be left of the
gleba except the basidiospores, and in a few cases the basidia also, or also
a few stiff threads, the capillitium. These are simple or branching thick-
walled hyphae, which develop in the interhymenial tissues of the gleba
before maturity of the spore fruit. They are rarely septate. The tangled
mass of capillitial hyphae prevents the escape of all the spores at once,
permitting them to sift out a few at a time. In some cases hymenial
cavities with a definite layer of hymenium are wanting but clusters of
basidia appear here and there in the gleba.

The ontogeny of the spore fruit exhibits an early gymnocarpic stage in
some species while in others the earliest stages are angiocarpic. The gym-
nocarpic stage usually becomes closed early so that the development cor-
responds to the pseudoangiocarpic mode of growth found in various

532 CLASS BASIDIOMYCETEAE

Boletaceae and Agaricaceae. The gymnocarpic origin or pseudoangio-
carpic origin of the hymenial surfaces is found in Hemigaster (Juel, 1895),
Chamonixia, Hydnangium, Arcangeliella, Elasmomyces, and probably
other forms. In the majority of genera however, the development is
angiocarpic.

The structure of the spore fruits is not at all on a common plan but
they are very heterogeneous. It is not certain that the 1200 to 1500 species
included in about 120 genera form a group of monophyletic ancestry.
Certain genera are beyond doubt closely related to the Agaricaceae and
may have arisen from that family or may be, as Lohwag (1925), Singer
(1936 and 1950) and others have suggested, in the ancestral line leading to
those fungi. If these are excluded from the Gasteromyceteae the re-
mainder form a more coherent group.

The structure of the basidiospores is of several types and probably
should be given further study with reference to its importance in the
determination of the relationship of the genera. The more or less lemon-
shaped spore with dark color and warty surface is very striking. This type
is found in Gasterella and Gasterellopsis, Hymenogaster, Chondrogaster,
Dendrogaster, and perhaps others. The spores in the Phallales are smooth,
ellipsoidal to cylindrical, and nearly colorless. In many genera the spores
are spherical and spiny, in others smooth. In Podaxis the spore is plainly
two layered, with a terminal germ pore. In Gautieria and Chamonixia the
spores are longitudinally ridged and furrowed, as in Clitopilus and Octo-
juga in the Agaricaceae. In Nigropogon they are angled, somewhat as in
Rhodophyllus {Entolmna) in that family. Until the ontogeny of the spore
fruits from their earliest stages has been studied it is uncertain to what
degree these different spore types represent relationships within or with-
out the Gasteromyceteae.

In Secotium, Elasmomyces and Gasterella, and some other forms cys-
tidia are present in the hymenium. This has been considered by some
investigators to indicate relationship with the Hymenomyceteae. In
Arcangeliella latex vessels are present and also in Lactariopsis (which may
perhaps better be placed in the Agaricaceae close to Lactarius). They are
also described in Battarrea and Phellorinia.

Since the youngest stages of development of the spore fruit have not
been studied, except in a relatively small number of the subterranean
genera, and also are unknown in many of the tropical species whose earlier
stages are not subterranean, the relationships of these fungi to other groups
and the interrelationships of the genera and families making up the
Gasteromyceteae are still far from settled.

In general there appear to be four types of structure, as pointed out by
Lohwag (1924b, 1925, 1926) and Eduard Fischer (many publications,
especially in Engler and Prantl, 1933). These may be called the lacunar,

GASTEROMYCETEAE

533

coralloid, multipileate, and unipileate types. The first stands rather by
itself but the other three represent a gradation from coralloid to unipileate
types. Briefly they are as follows: (1) Lacunar type. Within the fleshy mass
of hyphae destined to form the gleba the tissues pull apart at various
places to form cavities which become lined by a palisade of basidia. A
modification of this type is found in scattered species, genera, and whole
families in which the hyphae producing the tufts of basidia instead of
forming an even layer around a cavity grow irregularly into it, thus form-
ing nests of basidia which obliterate the hymenial cavities. Such a struc-
ture is called "plectobasidial." (2) Coralloid type. From the basal portion
of the spore fruit there grow upward and outward branching masses of
hyphae, all within the enlarging peridium so that a coralloid structure
develops. The ends of these tramal branches may unite with the inner
surface of the peridium and spread along it to form the inner peridial
layer, which is therefore of tramal origin. The spaces between the branches
are at first continuous so that there is in fact only a single cavity very
much interrupted by the coral-like tramal branches which are covered by
a continuous hymenium. Probably the lacunar and coralloid types grade
into one another. Lohwag believes that the former is derived from the
latter. In Lycopcrdon the basal part of the spore fruit is built on the
lacunar plan and the upper, fertile part develops, according to Reh-
steiner (1892) in the coralloid manner. (3) Multipileate type. In this type,
clearly derived from the coralloid type, a number of the tramal branches
are enlarged and reaching the inner surface of the peridium spread out
along it, producing abundant tramal development on the under side of
each "pileus." (4) Unipileate type. The apical pileus of a spore fruit of the
multipileate plan becomes large, clothing the inner surface of practically
the whole peridium, the other pilei remaining rudimentary and not pro-
ducing fertile glebal branches. The main central stalk of the unipileate
type may exist as a "columella" or may elongate downward so that a
stipe is formed. (Fig. 171.)

Fig. 171. Schematic representation of the basic structures of the spore fruits of
various Gasteromyceteae. (A) Lacunar type. (B) Coralloid type. (C) Multipileate
type. (D) Unipileate type. (After Fischer, in Engler und Prantl: Die Naturlichen
Pflanzenfamihen, Zweite Auflage, vol. 7a, Leipzig, W. Engelmann.)

534 CLASS BASIDIOMYCETEAE

The logical classification into orders, families, and genera of the
Gasteromycetes depends upon the probable course of evolution in the
group. The undeniable points of similarity between some of the Agari-
caceae and various unipileate Gasteromyceteae, e.g., Elasmomyces, Seco-
lium, Montagnea, etc., have already been mentioned. Heim (1934) in fact
includes Family Hydnangiaceae in the Agaricales, and others have placed
Secotium and Montagnea in the same order. The fact that the basidiospore
in all Gasteromyceteae is symmetrically perched at the tip of the sterigma
and is not shot off from it makes a derivation of the Agaricales from them
difficult, while the loss of these characters would not be so difficult to
imagine if some Agaricaceae became gasteromycetous in structure and
habit. It must be remembered that the genus Hyaloria in the Hetero-
basidiae has the basidia enclosed in a sort of loose peridium and the
basidiospores there are symmetrically placed and not shot away. Prac-
tically all of the other Heterobasidiae have the usual type of basidium.
The occurrence of transitional forms from Secotium and the apparent
close relationship to some other genera of Gasteromyceteae is an argument
in favor of the theory that there is a close connection between both
groups. Yet it is difficult to imagine simple forms like Protogaster and
Gasterella as being derived from the very complicated Secotiaceae. In view
of the fact that so much investigation still remains to be done on the
development of the spore fruit of many species and genera before a com-
pletely satisfactory conclusion can be arrived at as to the classification
and arrangement within the group the author has decided to follow
Eduard Fischer (1933) in the main, with modifications that seem to him
to be necessary because of more recent studies, especially on Protogaster,
Gasterella and Gasterellopsis, and some of the more secotioid genera,
Gyrophragmium, Battarrea, etc.

The Gasteromyceteae may be arranged in several parallel series, each
ranging from simple to complex structures (or possibly the reverse). As
outlined below the groups with simple structures are placed first. In one
series the spore fruit is mainly fleshy, with well-developed hymenial
chambers. It does not undergo partial autodigestion and depends upon
decay or mechanical destruction for the distribution of its spores. This
series contains forms of increasing complexity from Protogastraceae to
Hymenogastraceae and on to Sclerodermataceae on the one hand and on
the other through the Hydnangiaceae and finally to the Secotiaceae,
which last family has close connection with the Agaricaceae. Another
series leads from the partially gelatinous or cartilaginous Ilysterangiaceae
to the Clathraceae and Phallaceae, with increasing tendency to the auto-
digestion of the gleba to form a slimy, evil-odored mass attractive to
insects which distribute its spores. A third series, arising again in the
Hymenogastraceae leads to the Lycoperdaceae and Geastraceae, and

ORDER PROTOGASTRALES 535

Tulostomataceae and possibly the Podaxaceae. Some genera usually
assigned to the Sclerodermatales probably belong in this series. The gleba
breaks down by autodigestion and leaves the spores and sometimes also
the basidia and the hyphae making up the capillitium, as a dry powdery
mass, spore distribution taking place by air currents. A fourth tendency
shown by the Sphaerobolaceae, Nidulariaceae and the genus Pisoliihus, in
the Sclerodermataceae, is the formation of firmer walls around definite
regions of the gleba so that these are distributed as units, either by
mechanical means or currents of water or by violent expulsion from the
fruit body. These probably do not form a connected series but represent
separate evolutionary changes from both the Hymenogastrales and the
Sclerodermatales.

The orders tentatively recognized here are Protogastrales, Hymeno-
gastrales, Sclerodermatales, (possibly better distributed among other
orders), Lycoperdales, Nidulariales, Sphaerobolales, Phallales. The late
Sanford M. Zeller (1948, 1949) recognized 9 orders and 32 families instead
of the 7 orders and considerably fewer families recognized in this work.
Until much more intensive study has been given to the ontogeny of the
sporocarps of the G aster omyceteae any arrangement must be considered
to be more or less tentative.

Order Protogastrales. In this order the minute spore fruits have but a
single hymenial cavity. The hymenium consists of basidia without cys-
tidia, and the basidiospores are light-colored and smooth. Protogaster was
described by Zeller (1934). It is a minute fungus growing on the roots of
plants in Maine and has been found but once and then only in the mature
stages. It consists of a nearly spherical spore fruit, less than a millimeter
in diameter. It contains a single large cavity lined by basidia bearing

^^IX I

'% ^^

^^:^ J^l^""^

•''>;?:■• •

Fig. 172. Protogastrales. (A) Family Protogastraceae. Protogaster rhizophilus
Thaxt.; diagrammatic median section of spore fruit. (B-D) Family Hemigastraceae.
Hemigaster candidus Juel; three stages in the pseudoangiocarpic development of the
spore fruit. (A, courtesy, Zeller: Ann. Missouri Botan. Garden, 21(2):231-240. B-D,
from Comparative Morphology of Fungi by Gaumann and Dodge, New York,
McGraw-Hill Book Co., Inc.)

536 CLASS BASIDIOMYCETEAE

colorless ellipsoid spores. There are no noticeable projections or folds on
the surface of the hymenial cavity and no cystidia have been observed. In
the absence of younger stages it is impossible to locate this with certainty.
Perhaps it is wiser to place this in a distinct family Protogastraceae in the
Order Protogastrales. (Fig. 172 A.)

Possibly Juel's genus and family, Hemigaster and Hemigastraceae,
respectively, should be placed in this same order. The minute spore fruit
grows on rabbit excrement. It is 2 to 3 mm. tall and at maturity about as
broad. It arises as an upright tuft of parallel hyphae which spread at the
top like a sheaf of grain. The spreading hyphae curve downward and in-
ward and eventually join with hyphae growing out from the stipe to form
a circular chamber centrally pierced by the percurrent columella which is
in reality the upper portion of the stipe. Two layers are visible in the
peridium, an outer loosely woven portion and an inner denser subhy-
menial layer from which arise the basidia which cover the upper and outer
side of the circular hymenial cavity, but not the inside formed by the
columella. The basidia bear four nearly spherical, pale, flesh-colored
basidiospores with smooth surfaces. There are no cystidia or paraphyses in
the hymenium. From the columella there grow out into the cavity slender
hyphae which bear colorless chlamydospores wound about by slender
hyphae. Eventually the basidiospores and chlamydospores fill the cavity.
The mode of development of the spore fruit is of the type called pseudo-
angiocarpic. Since the younger stages of Protogaster are not known it is
not possible to determine definitely whether these two fungi are related or
not. Juel (1895) concluded that Hemigaster is related to the Thelepho-
raceae but it seems to the author that it belongs rather in the Gastero-
myceteae. (Fig. 172 B-D.)

Order Hymenogastrales. These are mostly subterranean, rarely super-
ficial, when young, growing so as to become external at maturity in many
cases. The spore fruits are fleshy to cartilaginous or somewhat gelatinous.
The spore dispersal is not by means of digestion of the gleba into an
insect-visited slimy liquid or by the production of a dry mass of wind-
conveyed spores. The gleba retains its structure essentially till maturity
of the spores. The thin (or evanescent) or firm peridium surrounds a gleba
of uniform structure or traversed by spreading "veins" or with a central
columella which in one family reaches the apex, i.e., a percurrent colu-
mella. The development of the gleba is possibly lacunar or more often coral-
loid, multipileate or unipileate. By the usual classification, that of Eduard
Fischer (1933), four families are recognized, depending upon the type of
development. Many spore types are found, colorless or colored, smooth or
verrucose, or ribbed, etc. Possibly it may be feasible, when the youngest
stages of development have been studied for most of the genera, to cor-
relate development, spore type and mature morphology to produce a more
satisfactory system of classification.

ORDER HYMENOGASTRALES 537

Family Hymenogastraceae. The type of this family is the genus
Hymenogaster. This is subterranean or with the upper surface projecting
above the ground at maturity. In very young specimens Rehsteiner
(1892) showed that a pahsade layer of cells develops as a boundary setting
off the thick sterile base from the upper peridium. This layer arches up-
ward with the growth in size of the spore fruit forming a single central
cavity into which grow downward from the roof various branches and
plates which extend to and apparently in places grow fast to the base.
These plates anastomose with one another so that labyrinthiform hy-
menial cavities are formed. With the increase in size of the spore fruits the
relative size of the sterile base becomes much smaller. Eventually the
spore fruit shows very numerous irregular cavities, sometimes radiating a
little from the sterile base. The peridium is rather firmly attached to the
outer side of the gleba. The hymenial cavities are lined with clavate
basidia which bear two, rarely four, spores. These vary with the species
but are mostly ellipsoidal, ovoid or limoniform, mostly yellow to brown in
color, smooth or more often verrucose or wrinkled. No cystidia are de-
scribed for this genus. Apparently closely related to the foregoing is the
unilocular genus Gasterella, described by Zeller and Walker (1935) and
Miss Walker (1940). It grows on the surface of the soil and reaches the
diameter of 300 to 700 ju and even up to over 1200 m- According to Routien
(1939) rhizomorphs are attached to the spore fruits and their hyphae show
clamp connections although these are not visible in the basidiocarp. In
the specimens that reach maturity in the smaller dimensions the cavity
may have smooth walls but in the larger spore fruits the hymenium sends
folds and projections into the cavity from above and from the sides. These
greatly increase the hymenial surface but do not reach the bottom, so that
the spore fruit remains unilocular. The basidiospores arise in twos or more
often fours and are dark-colored, verrucose and somewhat apiculate.
When detached a piece of the sterigma often remains attached to the
spore. Cystidia with black verrucose heads are sometimes found, espe-
cially in the smaller specimens that have developed under less favorable
conditions. They appear to be, perhaps, aborted basidia. In the very
young spore fruits an arching palisade layer of densely staining cells ap-
pears in the midst of the loose tuft of hyaline hyphae. As it broadens and
arches up further a cavity is formed into which eventually push the plates
or folds which partially divide up the single cavity. Miss Walker (1940)
suggested that Gasterella should be placed in Order Protogastrales but in
a separate family Gasterellaceae. In the author's opinion the similarity
of Gasterella to the young stages of Hymenogaster rehsteineri Bucholtz is
too great to allow their separation into different families and orders.
(Fig. 173 A-E.)

Gasterellopsis (Routien, 1940) begins its development much as in
Gasterella except that there is a central percurrent columella so that when

538

CLASS BASIDIOMYCKTEAE

Fir,. 173. Hymenogastrales, Family Hymenogastraceae. (A) Gasterella lutophila
Zeller & Walker; vertical section through a spore fruit showing the tendency toward
the formation of folds from the hymenial wall. (B, C) Hymenogaster rehsteineri
Bucholtz, showing vertical sections of very young and somewhat further developed
spore fruits. (D, E) Hymenogaster tener Berk. (D) Section through mature spore fruit.
(E) Portion of mature gleba. (A, courtesy, Zeller and Walker: Mycologia, 27(6) :573-
579. B-E, after Tulasne, from Fischer, in Engler und Prantl: Die Natlirhchen Pflanzen-
familien, Zweite Auflage, vol. 7a, Leipzig, W. Engelmann.)

OKDER HYMENOGASTRALES 539

the hymenial layer begins to appear it forms the roof of a circular cavity
instead of a depressed spherical one. From all sides of this cavity, except
the columella, centripetally directed folds begin to form and become
covered by the mature hymenium. They may stop short of the columella
leaving the circular cavity not completely divided but with numerous
radial lobes, or some or all of these folds may grow to the columella in
which case separate radial cavities are produced. Eventually the peridial
tissues in contact with the basal portion of the columella dissolve, thus
forming a basal circumscissile opening and then the remainder of the
peridium and finally of the tramal tissues dissolve so that a single cavity
filled with spores is left. Its outer wall then is the remains of the sub-
hymenial layer. The spores are like those of Gasterella and are produced by
twos or fours on the basidia. No cystidia were observed. Routien sug-
gested that this represents a further step in complication from Gasterella,
and perhaps should be placed in the same family with it, since it starts as
a unilocular fungus and often remains so. On the other hand it shows
great similarities, although it is much simpler in structure, to the Se-
cotiaceae. Rhizopogon is a genus of thirty or more species with subter-
ranean basidiocarps whose surface is covered with numerous loose or
adherent branching fibrils which lead into rhizomorphs. Its spores are
more or less ellipsoidal and smooth. The young spore fruit has a central
portion of loosely branching coralloid structure with the interconnecting
open spaces Hned with hymenium. The basidia are two- to eight-spored.
If a much enlarged Protogaster should develop invaginating and branched
ridges and lobes it would show many of the characteristics of Rhizopogon.

Pilat (1934) discussed the genus Gastrosporium and based upon it the
family Gastrosporiaceae. In the latter the peridium is double while in the
Hymenogastraceae it is, according to him, simple.

Family Melanogastraceae. This is an assemblage of several more
or less related genera that differ from the Hymenogastraceae in having
their lacunar hymenial cavities more or less filled or obliterated by a
gelatinous mass which in Leucogasier appears, according to Zeller and
Dodge (1924), to be the product of the gelification of conidia or chlamydo-
spores which were produced prior to basidial development. Into these jelly
filled cavities long, slender basidia push their way, partially filling them
with a crisscross tangle. The basidiospores are often coated with a gelati-
nous layer and arise two to eight perbasidium. They are almost colorless in
Leucogasier and dark brown in Melanogaster. The spore fruits are sub-
terranean or partially emerging at maturity, and without a stalk. The
gleba is traversed by veins or sheets of tramal tissue that divide it into
polyhedric or rounded units each of which is a "basidial nest" as some
authors call it. Fischer (1933) places Alpova tentatively in this family. It
has been reported only from the United States so far. It is partially sub-

540 CLASS BASIDIOMYCETEAE

terranean at maturity and reaches a diameter of 5 to 20 mm. The glebal
chambers are at first filled with large spherical cells which then gelatinize.
Long hyphae traverse these cavities and on them sit the long, slender
basidia which bear 5 to 11 pale brown, almost sessile, ellipsoid, smooth
spores. Dodge (1931) who described the genus considered it to belong to
the Rhizopogonaceae, a family segregated from the Hymenogastraceae,
and including a number of genera placed by Fischer in the Melanogas-
traceae. Zeller (1939) held that its development suggests closer relation-
ship to the latter family than to Rhizopogon.

Family Hydnangiaceae. In this family of Hymenogastrales the
coralloid development of the gleba has become unipileate. Like Hemigaster
the spore fruits are pseudoangiocarpic in their development. The upper
portion of the stipe becomes the percurrent columella. Unhke Hemigaster
the palisade layer of the under side of the recurving pileus is thrown into
folds which anastomose with one another and with the columella so that a
multilocular gleba is produced. The columella may become reduced with
age to a slender, scarcely recognizable strand, in some species. The stipe
below the pileus is represented by only a small projection, if visible.
Cystidia and spiny basidiospores two to four per basidium on long sterig-
mata are found in Hydnangium and Arcangeliella. In the latter genus
laticiferous tubes are present. In Chamonixia the spores are longitudinally
ribbed, much as in Gautieria.

Family Secotiaceae. In this family the general plan is much like that
of the preceding one, but the stipe is more pronounced in most forms.
Development is pseudoangiocarpic in Elasmomyces and angiocarpic in
most of the remaining genera. Elasmomyces represents probably an inter-
mediate form between Hydnangium and Secotium. The fruit body is
mostly eventually epigeous. Its development is pseudoangiocarpic like
that of Hydnangium. In the tissue of the stipe are nests of enlarged,
bladder-like cells, resembling those of Russula in the Agaricaceae. The
spores are marked with verrucosities, sometimes connected by ridges, as in
Russula, and as in that genus they are stained blue with reagents contain-
ing free iodine. Bucholtz (1903) considered these two genera, as did
Malengon (1931), to be closely related. Heim (1938) also emphasizes the
relationship of Lactarius and Russula to Elasmomyces.

The genus Secotium, with which Elasmomyces is sometimes united, is
angiocarpic in its development in the species studied (*S'. agaricoides
(Czern.) Hollos, by Conard, 1915, S. novae-zelandiae Cunningh. and *S'.
erythrocephalum Tul., by Cunningham, 1924 and 1925). (Fig. 174.) In the
earlier stages of development it shows great resemblance to that of
Agaricus (Atkinson, 1906, 1915), but instead of forming radial lamellae
separating the annular opening into radial cavities the tramal plates are
irregular in the direction of their growth and anastomose to form closed

ORDER HYMENOGASTRALES

541

Fig. 174. Hyraenogastrales, Family Secotiaceae. Secotium erythrocephalum Tul.
(A) Mature plants. (B) Section of tramal plate. (C, D) Vertical sections through young
and nearly mature plants. (Courtesy, Cunningham: Brit. Mycol. Soc. Trans., 10:216-
224.)

cavities. Apparently within the main tissues of the pileus other hymenial
cavities develop as well as in the thick tramal plates in the gleba so that
finally the latter is made up of very many narrow, hymenium-lined cav-
ities, separated by thin tramal plates. As the stipe elongates the edge of
the pileus pulls loose from it and the gleba becomes partly exposed to the
air. Since the glebal cavities are not continuous the spores are distributed
by the action of insects infesting the spore fruits and also by the decay of
the latter. The presence of cystidia in some species of Secotium is another
point of similarity to the Agaricaceae. There are some very small species,
e.g., S. coprinoides Routien (1940), about 4 mm. tall with a pileus about
2 mm. broad. Its gleba consists of numerous (about 18) radial hymenial
cavities extending from the lateral peridium to the stipe. The hymenium
consists of two- or four-spored basidia intermixed with paraphyses. The
ellipsoidal spores are smooth, black, and with a short pedicel formed by
the apical portion of the sterigma. S. olbium Tul. is 4 to 6 mm. tall but
the spores are smaller, spherical, and wrinkled. The structure of the gleba
of the former is somewhat similar to that of Gasterellopsis, but is not some-
times unilocular and the spores are of different types. There is no deli-
quescence so that this is not a Coprinus and the position of the spore on
the sterigma is typical of the Gasteromyceteae. At maturity the edge of

542

CLASS BASIDIOMYCETEAE

the pileiis breaks loose from the base of the stipe as the latter elongates
and leaves no volva.

Possibly closely related to Secotium are Gyrophragmium, Longula, and
Montagnea {Montagniies). In them the mature spore fruit is more highly
organized than in Secotium. When the pileus expands, exposing the under
side of the gleba, it leaves a volva, and sometimes an annulus, in Gyro-
phragmium, and a two-layered annulus, but no volva in Longula. In these
genera the expanded pileus is convex. The gleba is lamellar, but consider-
ably anastomosed. In Montagnea the pileus at maturity is a small disk at
the apex of the stipe with the black radially lamellar gleba hung beneath
it and free from the stipe as in the foregoing genera. This extends beyond
the disk as separate lamellae. There is a volva but no annulus. The de-
velopment oi Longula texensis (B. & C.) Zeller has been studied by Barnett
(1943). It is angiocarpic and similar to that of Agaricus.

Family Hysterangiaceae. In this family the coralloid structure of
the developing gleba is very marked. The enlarged end of a rhizomorph
develops into a body with a peridium and a central core. The enlarging
core begins to form folds and plates under the expanding peridium. These
anastomose so that eventually a multilocular gleba is formed with a

Fig. 175. Hymenogastrales, Family Hysterangiaceae. Gautieria plunibea Zell. &
Dodge. (A) Vertical section of fruiting body. (B) Portion of hymonium. (C) Basidio-
spores. (Courtesy, Zeller and Dodge: Ann. Missouri Botan. Garden, 5(2):133-142.) J

ORDER PHALLALES

543

dendroidally branching system of supporting branches arising at the base.
The peridium disappears at an early stage of development in Gautieria so
that the gleba is exposed. The spores in this genus are ribbed longitudi-
nally and resemble those of Clitopilus and Octojuga in the Agaricaceae and
Chamonixia in the Hydnangiaceae. In Hysterangium the peridium persists
and the glebal branches may grow fast to and spread along its inner
surface. A gelatinous subperidial layer may develop from the trama where
it comes into contact with the peridium. The cartilaginous -gelatinous
tramal character and the production of the gelatinous subperidial layer
are the main distinctions between this family and the Hymenogastraceae.
Protubera and Phallogaster are genera that show further transitional steps
toward the Clathraceae in the Order Phallales. (Fig. 175.)

Order Phallales. These are noteworthy because of the dissolution of
their gleba into a usually evil-smelling slimy mass filled with spores. This
attracts flies, especially those that feed upon and lay their eggs in carrion.
They serve to carry the spores far and wide. It was shown by Cobb (1906)
that the spores were not injured in their passage through the alimentary
canal of these insects. At first the spore fruits are completely or partially
subterranean, more or less spherical, with a firm, somewhat leathery
peridium, underneath which is a thick layer of slime resembling the white
of a raw egg. This, according to Lohwag (1925), is a modified outer por-
tion of the gleba. The functional portion of the gleba is supported upon
the surface of a pileate "receptacle" in the Phallaceae or upon or between
a framework of radiating or anastomosing branches in the Clathraceae. In

Fig. 176. Phallales, Family Clathraceae. (A) Clathrus ruber Mich, ex Pers. (B, C)
Pseudocolus javanicus (Penz.) Lloyd. (B) Fully expanded spore fruit. (C) Cross section
of "egg" just about to open, showing volva and four receptacular arms surrounding
the mass of gleba. (A, courtesy, Lloyd: Mycological Writings, vol. 3. B-C, after
Bernard: Ann. Jardin Botan. Buitenzorg, 31:93-102.)

544 CLASS BASIDIOMYCETEAE

the former there is a stout, rapidly expanding stipe whose expansion tears
open the peridium at the top leaving it around the base of the stipe as a
volva. (Fig. 178.) In the Clathraceae the stipe may be present {Simhlum,
Pseudocolus (Fig. 176 B-C), Aseroe, etc.) bearing the receptacle at its top,
or absent, the enlarging of the network of the receptacle rupturing the
volva (Clathrus). The young gleba is more or less coralloid in its develop-
ment and forms a complicated system of branching and anastomosing
branches and plates covered by the hymenium. These dissolve completely,
except the spores, to form a slimy malodorous mass.

The simple Phallales have many points of resemblance to the Hyster-
angiaceae, in some of the genera of which the gleba eventually dissolves
(e.g., Phallogaster) . The majority of the order are tropical or subtropical,
but several genera are common in the temperate regions. Among these
are, in the Clathraceae, Clathrus ruber Mich, ex Pers. (Fig. 176 A) which
forms a pyriform coarse net with thick receptacular branches, arising
from the ruptured volva. The dissolved gleba lines the inner surface of the
hollow receptacle. The fungus is 6 to 8 cm. tall and red, rarely yellow, in
color. Lysurus australiensis Cke. & Mass. (Anthurus horealis Burt) has a
white to pink stipe with several connate, outwardly furrowed receptacular
arms at its top, the whole reaching a height of 10 cm. In the Phallaceae
the commonest genera of the temperate regions are Mutinus, Phallus, and
Didyophora. In the first the receptacle is a closely appressed cap on the
upper portion of the stipe which, as in all the stalked members of the
order, stands in the ruptured volva. The pileus is usually some shade of
red as is often the case for the upper portion of the stipe. The mainly
European M. caninus (Huds. ex Pers.) Fr. is usually without offensive
odor while M. ravenelii (B. & C.) Fisch., which is the commoner form in
the United States, has a foul odor. Phallus and Didyophora have a bell-
shaped pileus free from the stipe except at the top. The dissolved gleba
covers the pileus which may be smooth but in most species is reticulate
with large shallow pits. The commoner species in Europe is P. impudicus
L. ex Pers., sometimes attaining a height of 15 cm. or more with a pileus 3
to 3.5 cm. broad. The stipe is white and 2 to 3 cm. thick. The color of the
pileus when free from the spores is mostly white. In the eastern United
States the commoner species is P. ravenelii B. & C, with a reddish stipe
and the surface of the pileus not strongly reticulately marked. Didyophora
differs from Phallus by the formation of a beautiful skirt-like "indusium"
attached near the top of the stipe beneath and free from the pileus. This
is white in color and reticulate with large meshes. The troi)ical D. indu-
siata (Pers.) Fisch. has a larger indusium than D. duplicata (Bosc ex Fr.)
Fisch. which is frequent in the eastern United States and occasional in
Europe. (Figs. 177, 178.) The tropical genus Itajahya, first described from
Brazil, has recently been discovered in Arizona and New Mexico by Long

OEDER PHALLALES

545

Fig. 177. Phallales, Family Phallaceae. Dictyophora duplicata (Bosc ex Fr.) Fisch.

(Courtesy, Walters: Mycologia, 35(1).)

and Stouffer (1943). In it the pileus has numerous overlapping trama
plates between which the gleba is formed. When the latter is washed
away the pileus resembles a wig perched at the apex of the stipe.

The genus Claustula, whose ovoid receptacle remains enclosed in the
volva until the gleba is completely mature, has been placed by Cunning-
ham (1931) in a separate family, the Claustulaceae, considered by him to
be more primitive than the remainder of the order. Fischer (1933) includes
this genus in the Clathraceae with a total of 15 genera and recognizes 10
genera in the Phallaceae.

546

CLASS BASIDIOMYCETEAE

Fig. 178. Phallales, Family Phallaceae. Didyophora indusiata (Pers.) Fisch. (A)
Vertical sections through two unexpanded eggs, the lower one further advanced.
(B) Vertical section through an expanded spore fruit whose indusium has not yet
expanded. (G) Gelatinous layer of volva. {H) Pileus. (/) Indusium. {Sw) Wall of
stalk. {S) Stalk axis becoming the hollow at maturity. (P) Primordial tissue between
stalk and indusium. {Pi) Primordial tissue between pileus and indusium. (a) Gleba.
(After Fischer: Ann. Jardin Botan. Buitenzorg, Serie I, 6:1-51.)

Order Sclerodermatales (Plectobasidiales). These were set apart from
the Hymenogastrales by Schroeter (1897) and recognized by Fischer
(1933, 1936). The distinction is based upon the structure of the gleba
which mostly does not exhibit sharply defined hymenial cavities lined by
an even layer of basidia as is typical of most Gasteromyceteae. Even in
this order a tendency toward chamber formation is apparent in the younger
basidiocarps but the hyphae whose terminal cells are destined to become
basidia grow out into these incipient cavities to different lengths so that
they are more or less completely obliterated, being represented by nests

ORDER SCLERODERMATALES (PLECTOBASIDIALES)

547

of basidia. According to Fischer these chambers arise in the lacunar
manner. There is great need of developmental studies before the relation-
ships within this order can be determined as well as to the Hymeno-
gastrales, from which they undoubtedly have arisen. At maturity the
gleba mostly becomes a powdery mass of spores with more or less capil-
litium. The peridium may be thin but is often several-layered and thick
and firm, hence the name of the principal genus, Scleroderma. The spores
are from four to six or more on the basidium, sessile or nearly so. They are
usually dark-colored. Only one family, Sclerodermataceae, appears to the
author to belong in this order. It is really doubtful whether the oblitera-
tion of the hymenial cavities is of sufficient importance to warrant the
removal of this family from the Hymenogastrales, especially in view of
the fact that Melanogaster, Leucogaster, Alpova, and others have a tend-
ency toward this structure. (Fig.* 179.)

Fischer (1933) included 10 genera in the family, of which he indicates
five to be in doubt. Of the typical members of the family Scleroderma and
Pisolithiis may be noted. The former is subterranean or growing on the
surface of the soil. It forms rounded spore fruits, in some species reaching
a diameter of 10 cm. If superficial, there is a rooting mass of mycelial
strands, the base being sometimes slightly stipe-like. The peridium is
thick and at maturity more or less leathery. The surface may be smooth
or roughened or in some cases forming large overlapping scales. The
mature gleba shows numerous dark basidium-producing areas, separated
by sterile veins or sheets. There is no definite hymenial layer in the
hymenial cavities. The pyriform basidia bear two to five nearly sessile,
rounded or ellipsoid, smooth or sculptured spores. The basidia disappear

Fig. 179. Sclerodermatales, Family Sclerodermataceae. Scleroderma aurantiacum
Pers. (A) Vertical section through an almost mature spore fruit. (B) Basidia, with
sessile spores, with completely obliterated hymenial cavity. (After Tulasne, from
Fischer, in Engler und Prantl: Die NatiirUchen PfianzenfamiUen, Zweite Auflage,
vol. 7a, Leipzig, W. Engelmann.)

548

CLASS BASIDIOMYCETEAE

early and the spores are in some species enveloped in a sheath of nurse
hyphae till maturity when everything disappears except the spores and a
few capillitial threads. The spore fruit may crack open stellately or may
be opened by attacks of insects or rodents. Pisolithus does not have so
thick a peridium and the tramal sheets split in such a manner that the
basidium-producing areas form numerous ellipsoid or round "peridioles"
retained within the peridium. Finally these fall apart into a powdery spore
mass. The capillitial threads are few. The basidia bear two to six almost
sessile, rounded spores. There is no special means of dehiscence provided
in this genus.

Order Nidulariales. In this order the spore fruits are external, not
subterranean. They have a thin or thick peridium and the gleba contains
several to innumerable hymenial cavities, apparently formed in the lacu-
nar manner, each lined internally by a layer of basidia. The tramal tissue
in Family Nidulariaceae forms a firm several-layered wall around each
cavity with its contained spores and these hard-walled bodies, called
"peridioles" lie in the bottom of the cup formed by the dissolution of the
top peridium of the spore fruit and of the tissues surrounding the peridi-
oles. In Family Arachniaceae placed in this order by Fischer, the perid-
ioles are innumerable and the tramal wall of each is thin and fragile.

Family Nidulariaceae (Bird's Nest Fungi). Several peridioles,
over 20 in Nidularia, are formed in each spore fruit. At first connected in
a continuous gleba the peridioles early become separated from each other
and lie free in the cavity of the spore fruit or are connected to the peridium
by long slender strands, the "funiculi." The principal genera are Cruci-
bulum, Cyathus, and Nidularia. The spore fruits are several millimeters

Fig. 180. Nidulariales, Family Nidulariaceae. Cyathus striatus Pers. Three
opened fruit bodies showing peridioles and one unopened spore fruit. (Courtesy,
F. C. Strong.)

ORDER SPHAEROBOLALES

549

Fig. 181. Nidulariales, Family Nidulariaceae. (A, B) Cyathus stercoreus (Schw.)
DeToni. (A) Section through wall of mature peridiole. (B) Basidia with their sessile
basidiospores. (C) Crucihulum vulgare Tul.; vertical section through a portion of
immature spore fruit. (A, B, courtesy, Coker and Couch: The Gasteromycetes of the
Eastern United States and Canada, Chapel Hill, Univ. North Carolina Press. C, after
Sachs: Botan. Ztg., 13(48) :833-845; (49):849-861.)

up to a centimeter in height and funnel-formed or almost spherical, with
a flattened top. The peridium on this flattened upper portion ruptures
and exposes the peridioles lying like eggs in a nest, whence the common
name of the fungi. B. O. Dodge (1941) reports that they are discharged
from the spore fruit at maturity, in some cases to a height of 3 or 4 meters.
In forms with a funiculus (e.g., Cyathus) the latter remains attached to
the peridiole when it is discharged and, being sticky, attaches it to various
objects with which it may come in contact. Germination of the basidio-
spores occurs within the peridiole from whose outer surface numerous
germ tubes emerge in all directions. According to Martin (1927), in the
development of the peridiole the basidia collapse while the spores are not
yet mature or fully grown. The spores are then nourished by a weft of
hyphae surrounding each spore, much as occurs in some species of Sclero-
derma. (Figs. 180 and 181.)

Order Sphaerobolales. This order, included in the preceding one by
many mycologists, has but one family, the Sphaerobolaceae. In the two
genera Sphaeroholus and Nidulariopsis, the peridium has three or more
rather thick layers, the middle one of which is lacking in the apical region

550 CLASS BASIDIOMYCETEAE

of the second genus. The tension arising from the osmotic swelling of the
cells of one of these layers results in the violent eversion of the inner
peridial wall so that the whole glebal mass, 1 to 2 mm. in diameter, is
ejected. In Sphaeroholus Miss Walker (1927) has shown that this glebal
ball may be shot upward to a distance of over 4 meters. In Nidulariopsis,
according to Greis (1935), the distance is short. The basidia in S. stellalus
Tode ex Pers. are arranged irregularly in a number of clusters of hyphae
intermingled with the basidia, as in Scleroderma, these groups being
separated by thin hyphal sheets. In the genus Nidulariopsis there are
numerous definite hymenial cavities lined by basidia, as in the Hymeno-
gastraceae and Lycoperdaceae. The basidia bear from four to nine spores.
The ejected gleba germinates as in Nidulariales by numerous hyphae from
all sides. In the course of the rupture of the spore fruit the outer layers of
the peridium are split into five or more lobes, in some regards resembling
a partially opened Geastrum.

Order Lycoperdales. The remaining order of the Gasteromyceteae is
Order Lycoperdales. This carries on the tendencies noticeable in the
Sclerodermataceae of destruction of the glebal tissues to form a dry
powdery mass of basidiospores intermingled more or less with sterile
hyphae which form the capillitium. The function of the capillitial hyphae
in the young spore fruits may well be that of water and food conduction
as suggested by Fischer (1936) and Zeller (1939). At maturity these
hyphae, now empty and more or less thick-walled, serve to keep the
spores loosened up so that they do not escape all at once but gradually,
over a longer period. The spore distribution therefore has become entirely
dependent upon air currents. The fully grown gleba undergoes autodiges-
tion which involves the hyphae of the trama (except those that become
the capillitium) and mostly the basidia also. At this stage the contents of
the spore fruit form a soggy, water-soaked mass. The water is quickly
evaporated or perhaps also returned to the mycelium so that soon the
gleba is dry, colored brown to purple, depending upon the color of the
spores and of the capillitial threads.

Fischer (1933) recognized only two families in this order, Lyco-
perdaceae and Geastraceae, sometimes united into one family. In spite of
the difference in glebal structure in some genera it seems to the author
that the Tulostomataceae and Podaxaceae perhaps should find their posi-
tion here, as representing in this order the same evolutionary trend toward
the stipitate pilear structure as is shown in the series Hydnangiaceae to
Secotiaceae in the Hymenogastrales. The knowledge of the early develop-
mental stages is lacking in many of the genera but in those that have been
studied it seems that the gleba may be lacunar or coralloid in its develop-
ment, often the former in the basal, first developed portion of the gleba
and the latter in the upper, fertile portion. Whether Podaxis represents a

OEDER LYCOPERDALES

551

Fig. 182. Lycoperdales, Family Lycoperdaceae. Lycoperdon pyriforme Pers. (Courtesy,

F. C. Strong.)

unipileate modification of the coralloid type of development needs careful
study of the very early stages.

Family Lycoperdaceae. The spore fruits are external from the first
or may be shallowly subterranean when young, becoming external at
maturity. They consist of a flexible peridium of two or three well-marked
layers enclosing the gleba. The basidia are ovoid with short or long sterig-
mata and four to eight basidiospores. After the spores are mature the
basidia and the tramal tissues dissolve, except for the brown, thick-walled
capillitial threads. As the spore fruit enlarges the outer peridium ruptures
in various ways, scaling off in granules or larger pieces. The inner peridium
may also break up in pieces or more often forms one or more ostioles, usu-
ally in the apical region. As wind or firmer objects strike the spore fruit
the spores are puffed out through these ostioles. The spore fruits vary
from a few milhmeters in diameter up to 1.6 m. in length, 1.35 m. in width
and 24 cm. in height, in the case of a specimen of Calvatia gigantea (Batsch
ex Pers.) Lloyd, collected in New York State many years ago and reported
by C. E. Bessey (1884). Such a puffball would produce approximately

552 CLASS BASIDIOMYCETEAE

160,000,000,000,000 spores. As in some of the Hymenogastrales the basal
portion of the spore fruit may remain sterile, not forming basidia in the
cavities which are produced in the sterile base. This is particularly charac-
teristic of the genus Lycoperdon in which the sterile base may be narrower
and resemble somewhat a broad stipe. In Calvatia the basal portion is
sterile in some species but not so markedly narrowed. The spore fruits of
all the species of this family appear to be edible when young, while still
white and rather brittle. The genus Lycoperdon has many species vary-
ing in size from 1 cm. to 5 cm. or more. The spore fruits are more or less
pear-shaped with a large sterile base. The exoperidium scales off as granules
or scales. The endoperidium has a single apical ostiole. They grow scattered
or in closely crowded masses on the ground or on decaying wood. (Fig.
182.) Calvatia differs externally from the preceding in the less pronounced
narrowing of the sterile base and in the absence of an ostiole in the endo-
peridium. The latter breaks off in large pieces. The capiUitium consists of
long, tangled, somewhat branched threads which usually break up into
short pieces at maturity. C. gigantea, occurring in the Fall in fields and
pastures, is collected for food while still firm and white. Bovista has a thin
exoperidium which sloughs off and a slightly thicker endoperidium with an
apical ostiole. There is no sterile base, the gleba filling the whole spore
fruit. The branched capillitial hyphae are slender and smooth and usually
not breaking into pieces. The sterigmata break loose from the basidium
and remain attached to the basidiospores. B. plumhea Pers. is 3 to 5 cm. in
diameter, nearly round and with a lead-gray endoperidium. Mycenastrum
has a rather thick endoperidium which cracks open in a more or less stel-
late manner. The capiUitium is composed of thick, branched, spiny
hyphae, tapering from the middle to the acute tips. The sterigmata are
very short so that the spores are almost sessile. Disciseda (Catastoma) has
a firm exoperidium which splits equatorially. The endoperidium pulls free
from the basal half of the exoperidium while remaining attached to the
upper half. The result is that the spore fruit escapes and blows around,
leaving the basal portion of the exoperidium still attached to the ground.
The ostiole pierces the endoperidium at the center of the exposed portion,
which is morphologically its base. Disciseda Candida (Schw.) Lloyd (C.
circumscissum) is common in grassy places in the prairie regions of the
United States and in similar regions in Eastern Europe. (Fig. 183.) Lan-
opiUi has a very thin peridium which separates in irregular pieces from the
gleba. This consists at maturity of a capiUitium of much entangled,
slender hyphae with the intermingled spores. The generic name was given
because of its resemblance to a ball of wool. Swoboda (1937) studied the
structure of the not quite mature spore fruits of L. bicolor (Lev.) Pat. and
found that the spores are borne on one side of the basidium on short

ORDER LYCOPERDALBS

553

Fig. 183. Order Lycoperdales, Family Lycoperdaceae, Disciseda Candida (Schw.)
Lloyd {Catastoma circumscissum (B. & C.) Morgan). Spore fruits in various stages of
dehiscence. (After Morgan, from Fischer, in Engler und Prantl; Die Natiirlichen
Pflanzenfamilien, Zweite Auflage, vol. 7a, Leipzig, W. Engelmann.)

sterigmata. The basidia are produced sympodially on branched hyphae of
a finely divided coralloid gleba, not on a hymenium clothing definite
cavities. He concluded that the genus should form the type of a family to
be named Lanopilaceae in the Sclerodermatales. Its abundant capillitium
and the position of the spores on the basidium would seem to indicate
possible relationship to Tiilostoma.

Two genera, Broomeia and Diplocystis, from the warmer parts of the
world, are characterized by the production of their spore fruits crowded
side by side on a stroma, which is thick and often with a stout stalk in the
former and thin and saucer-shaped in the latter. A few other genera are
recognized. In the mountains in the western part of the United States
occurs Calhovista, described by Miss Morse (1935). It resembles Calvatia
sculpta (Hark.) Lloyd, but has a capillitium resembling that of Bovista.
Zeller (1044) segregated four genera from the Lycoperdaceae to form
the Mesophelliaceae. These are Radiigera, so far only found in the United
States, Ahstoma, from New Zealand, Australia, and California, Meso-
phellia, from Europe and Australia, and Castoreum, from Australia. The
distinction is based on a usually three-layered peridium which is indehis-
cent or rupturing irregularly at the apex. In this group the hymenial
cavities are not well defined. The basidia are borne in clusters on short
branches of radial hyphae. This seems to indicate a tendency toward the
basidial arrangement found in Phellorinia and Podaxis.

Family Geastraceae. In most mycological works the genera included
in this family are placed in the Lycoperdaceae, but the author follows
Fischer (1933) in making the segregation. The chief distinction is that in
the latter the outer layer of the peridium lacks a fibrous layer and dis-
integrates at maturity, while in the Earthstars, as the Geastraceae are
called, the outer peridium does possess such a layer and splits stellately
from the top toward the base, spreading out in a star-like manner. The

554 CLASS BASIDIOMYCETEAE

Fig. 184. Lycoperdales, Family Geastraceae. Geastrum rufescens Pers. (Courtesy,
Coker and Couch: The Gasteromycetes of the Eastern United States and Canada,
Chapel Hill, Univ. North Carolina Press.)

chief genus is Geastrum.^ In this genus the inner peridium remains intact
when the outer peridium sphts open. It is sessile or on a short stalk and
has a single apical ostiole. (Fig. 184.) In Myriostoma the inner peridium
stands upon several slender stalks and there are several to numerous

ostioles.

In the two foregoing genera the gleba is like that of the Lycoperdaceae,
made up of tramal tissue with numerous closed, basidium-lined hymenial
cavities. There is usually but not always a columella. Formerly included
in this family was Astraeus hygromelricus (Pers.) Morg., in which the
chief difference is the partial obliteration of the hymenial • cavities by
ingrowing tufts of basidia. The outer peridium is exceedingly hygro-
metric, opening out when moist, closing to almost the original position
when dry. Because of the difference in glebal structure Fischer (1899,
1933) placed this genus in Family Calostomataceae in the Order Sclero-
dermatales. In view of the fact that in the family Sphaerobolaceae both
types of gleba are present without even leading to the division of the
family it appears best to retain Astraeus in the Geastraceae. (Fig. 185 A.)

The two remaining genera of the family were called by Fischer Geas-
teropsis and Trichaster, but Long (1945) pointed out that the former name
is not available nor is his own earlier name {Geasteroides, 1917) so that he
applied a new name Terroslella. This differs from Geastrum in the pos-

1 In most publications and in the author's earlier book this genus is called
Geasler. Since, however, the international rules of botanical nomenclature designate
Persoon's Synopsis Methodica Fungorum (1801) as the basis for the nomenclature
of the Gasteromyceteae his name Geastrum must be used, not the name preferred by
later mycologists.

ORDER LYCOPERDALES 555

session of a prominent sterile base, and a central ostiole on the endo-
peridium, whose whole upper portion soon is more or less caducous. In
Trichaster, apart from a subligneous columella there is no sterile base, and
the endoperidium has no ostiole and either falls off in pieces or adheres to
the exoperidium when the latter dehisces.

Fig. 185. Lycoperdales. (A) Family Geas-
traceae. Astraeus hygrometricus (Pers.) Morg. ;
portion of gleba showing partial obliteration
of hymenial cavities. (B) Family Tulosto-
mataceae. Tulostoma simulans Uoyd; portion
of gleba showing basidia with laterally pro-
duced spores. (Courtesy, Coker and Couch:
The Gasteromycetes of the Eastern United
States and Canada, Chapel Hill, Univ. North
Carolina Press.)

Family Tulostomataceae. This family is based upon the genus
Tulostoma} These fungi are popularly called the stalked puffballs. The
spore fruits of this family originate hypogeously but by the elongation
of a basal stalk become epigeous. The commonest genus is Tulosloma with
a spherical spore fruit 1 to 3 or 4 cm. in diameter and a slender stalk
sometimes 5 or more cm. in length and 2 to 4 mm. thick, often with a
small volva remaining at its base. (Fig. 186.) The endoperidium opens by
an apical ostiole which may be irregular or which may have a projecting
striate margin. The 40 or more species are mostly found in drier regions.
In some species the basidia are described as producing four basidiospores
laterally instead of apically. Whether this is true for the whole genus
or for all other genera of the family is not known. (Fig. 185 B.) The hy-
menial cavities in this genus and possibly other genera become obliterated
by the growth into them of hyphae bearing the irregularly arranged
basidia as in Scleroderma and Sphaeroholus stellatus (not as in Nidulari-

2 Persoon (1801) used this spelling instead of the more usual Tylostoma and for
this reason his spelling must be followed.

556

CLASS BASIDIOMYCETEAE

Fig. 186. Lycoperdales, Family Tulostomataceae. Tulostoma campestre Morgan.

opsis) and in Astraeus. For this reason the family is sometimes removed
from the Order Lycoperdales.

The stipe may be a rather broad and relatively short one or tall and
slender. In Tulostoma the rounded main body of the spore fruit reaches its
full size before a small mass of tissue at its base, and within the exoperid-
ium, begins its rapid elongation to become the stipe. The exoperidium
may remain as a fragmentary cup at its base and shred off from the main
portion of the body. The capillitium is abundant and grows fast to the
peridium. The basidia bear the spores, four in number, on short sterig-
mata laterally. This led von Tavel (1892) to suggest the origin of this
genus from Phleogena in the Auriculariales, with the loss of the cross septa
in the basidia of the latter.

Queletia mirabilis Fr., the only species of this genus, has been found in
France, England, and the United States, but appears to be rare. Like
Tulostoma its spore fruits develop underground. Only as the gleba ap-
proaches the spore-forming stage does the base start to elongate, breaking
through the peridium and forming a stout stipe 8 to 15 cm. tall and 3 to
4 cm. thick, bearing a rounded sporocarp 3 to 7 cm. in diameter. There is
a sharp line of distinction between the concave base of the latter and the
rounded apex of the stipe. The gleba is similar to that of Tulostoma, with
no trace of hymenial cavities at maturity. The 1 to 4 (mostly 3) basidio-
spores are terminal and lateral as in that genus. Normally the head breaks
away from the stipe and is blown about in the wind, scattering its spores,
but if the attachment is too firm the stipe shreds away and thus exposes
the under side of the powdery gleba which consists only of spores and
capillitium. The genus Calostoma was placed by Fischer (1933) in the
Family Calostomataceae along with Astraeus, with which it has little in
agreement except the plectobasidial type of gleba. In Calostoma the stipe

ORDER LYCOPERDALES

557

Fig. 187. Lycoperdales, Family Tulostomataceae. (A, B) Calostoma cinnabarinurn
Desv. (A) Spore fruit from which most of the volva has disappeared, through dehques-
cence. (B) Several basidia showing the sessile spores on all sides. (C, D) Battarrea
phalloides (Dicks.) Pers.; two specimens which grew in different environments.
(A-B, courtesy, Burnap: Botan. Gaz., 23(3):180-192, Univ. Chicago Press. C-D,
courtesy, Rea: Mycologia, 34(5):563-574.)

is mostly subterranean. The basidia have 5 to 12 sessile spores borne on
the apex and along the sides, in which they give some suggestion of their
relationship to Tulostoma. The capillitium is sometimes marked with an-
nular or spiral thickenings, resembling the elaters of some fungi. In the
author's opinion Calostoma probably belongs in the Tulostomataceae. In
this he follows Burnap (1897). (Fig. 187 A-B.)

Battarrea has a stout scaly stalk that may reach over 30 cm. in height
and with a large volva at its base. (Fig. 187 C-D.) The endoperidium of B.
phalloides (Dicks.) Pers. splits circumscissilely, rolling upwards a httle at
a time as the spores and capilUtium escape and in B. digueti Pat. & Har. is
perforated by many pores. The gleba in this genus contains typical hy-
menial cavities. It is found in the sandy or gravelly soil of the foothill
regions of the southwestern United States and in Europe, South America,
Austraha, etc. Rea (1942) recognizes two species in the United States with
one or the other of which perhaps all the other described species are syn-
onymous. There is some capillitium, evidently the remains of the tramal
tissues. In addition there are numerous elaters with annular or spiral
thickenings, reminding one of those of the Trichiaceae among the Myce-
tozoa or in the Hepaticae. In the younger, not fully developed specimens
single cells or several in succession in a hypha enlarge and become "lati-
ciferous." These do not become transformed into capillitial hyphae as in
Phellorinia. Clamp connections are numerous throughout the spore fruit.

558 CLASS BASIDIOMYCETEAE

Maublanc and Malengon (1930) made an extensive study of the anatomy
and development of the fungus.

Family Podaxaceae. Long and Stouffer (1946) place together in the
tribe Phellorinieae the genera Phellorinia, Didyocephalos, and Chlamy-
dopus. These three monotypic genera have their sporocarps elevated at
maturity on definite elongated stipes. The basidia are in fasciculate clus-
ters and remain undissolved when other tissues of the gleba undergo
autodigestion. Capillitial threads are present. The stipe may rarely extend
into the base of the sporocarp as a low columella in Phellorinia but more
often is absent. Probably to be associated with these is Podaxis in which
the stipe extends well up into the gleba, in most cases reaching clear up to
and uniting with the peridium at the apex. The persistent clustered
basidia are present as in the other genera. All four are hypogeous at first
and surrounded by a universal veil, part of which usually remains as a
volva at the base of the stipe and as quickly disappearing patches on the
sporocarp. In Phellorinia the universal veil is continuous as an exoperid-
ium and the outer layer of the stipe, rarely breaking away well up on the
stipe to show indications of a tightly adhering volva. The endoperidium
is an extension of the stipe surrounding the gleba for two-thirds or more of
its height, with a thin inner layer extending over its top. This falls away
at maturity. Malengon (1935) showed that in the immature gleba there
are numerous sinuously curved and branching hymenial cavities lined by
the basidial primordial cells. The latter branch sympodially and form
clusters of basidia on hyphae of various lengths which eventually obliter-
ate these cavities. This led earlier investigators who studied only the
practically mature fungi to classify them with plectobasidial forms such
as Sclerodermatales. At maturity the major part of the hyphae making up
the trama between the cavities dissolves, leaving spores and basidia and
the hyphae upon which these arose undissolved, as well as a few of the
thicker tramal hyphae which in the earlier stages of development served
as laticiferous tubes. These hyphae and those bearing the clusters of
basidia make up the so-called capillitium. This genus occurs in North and
South America, Europe, Asia, and Australasia.

Didyocephalos (see Long and Plunkett, 1940) occurs in the south-
western part of the United States and also northern Africa. There is but
one species, D. attenuatus (Pk.) Long and Plunk. {D. curvatus Underw.).
It originates 4 to 20 cm. beneath the surface of the soil and may attain a
height of 7 to 56 cm. with a head 5 to 13 cm. broad. The universal veil
breaks as the stipe elongates, leaving a volva at the base and a fleshy or
gelatinous exoperidium over the sporocarp. This at maturity becomes a
series of hairy scales which break off and expose the tough endoperidium.
The latter breaks away irregularly exposing the gleba. When young the
latter is cellular. Chlamydopus, with the single very variable species C.

ORDER LYCOPERDALES

559

ft- '"' V . ■ M\

1 r ■■ i ■ ■' ■ ■?ilJ

Fig. 188. Lycoperdales, Family Podaxaceae. Podaxis pistillaris (L. ex Pers.) Fr.
(A) Spore fruit, external view. (B) Somewhat diagrammatic longitudinal section of
spore fruit. (C) Basidium with sessile spores and single spore more highly magnified.
(D) Cluster of basidia on piece of capillitial thread. (After Fischer, in Engler und
Prantl: Die Naturlichen Pflanzenfamilien, Zweite Auflage, vol. 7a, Leipzig, W.
Engelmann.)

meyenianus (Klotzsch) Lloyd, has been found in North and South
America and Austraha (Long and Stouffer, 1946). In this genus the por-
tion of the universal veil which forms the exoperidium is verrucose and
fragile and brittle, usually breaking very early and exposing the tough
endoperidium which at maturity produces a single apical mouth. The
large volva is conspicuous. The stout clavate stipe is 4 to 15 cm. tall,
tapering toward the base and becomes woody to corky. The sporocarp is
depressed globose up to 30 mm. wide. The gleba shows no signs of having
been cellular. Podaxis has a slender woody stipe, arising from a narrow
volva and bearing a more or less pyriform or rounded head. The central

560 CLASS BASIDIOMYCETEAE

part of the stipe penetrates the gleba to the apical portion of the peridiiim,
rarely not quite reaching this far. The peridium pulls free from the stipe
below or splits near the latter exposing the powdery gleba. In 1933 Fischer
placed Podaxis in a separate family but in 1934 he recognized its close
relationship to Phellorinia. Miss Morse (1933) made an extensive study of
Podaxis and came to the conclusion that the various species that have
been described are all based upon specimens of various ages and grown
under extremes of habitat. She recognizes therefore only the one species
P. pistillaris (L. ex Pers.) Fr. (Fig. 188.)

Key to the Orders and Families of Gasteromyceteae

Small, hypogeous or epigeous, with a single hymenial cavity lined by an even
hymenium. Order Protogastrales

No columella, hymenial cavity more or less spherical.

Family Protogastraceae
Hymenial cavity annular, surrounding the percurrent columella.

Family Hemigastraceae
Small to large, hypogeous or epigeous, with one to many hymenial cavities pro-
duced in the lacunar or coralloid manner. If only one hymenial cavity
is formed that is lined at maturity by a folded or lobed hymenial
layer. In some genera the cavities are filled with a gelatinous sub-
stance into which the basidia project.
Gleba at maturity not undergoing much change. With or without a stipe.
Hymenial cavities typical or gelatine-filled.

Order Hymenogastrales
Columella mostly lacking (present in some of the Hysterangiaceae and in
Gasterellopsis in the Hymenogastraceae) .
Hymenial cavities typical, lined by hymenium.
Gleba fleshy, not clearly coralloid in development.

Family Hymenogastraceae
Gleba cartilaginous to gelatinous, plainly coralloid.

Family Hysterangiaceae
Hymenial cavities filled with gelatinous substance or with irregular masses
of basidia. Family Melanogastraceae

Columella reaching the apex and spreading to form the pileus, no marked
stipe at maturity. Development pseudoangiocarpic.

Family Hydnangiaceae
Columella produced downward below the pileus to form a distinct stipe.
Gleba free from the stipe at maturity, at least below.

Family Secotiaceae
Gleba at maturity deliquescing to a slimy, usually evil-smelling mass, which
covers or is supported by a definite framework (the receptacle).

Order Phallales
Receptacle lattice-like or irregularly branched or lobed, with or without a

stipe. Family Clathraceae

Receptacle occupying the upper portion of a stout, hollow stipe, either grown
fast to it or forming a bell-shaped structure (pileus) attached at the
top of the stipe. Family Phallaceae

Medium to large, mostly epigeous at maturity and usually with a thick peridium.
Hymenial cavities lacunar in origin, obliterated or replaced by nests

KEY TO THE MORE IMPORTANT GENERA OF GASTEROMYCETEAE 561

or clusters of basidia. At maturity the whole gleba or portions of it
become a powdery mass of spores with only rudimentary or lacking
capillitium. Order Sclerodermatales

Only family. Family Sclerodermataceae

Medium-sized to small, not hypogeous at maturity or not at all. Hymenial cavities

with definite lining of basidia. Toward maturity the tramal tissue

surrounding each cavity encloses it in a thin or thick wall, producing

separate structures called peridioles. Order Nidulariales

Peridioles very numerous, with thin walls, escaping by irregular breaking of the
thin peridium. Family Arachniaceae

Peridioles few, with thick hard walls. Peridium beaker-like, opening by rupture
of the diaphragm-like top, leaving the peridioles like eggs in a nest.

Family Nidulariaceae
Small (up to 5 mm.), mostly growing on decayed wood or on manure, the whole
gleba being expelled as a single ball by the eversion of the inner layer
of the thick peridium which splits stellately at the top. Gleba with
many distinct hymenial cavities or these obliterated by the ingrowing
basidia. Order Sphaerobolales

Only family. Family Sphaerobolaceae

Medium-sized to large, epigeous, at least at maturity (a very few exceptions).
Gleba mostly with definite hymenial cavities, or these sometimes ob-
literated. At maturity the gleba becomes a dry powdery mass of spores
and capillitium, in one family the basidia also remaining intact.

Order Lycoperdales

Without stipe. Outer peridium shedding in patches or granules, inner peridium
mostly thin (thick in Mycenastrum) , opening by a mouth (ostiole) or
by breaking away in pieces. Columella sometimes present.

Family Lycoperdaceae

Without stipe. Outer layers of the peridium splitting and turning back stel-
lately. Inner peridium opening by an ostiole or falling away in pieces.
Columella mostly present. Family Geastraceae

With stipe. Basidia lining definite cavities or these more or less completely
obliterated. At maturity the whole gleba except capillitium and spores
dissolving into a powdery mass. Family Tulostomataceae

With stipe. Basidia occurring in clusters on glebal hyphae. At maturity the
basidia and their supporting hyphae are not destroyed but make
up, with the spores and capillitium, the powdery contents of the
sporocarp. Family Podaxaceae

Key to the More Important Genera of Gasteromyceteae

Family Protogastraceae.

Single genus, United States. Protogaster

Family Hemigastraceae.

Single genus, Europe. Hemigaster

Family Hynienogastraceae.
No percurrent columella.

At maturity a single hymenial cavity with folded or lobed hymenial lining.

United States. Gasterella

At maturity many hymenial cavities.

Peridium almost absent at maturity. Spores colorless or pale brown, spiny
or verrucose. North America, Australia. Gymnomyces

562 CLASS BASIDIOMTCETEAE

Peridium well developed at maturity.

Sporocarp without root-like mycelial strands.
Spores ellipsoid, ovoid or fusiform.

Trama plates irregular or radiating somewhat from a small sterile

base. Old and New World. Hymenogaster

Trama plates arising from a dendroidally branching axial strand
(possibly better assigned to the Hysterangiaceae). Europe.

Dendrogaster
Spores spherical, spiny.

No sterile base, but glebal chambers converging toward the center

of the base. Europe. Martellia

Glebal chambers converging toward a sterile base. Europe, North
America, Australia. Odaviania

Sporocarp with root-like mycelial strands.

Spores spherical, verrucose, cystidia present. Europe and North

America. Sclerogaster

Spores ellipsoid, smooth, no cystidia. Europe, North and South

America, Asia, Australia. Rhizopogon

Spores angular, no cystidia. North America. Nigropogon

Percurrent columella. Annular glebal cavity divided by centripetally develop-
ing plates into several lobes or separate cavities. North America.

Gasterellopsis
Genera of doubtful relationship, fusiform to pyriform, sometimes shortly
stalked: Gymnoglossum, Protoglossum, LeRatia, Clavogaster.
Family Hysterangiaceae.
Spores spiny or ribbed.

Peridium wanting at maturity, spores oblong, ribbed. Europe, North Africa,

and North America. Gautieria

Peridium present at maturity, spores spherical.
No gelatinous layer beneath exoperidium.

Basidia two- to four-spored. Europe. Maccagnia

Basidia five- to six-spored. Java. Hoehneliogaster

A gelatinous layer present beneath the thin exoperidium. Basidia two-
spored. East India. Clathrogaster

Spores smooth, ellipsoid or rod-shaped.

Columella a basal cushion or a mycelial strand, often branched, not per-
current to the apex.
Peridium with large gelatinous outgrowths. Australia. Phallobata
Peridium more or less uniformly thick.

Gleba at maturity collapsing into a thin layer on the inner surface of the

peridium. Australia. Gallacea

Gleba not collapsing.

Branches of the columella not dividing the gleba into sharply bounded
portions.
Sporocarp sessile. Europe, Africa, North and South America,

Australia. Hysterangium

Sporocarp stalked. Europe. Jaczewskia

Branches of the columella dividing the gleba into sharply bounded
portions.
Sporocarp tuberoid, sessile. South America, Ceylon.

Protuhera
Sporocarp pyriform, stalked. North America. Phallogaster

KEY TO THE MOEE IMPORTANT GENERA OF GASTEROMYCETEAE 563

Columella unbranched, percurrent to the apex. North America.

Rhopalogaster
Family Hydnangiaceae.

Spores ovoid to fusiform, with longitudinal furrows. Europe. Chamonixia
Spores spherical, spiny, laticiferous vessels wanting. Europe. Hydnangium
Spores spherical or oblong, spiny or verrucose, laticiferous vessels present.
Europe, North America, South America. Arcangeliella

Family Melanogastraceae.

Gleba not clearly chambered, basidia in nests between sterile veins. Africa.

Corditubera
Gleba with numerous cavities which become filled with hyphae or gelatinous
substance.
Without stipe.

Chambers loosely filled with hyphae among which the basidia are placed,

spores roughened. North Africa. Chondrogaster

Chambers formed by enlarged cells which then gelatinize. On hyphae
traversing the chambers clusters of basidia arise. Spores smooth.
North America. Alpova

Basidia forming an irregular hymenium surrounding the hymenial cavities
filled with loose hyphae or gelatine.
Spores ellipsoidal, brown. Europe and North America. Melanogaster
Spores spherical, almost colorless. Europe and North America.

Leucogaster
With stipe at maturity, the exoperidium remaining as a basal volva and as
shreds on the sporocarp which is hemispherical, and concave
below. Relationship uncertain. Torrendia

Family Secotiaceae.

Fleshy. Development pseudoangiocarpic. Spores hyaline or light-colored, spiny
or reticulate.
Numerous groups of pseudoparenchymatous cells in the tissues. Cystidia

present. Europe and North America. Elasmomyces

Groups of pseudoparenchymatous cells wanting. South Africa.

Macowanites
Firm, stipe almost woody. Development angiocarpic. Gleba chambers more or
less radially elongated, sometimes lamelloid. Spores colored at
maturity.
Lower edge of the peridium pulling loose from the stipe, leaving no volva
(at least not conspicuous), cystidia often present. World-wide in
its distribution. Secotium

Lower edge of peridium pulling loose, leaving a distinct annulus but no volva.

Western United States. Longula

Lower edge of slender conical peridium pulling loose and leaving a cortina,
but no annulus or volva. Spores resembling those of Galerula and
Bolbitius. Europe, Asia, United States. Galeropsis

Peridium opening circumscissilely, leaving a large volva. Europe, Africa,
North and South America, Asia. Gyrophragmium

(including Polyplocium)
Peridium leaving a large volva, persisting on the pileus only as a small central
patch beyond which the slender radial glebal lamellae project.
No distinct annulus. Europe, Africa, North America, Australasia.

Montagnea
{Montagnites)

564 CLASS BASIDIOMYCETEAE

Family Clathraceae.

Receptacle lattice-like or of meridionally curved branches united at their tips.
Without stipe, not strongly narrowed below.

Branches of the lattice-like receptacle heavy and thick. Europe, United

States, Ceylon. Clathrus

Branches of the lattice-like receptacle slender, meshes large. Tropics of

Australia and Asia. lleodictyon

Meridionally curved branches heavy. Tropical regions of Old and New

World. Colonnaria

Meridionally curved branches slender. West Indies. Laternea

With stipe or strongly narrowed below.

No true stipe, lower meshes like the upper ones or slightly elongated.

Tropics of Old and New Worlds. ClathreUa

Stipe short. Lower series of meshes much elongated, upper ones isodia-

metric. Mediterranean region. Colus

Stipe tall, stout. Lattice-like receptacle with isodiametric meshes. Tem-
perate and tropical regions of Old and New Worlds.

Sirnblum
Stipe distinct. Several arms bowed out and united at the tip. Warmer
regions of Old and New Worlds. Pseudocolus

Receptacle stipitate, of arms not united at their tips at maturity.

Arms spreading horizontally from the margin of a disk-like widening of the
upper end of the stipe. Tropical regions of Old and New Worlds.

Aseroe
Arms vertically parallel, spreading at their tips.

Arms surrounded by gleba laterally, and wholly or partially dorsally.

Australia, Asia, America, Europe. Lysurus

Gleba only on inner side of arms. Africa, Australia, Europe.

Antkurus
Arms projecting in all directions, knobby. Africa. Kalchbrennera

Receptacle stipitate and surrounding the gleba, splitting stellately into five

lobes. New Zealand. Claustula

The tropical genera Blumenavia and Mycopharus closely resemble Colonnaria
and Pseudocolus, respectively.
Family Phallaceae.

Receptacle closely clothing the upper part of the stipe, not on a campanulate
pileus.
No pseudoparenchymatous ridges or reticulations on the gleba at maturity.
Gleba covering the tip as well as the upper portion of the stipe. South

America. Xylophagus

Gleba forming a belt some distance below apex of stipe. South America.

Staheliomyces
Gleba closely investing the upper portion of the stipe but not overrunning
the tip. Old and New Worlds. Mutinus

Like Mutinus, but with pseudoparencliymatous projections or ridges from

the gleba. East Indies and Australasia. Jansia

Like Mutinus, gleba invested with a loose net. Africa. Floccomutinus

Receptacle forming a campanulate pileus, attached centrally at the upper end
of the stout stipe.
Indusium growing from between pileus and stipe.

Pileus perforated like lattice-work. Volva mostly spiny. East Indies.

Echinophallus

KEY TO THE MORE IMPORTANT GENERA OP GASTEROMYCETEAE 565

Pileus not perforated like lattice-work. Volva not spiny. Tropics of Old and
New World, one species in temperate North America.

Dichjophora
No indusium between pileus and upper portion of stipe.

Pileus and gleba continuous over the apex of the stipe, leaving no apical

perforation. Brazil. Aporophallus

Apex of pileus perforate, rarely covered temporarily by a fragment of the

volva. Most regions of the world. Phallus

Apex of the pileus covered by a sort of cap, the receptacle, with many lobes
or branches so that at maturity, after the gleba has disappeared,
it resembles a wig. Southwestern United States and tropical and
temperate South America. Ilajahya

Family Sclerodermataceae.

Gleba marked by veins into many distinct regions. Capillitium rudimentary.
Gleba breaking up into a powdery mass, the veins remaining or disappearing.
Peridium firm, thick.
Peridium covered externally by conical spines. The spiny spores borne on

long sterigmata. East Indies. Caloderma

Peridium not spiny, thick.

Spores before maturity surrounded by a coat of hyphae. All over the

world. Scleroderma

Spores without hyphal sheath. Europe. Pompholyx

Gleba breaking up into numerous separate, thin-walled peridioles. Peridium
thin, falling to pieces at maturity. Europe, America, Asia,
Australasia. Pisolithus

Gleba not marked by veins, capilhtium well developed. Peridium simple, thin,
opening by an apical pore. Spores borne laterally on the basidia.
_ Europe. Glischroderma^

Family Arachniaceae.

Single genus. Americas, Africa, Australasia. Arachnion

Family Nidulariaceae.

Peridioles without funiculus.

Spore fruit roundish, without typical epiphragma. All continents.

Nidularia
Spore fruit beaker-formed, with epiphragma. North and South America,
Asia, Australasia. Nidula

Peridioles with funiculus.

Spore fruit cup-shaped, peridium of one layer, peridioles with a thick white

tunica. All continents. Crucihidum

Spore fruit bell- or goblet-shaped, peridium of three layers, tunica of perid-
ioles thin, hence their color, black or gray. All continents.

Cyathus
Family Sphaerobolaceae.

Peridium of three layers, including the apical region. Widely distributed.

Sphaeroholus
Middle layer of peridium lacking in apical region. Europe and North America.

Nididariopsis
Family Lycoperdaceae (including Mesophelliaceae). The little known tropical or
African genera, Lycoperdopsis, Lasiosphaera, Hippoperdon, and
Bovistoides, are not included in the key.

3 This genus is put in a separate family, Glischrodermataceae, by Rea (1922)
and Fischer (1933).

566 CLASS BASIDIOMYCETEAE

Sporocarps single or gregarious, not on a stroma.
Inner peridium thin, opening variously.

Capillitium of more or less uniform hyphae, branched or simple.

Outer peridium separating in granules or flakes from the inner peridium.
Inner peridium breaking up into flakes or fragments.

Capillitial hyphae interwoven into a woolly ball. North and South

America, Africa. Lanopila

Capillitial threads not tightly interwoven, breaking into short
pieces at maturity. In most continents. Calvatia

Inner peridium opening by an apical pore. The world over.

Lycoperdon
Outer peridium firm, remaining attached to the upper half of the inner
peridium and splitting circumscissilely. Inner peridium break-
ing free from the basal portion of the outer peridium, open-
ing by a pore in the originally basal portion. Europe, America,
Australasia.

Disciseda
(Catastoma)
Outer peridium firm, remaining attached to the inner peridium. At first
hypogeous in most cases. The whole peridium rupturing irregu-
larly. (This is Family Mesophelliaceae of Zeller.)
Spores spherical, echinulate, reticulated or verrucose.

Gleba without a sterile base. Australia and Western United States.

Absto77ia
Gleba with a sterile base. Western United States. Radiigera
Spores eUipsoidal, smooth or irregularly roughened.
Gleba with a central core. Australasia and Europe.

Mesophellia
Gleba without core. Australasia. Castoreum

Capillitium hyphae much branched, consisting of a thicker main stem and
tapering branches.
Inner peridium opening by apical pore.

At maturity attached to the ground. Eurasia and Australasia.

Bovistella
At maturity breaking loose and blown by the wind. Europe, North
America, and Australasia. Bovista

Inner peridium breaking up in flakes. Western United States.

Calbovista
Inner peridium thick and corky, opening irregularly or somewhat stellately.
Capillitium threads thick, short, spiny, with numerous thorn-like
processes. The world over. Mycenastrum

Sporocarps perched close together on a stroma.

Stroma thick, often columnar. South Africa. Broomeia

Stroma shallow, thin, shell-like. South Africa and West Indies.

Diplocystis
Family Geastraceae.

Columella wanting. Hymenial chambers lacking (plectobasidial). Europe and

North America. Astraeus

Columella present. Mostly with typical hymenial chambers.

A prominent sterile base in addition to the columella. Endoperidium with an
apical pore but soon the whole upper part more or less caducous.
America and Africa. Terrostella

(Syn., Geasteropsis and Geasteroides)

LITERATURE CITED 5G7

No sterile base in addition to columella.

Endoperidium sessile or on a single short stalk, opening by a single apical

pore. All parts of the world. Geastrum

Endoperidium on several slender stalks, opening by several pores. Europe,

South Africa, North and South America. Myriostoma

Endoperidium with no pore, breaking away in pieces or adhering in pieces
to the dehisced exoperidium. Europe and South Africa.

Trichaster
Family Tulostomataceae.

Stipe (largely subterranean) stout, composed of parallel or interwoven carti-
laginous strands growing from the base of the endoperidium.
Basidia bearing 5 to 12 sessile spores. Ostiole stellately bordered
by several, usually colored, lobes. North and South America,
Asia, East Indies, Australia. Colostoma

Stipe stout or slender, of parallel hyphae, sometimes scaly with age.

With elaters marked with rings or spirals. Gleba with hymenial cavities. At
maturity stipe with a large volva and covered with overlapping
scales.
Sporocarp low bell-shaped (convex above and concave below). Widely

distributed. Battarrea

Sporocarp spherical (by some considered a species of Battarrea). Africa.

Sphaericeps
Elaters lacking, but with typical capillitium. Hymenial cavities obliterated.
Inner peridium with apical mouth. Stipe slender. Widely distributed.

Tulostoma
Inner peridium opening by stellate lobes. Stipe slender. Africa.

Schizostoma
Inner peridium opening irregularly. Stipe stout. Europe and United States.

Queletia
Family Podaxaceae.

Columella wanting or very low.
With volva.

Dehiscing by an apical pore. North America, Africa, Australasia.

Chlamydopus
Dehiscing by irregular rupture of upper portion of endoperidium. W^estern
North America, and Africa. Dictyocephalos

Volva wanting; urceolate at maturity. North and South America, Africa,
Asia, Australasia. Phellorinia

Columella continuing as an extension of the stipe to or nearly to the top of the
sporocarp.
Peridium pulling loose at its lower edge. Spores sessile. North America,

Africa, Asia, Australia. Podaxis

Peridium remaining attached below, splitting laterally. Spores on sterigmata.
Australia. (Sometimes considered a form of Podaxis.)

Chainoderma

(For a more detailed key to the genera of the Gasteromyceteae the student is
referred to the latest publication by the late Dr. S. M. Zeller (1949). His arrange-
ment is not entirely in agreement with the ideas of the author.)

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VON Tavel, Franz: Vergleichende Morphologie der Pilze, 208 pp. 90 figs. Jena,

Gustav Fischer, 1892.
Tulasne, Louis Rene: Fungi hypogaei. Histoire et monographic des champig-
nons hypog^s, xix + 222 pp. 21 pis. (9 colored). Paris, Fr. Klincksieck, 1851.

(Second edition in 1863.)
Walker, Leva B.: Development and mechanism of discharge in Sphaerobolus

iowensis n. sp. and S. stellatus Tode, /. Elisha Mitchell Sci. Soc, 42(3-4) :151-

178. Pis. 16-25. 1927.
: Development of Gasterella lutophila, Mycologia, 32(l):31-42. Figs. 1-45.

1940.
White, V. S. : The Tylostomaceae of North America, Bull. Torrey Botan. Club,

28(8) :421-444. PZs. 31-40. 1901.
Zeller, S. M. : Protogaster, rei)rescnting a new order of the Gasteromycetes, Ann.

Missouri Botan. Garden, 21(2) :23 1-240. 2 pis. 1934.
• : Developmental morpliology of Alpova, Oregon State Monographs.

Studies in Botany, 2:1-19. Pis. 1-4. 1939.
: Representatives of the Mesophelliaceae in North America, Mycologia,

36(6) :627-637. Figs. 1-6. 1944.
— : A new name, Mycologia, 37(5) :636. 1945.

LITERATURE CITED 571

— : Notes on certain Gasteromycetes, including two new orders, ibid.,

40(6) :639-668. 1948.
— : Keys to the orders, families, and genera of the Gasteromycetes, ibid.,

41(l):36-58. 1949.
— , AND Carroll W. Dodge: Gautieria in North America, Ann. Missouri

Botan. Garden, 5(2) :133-142. PL 9. 1918.
— , AND : Leucogaster and Leucophlebs in North America, ibid.,

11(4):389-410. P/. 11. 1924.
- — ■, AND Leva B. Walker: Gasterella, a new uniloculate Gasteromycete,

Mycologia, 27(6) :573-579. Figs. 1-13. 1935.

FUNGI IMPERFECTI: THE IMPERFECT FUNGI

THERE are a great many species of fungi of which the perfect stage is not
known and which therefore cannot find a place in the classes already
discussed. By the term ''perfect stage," as here used, is meant that stage
in which the ultimate sexual structures are formed, e.g., zygospores,
oospores, asci, basidia, and teliospores. Most of the Phycomyceteae are
so characteristic in their mycelial structure as well as in their modes of
asexual reproduction that ordinarily the genus and often even the species
can be determined from the asexual stage alone. Thus the Imperfect
Fungi are practically confined to those Higher Fungi in which the stage
is lacking in which the asci, basidia, or teliospores are produced. Since
most of the Uredinales have very characteristic asexual stages the im-
perfect forms of this order are readily assigned to that group and are
placed in one of the imperfect genera there, e.g., Aecidium, Uredo, Caeoma,
etc., if their host requirements and other features make it impossible to
assign them to described species in recognized perfect genera of that order.
Thus it comes about that the Fungi Imperfecti, as ordinarily considered,
include those fungi not otherwise referable to their natural relationship
(e.g., Phycomyceteae or Uredinales) whose true relationship cannot be
determined in the absence of the perfect stage. Judging by the rather
exceptional presence of clamp connections, as well as by the similarity of
the conidial stages to those in the Class Ascomyceteae, it is probable that
a great majority of species of imperfect fungi really belong to that class,
and the perfect stage is not present in the specimens examined. It is
possible that some fungi have lost entirely the power to produce a perfect
stage and so are truly imperfect fungi. A few assigned to this class are
doubtless imperfect stages of Ustilaginales or other groups of Basidio-
myceteae. In the first edition of Engler and Prantl, "Die Natiirlichen
Pflanzenfamilien," Lindau (1899, 1900) recognized about 600 genera and
15,000 to 20,000 species. In a more recent, as yet unpublished, work on
this group Dr. Harold B. Bender (1931) recognized as valid 1331 genera.
Since for a great many fungi the asexual and sexual stages of repro-

572

FUNGI IMPERFECT! : THE IMPERFECT FUNGI 573

duction may be separated in time and substratum, many fungi have been
described under different names according to whether one or the other
stage was studied. As time goes on the connection between the two stages
is recognized in many cases. Theoretically, therefore, the asexual stage
should cease to be known by its name among the Imperfect Fungi and it
should no longer be included in that group. Practically, however, it is
desirable to retain this name among the Fungi Imperfecti since it would
be sought there in attempts to identify it, unless the perfect stage were
found along with it. Thus we still seek for such genera as Aspergillus,
Penicillium, Sphaceloma, Ramularia, Cercospora, etc. in the manuals
describing the Imperfect Fungi although the perfect stages of many
species of these genera are known and provided with names.

The distribution of the many thousand species of Fungi Imperfecti
into genera, families, and orders must necessarily be based upon vegetative
and asexual reproductive structures instead of upon the perfect reproduc-
tive stages. Inasmuch as it has been demonstrated that fungi whose per-
fect stages show them to be of very different families may possess rather
similar types of asexual reproduction it follows that genera based upon
the asexual reproductive forms are not necessarily assemblages of related
species. As an example attention may be drawn to the genus Gloeosporium.
In this genus the one-celled, hyaline, ellipsoidal, straight or slightly curved
conidia are produced, usually embedded in a gummy substance, from
short conidiophores packed in a palisade underneath the host epidermis
which is ruptured by the developing mass of conidia. This acervulus type
of asexual reproduction is found in some species of Gnomonia and Glom-
erella, both being genera in the Gnomoniaceae of the Sphaeriales. The
fungus commonly known as Pseudopeziza ribis Kleb., of Family Mol-
lisiaceae, Order Pezizales, also has a similar type of asexual reproduction.
A number of similar cases are known. Since, then, the genera based upon
asexual structures do not necessarily indicate true relationships of the
included species the term "form genus" was suggested for such groups by
Schroeter. It is in this sense that the term genus is used in this class. On
the other hand, as has been pointed out by several authors including
Petrak and Sydow (1926-1927) there is frequently a similarity in asexual
structures among fungi considered to be closely related as judged by their
perfect stage. Careful study has revealed that in many cases some of the
form genera of the Imperfect Fungi can be subdivided into groups of
species correlated with the perfect stages. Thus has come about on the
part of some mycologists the breaking up of the larger genera into smaller
more compact ones, on characters that would otherwise be considered of
rather minor importance except for their correlation with groups of
perfect fungi.

The fact that many Imperfect Fungi possess several different spore

574

FUNGI IMPERFECTi: THE IMPERFECT FUNGI

Fig. 189. Moniliales, Family Moniliaceae. Various types of adaptation for the
production of submerged spores. (A) Varicosporium elodeae Kegel. (B) Tetracladium
marchalianum De Wild. (C) Clavariopsis aquaiica De Wild. (D) Tetrachaetum elegans
Ingold. (E) Lunulospora curvula Ingold. (Courtesy, Ingold: Brit. Mycol. Soc. Trans.,
25(4):339-417.)

FUNGI IMPERFECTi: THE IMPERFECT FUNGI 575

forms makes their recognition as definite species difficult, even where the
perfect stages are not discovered. Furthermore the adaptation of the type
of spore or sporophore to special habitats may be responsible for similar-
ities among these that perhaps do not reflect real relationships. Thus the
fungi that live in wet habitats and frequently produce their spores on
hyphae that are completely submerged show many points of likeness; as
in the genera Varicosporium, Tetracladium, Heliscus, Lemonniera, Tri-
cladium, Tetrachaetum, Lunulospora, etc. The spores are usually slender
and with long branches which give them a great abihty to float. In some
genera these are aleuriospores, in the sense of Vuillemin (1910, 1911) and
in others are phialospores and radulaspores, in the sense of Mason (1933,
1937). To these Ingold (1942) would add the type "aquatic spore." A
phialospore is borne at the apex of a phialide, a "fusiform truncate, fusi-
form beaked or acuminate terminal portion of a hypha, from the apex of
which, or within which, thin-walled conidia are abstricted" (Mason). The
septum separating the spore from the phialide is not produced until the
spore is fully grown. An aleuriospore is a terminal portion of a hypha that
is early separated by a septum from the parent hypha. A radulaspore is
according to Mason a type "in which each spore is borne on a little
sterigma, without any reference to the growing-point of a hypha," as in
Botrytis cinerea. There is no direct evidence that these various aquatic
genera are closely related except in habitat. Ingold (1942, 1943, 1944) has
studied many of these and described several new genera. (Fig. 189.)

As the life histories of various fungi are studied by pure culture
methods or by means of carefully controlled inoculations, from time to
time an imperfect fungus is connected up with its perfect stage. This may
perhaps be a species already known or may have been unknown there-
tofore. Klebahn (1918), the German mycologist, has made many such
connections. It often happens that the same species has several types of
asexual reproduction so that it may appear in several different form
genera.

The many species and genera are usually divided into four form orders
as follows :

Sphaeropsidales: conidia produced within pycnidia or niodifications of such
structures. A pycnidium is a perithecium-like structure and may be com-
plete, like the perithecium of the Sphaeriales and Hypocreales, or only the
top half may be present as in the perithecium-like spore fruit of the Hemi-
sphaeriales, or it may , open by a longitudinal slit as in the apothecia of
the Hysteriales or may be closed at first and finally open into a cup or
saucer-shaped structure, much like a miniature apothecium.

Melanconiales: conidia produced singly or in chains, often surrounded by a
gummy mass, from conidiophores packed closely in a usually subepidermal
or subcortical layer, the acervulus.

MoniHales: conidia formed on conidiophores which are separate, at least at

576

FUNGI IMPERFECTi: THE IMPERFECT FUNGI

their apical portions, or the vegetative mycelium breaking up into conidia.
The conidiophores may] be simple or branched, short or long, similar to
the ?,vegetative mycelium or very distinct from it, but are never enclosed
within a pycnidium nor packed laterally into a subepidermal or subcortical
acervulus. They are almost always external at time of conidium production.
Mycelia Sterilia: imperfect fungi which lack all conidial formation, and which
produce sclerotia, rhizomorphs, and various other forms of mycelium without
spores.

The usual extremely artificial classification of the Fungi Imperfecti
separates many genera which perhaps more logically should be placed

nearer each other. An example is the
series of fungi in which the conidia
("endoconidia") are produced in the
interior of the conidiophore and pushed
out successively from an opening at the
apex. These genera are found in several
different "form families" and have
conidia that are colored or colorless; one-
celled to several-celled; pushed out in
chains or singly; surrounded by slime or
not, etc. In some of these genera other
imperfect spore-forms also occur while
in some the endoconidia are the only
ones known. Some are believed to be
imperfect forms of Ascomyceteae while
it is suspected that the perfect stage of
others may be Basidiomycetous. The
following genera include most of the
endosporous Fungi Imperfecti: Cado-
phora, Thielaviopsis, Hymenella, Chalara,
Sporoschisma, Sporendonema, Endoco-
nidium, Chalaropsis, and probably Caten-
ularia and Phialophora. (Fig. 190.) In the last two the conidiophore
approaches the type of phialide (or sterigma) found in Cephalosporium,
Gliocladium, and Penicillmm, in which the conidia appear to be almost
endogenous in origin.

Order Sphaeropsidales. The 568^ genera (with over 2300 species in
North America alone) ascribed to this order are divided into four form

families.

Family Sphaeropsidaceae (Sphaerioidaceae of Some Authors).
Pycnidia resembling typical perithecia or forming pycnidial cavities in a

Fig. 190. Moniliales. Endoge-
nous production of conidia (endo-
spores). (A) Cadophora obscura
Nannfeldt. (B) Thielaviopsis para-
doxa (de Seynes) von Hohnel (co-
nidial stage of Ophiosloma {Cera-
tostomella) paradoxum). (A, after
Melin and Nannfeldt, Svenska
Skogsvardsforeningens Tidskrift,
Hafte III-IV, pp. 397-616. B, cour-
tesy, Dade: Brit. Mycol. Soc.
Trans., 13:184-194.)

1 The figures for this order are taken from H. B. Bender's pamphlet (1934) on
the Sphaeropsidales.

ORDER SPHAEROPSIDALES 577

stroma; tough leathery to brittle, and dark-colored. The spores often
exude from the ostiole in damp weather in a worm-like mass, or cirrhus,
consisting of gum and embedded spores. This family contains 359 genera
so that it has become necessary to devise some means of subdividing it in
a practical and easily applied manner. The scheme most often used was
suggested by the great Italian mycologist P. A. Saccardo (1899) and is
based on the structure and the color of the spores as follows :

Amerosporae: spores one-celled, spherical, ovoid or somewhat elongated.

Hyalosporae: spores hyaline.

Phaeosporae : spores colored some shade of light brown to black.
Didymosporae: spores similar to the foregoing, but two-celled.

Hyalodidymae: spores hyaline.

Phaeodidymae: spores colored.
Phragmosporae: spores three- or more-celled by transverse septa.

Hyalophragmiae: spores hyaline.

Phaeophragmiae: spores colored.
Dictyosporae : spores divided by both transverse and longitudinal septa.

Hyalodictyae : spores hyaline.

Phaeodictyae: spores colored.
Scolecosporae: spores very slender, thread- or worm-like, one-celled to several-
celled, hyaline or colored.
Helicosporae: spores cylindrical and more or less spirally coiled, one-celled to

several-celled, hyaline or colored.
Staurosporae: spores radiately lobed or star- or cross-shaped, one-celled to
several-celled, hyaline or colored.

This same scheme is used for other orders and families of Imperfect
Fungi in which conidia are produced, with the omission of such sub-
divisions as are not represented. Where only a few forms are present
in the major subdivisions the minor ones based on spore color are often
omitted. In the discussion of representative genera of this class the
group name, based on the foregoing scheme, follows the generic name in
parenthesis.

The following genera should be mentioned: Phyllostida (Fig. 191A-C),
Phoma, Dendro'phoma (Fig. 191D-E) and Macrophoma (all Hyalosporae).
Their 2500 or more species are parasitic on leaves and stems of plants. A
possible exception is the fungus named by L. R. Fitzgerald (1943) Phoma
stenohothri (Holl. & Mor.) Fitzg., which is parasitic upon grasshoppers.
The first genus produces leaf spots with definite borders, the others pro-
duce less definite spots and occur on other parts of the host as well.
Macrophoma, as the genus is usually defined, but incorrectly, according
to Petrak and Sydow (1926-1927), has large conidia, over 15 m long, but
otherwise is like the others. In these genera the pycnidium is buried in the
host tissue but the short ostiole emerges at maturity. The distinctions
between these genera are clearly entirely artificial. A leaf spot of the beet
(Beta vulgaris L.) is caused by Phyllostida tahifica Prill. The same fungus

578

FUNGI IMPERFECTi: THE IMPERFECT FUNGI

KV'rir^

Fig. 191. Sphaeropsidales, Family Sphaeropsidaceae. (A-C) Phyllostida tabifica
Prill. (A) Pycnidium viewed from above, showing cirrhus of spores. (B) Vertical sec-
tion through pj^cnidium. (C) Spores. (D, E) Dendrophoma convallariae Cav. (D)
Pycnidium from above. (E) Conidiophores. (F) Ascochyta dianthi (A. & S.) Berk.;
section through pycnidium. (A-C, after Prillieux: Bull. soc. mycol. France, 7(1):15-19.
D-E, after Briosi and Cavara: Fascicle IV, No. 89. F, ibid., Fascicle XIV, No. 342.)

on the roots and inflorescence has been called Phoma betae Fr. Various
species of these genera have been shown to have as their perfect stage
species of Guignardia, Mycosphaerella, etc., in the Sphaeriales or Pseudo-
sphaeriales. The specific distinctions have to a considerable extent been
based on the host species attacked. Until very extensive inoculation ex-
periments can be carried out, this basis for segregation of species will
probably continue to be used. Cyiospora and Ceuthospora (Hyalosporae)
produce their pycnidial cavities in subepidermal or subcortical stromata.
The spores emerge from the separate or united ostioles in an amber-
colored cirrhus. Many of the species are the imperfect stages of Valsa and
related genera in the Sphaeriales. (Fig. 192A.) Sphaeropsis (Phaeosporae)
corresponds to Macrophoma except that the conidia are dark-colored. S.

ORDER SPHAEROPSIDALES

579

malorum Pk. causes twig cankers on apple and quince and the black rot of
the fruits. Its perfect stage is Physalospora. The imperfect form genus is
broken up by the more recent authors into several form genera. The name
Sphaeropsis having been first applied to the perfect stage of an Asco-
mycete should, according to Petrak and Sydow, be replaced by Haplo-
sporella. Some of the species usually included here they transfer to
Botryodiplodia, including among those so transferred S. malorum. Conio-
thyrium has smaller pycnidia and very much smaller conidia which emerge
from the ostiole in a black cirrhus. C. fuckelii Sacc, the cause of the cane
blight of various species of Rubus, has as its perfect stage Leptosphaeria
coniothyrium (Fckl.) Sacc, in the Sphaeriales. Ascochyta (Hyalodidymae)

Fig. 192. Sphaeropsidales, Family Sphaeropsidaceae. (A) Ceuthospora abietina
Delacr.; section through stroma with several pycnidial cavities with one common
ostiole. (B, C) Septoria aesculi (Lib.) West. (B) Section through pycnidium. (Cj
Spores. (E-F) Phomopsis citri Fawcett. (E) Section through pycnidium. (F) Portion
of wall of pycnidium showing pycnospores and the long, slender, curved stylospores.
(D) Family Leptostromataceae. Leptothyrium acerimim (Kunze) Corda; section
through pycnidium. (A, after Delacroix: Bull. soc. mycol. France, 6(4);181-184. B-C,
after Briosi and Cavara: Fascicle V, No. 120. E-F, after Fawcett: Phytopathology,
2(3) -.109-113. D, after Briosi and Cavara: Fascicle II, No. 40.)

580 FUNGI imperfect:: the imperfect fungi

is essentially a Phoma with two-celled conidia. (Fig. 191F.) A. pisi Lib. is
very destructive to cultivated peas. Its perfect stage is Mycosphaerella
pinodes (B. & Bl.) Stone. Diplodia (Phaeodidymae) is like Macrophoma
with colored two-celled spores. In fact some species of Diplodia have been
described as Macrophoma, Sphaeropsis, and Diplodia, depending upon the
age of the conidia. Such species are placed by some authors in Botryo-
diplodia. All three types of spores may be present in the same pycnidium
and all are viable. The 500 or more described species occur on leaves,
stems, etc., and are often the cause of serious diseases of their host plant.
Septoria (Scolecosporae) is a genus of over 1000 species, practically all
parasites. (Fig. 192B-C.) Like Phoma the pycnidium is immersed in the
host tissue and the short ostiole projects to the surface. The conidia are
long and slender, often considerably longer than the diameter of the
pycnidium, hence are curved within the latter. Two serious diseases of
celery are caused by S. apii Chester, and S. apii-graveolentis Dorogin,
respectively. S. lycopersici Speg. defoliates the older plants of tomato
{Ly coper sicom) and causes enormous losses. The perfect stage of some
species of Septoria belongs to the genus Mycosphaerella. Phaeoseptoria
differs from Septoria in the possession of colored, instead of essentially
hyaline, spores. In the genus Phomopsis two forms of conidia are pro-
duced in the same pycnidium; long slender curved stylospores and short
ellipsoidal pycnospores. (Fig. 192E-F.) The perfect stage of some species
of Phomopsis belongs to the genus Diaporthe in the Sphaeriales.

Family Zythiaceae (Nectrioidaceae). The pycnidia are bright-
colored and waxy, like the perithecia of the Hypocreales. Bender recog-
nized 62 genera. Zythia (Hyalosporae) is essentially a Phoma with bright-
colored pycnidia. Some species are parasitic. Aschersonia (Hyalosporae)
produces its pycnidia buried in a bright-colored stroma with several
separate or united ostioles. The stroma is produced externally on leaves
or twigs and in some, probably all, cases is parasitic upon insects feeding
upon the supporting plant.

Family Leptostromataceae. The pycnidia have a well-developed
upper half but the basal portion is not well-developed. They resemble in
many respects the fruiting bodies of some of the Hemisphaeriales. Bender
recognizes 88 genera. They are largely leaf-inhabiting saprophytes and
parasites. Leptothyrium pomi (M. & F.) Sacc. (Hyalosporae) produces the
so-called "fly-specks" of apple fruits. These are the minute flattened,
round pycnidia. (Fig. 192D.) Leptostroma (Hyalosporae) produces elon-
gated pycnidia with slit-like ostioles, on leaves, stems, etc. Entomosporium
maculatum Lev. (Hyalophragmiac) forms four-celled conidia, the cells
forming a sort of sciuare, each with a bristle. It causes leaf and fruit spots
of the pear. Its perfect stage is Diplocarpon soraueri (Kleb.) Nannf.
{Fahraea maculata (Lev.) Atk.), Family Mollisiaceae, Order Pezizales.

ORDER MELANCONIALES

581

Family Excipulaceae. The pycnidia open out early to form a more
or less deep, cup or saucer-shaped structure, tough or hard and black,
either arising subepidermally or subcortically and breaking through to
the outside, or in some cases external from the first. Largely saprophytic
on twigs, stems, etc., less often on leaves. It is sometimes difficult to
distinguish certain subcortical species of this family from the following
order (Melanconiales). Bender recognizes 59 genera. Excipula and Discula
(Hyalosporae) and Discella (Hyalodidymae) are among the genera with
the greatest number of species.

Tehon (1940) separated from among the genera usually included in
the Leptostromataceae two families, Pycnothyriaceae and Rhizothyri-
aceae, which he combines in a distinct order Pycnothyriales. These differ
from the Leptostromataceae in bearing their spores on the under side of
the pycnidial cover instead of basally.

Order Melanconiales. This order of 92 genera^ and over 600 North
American species consists of but one family, Melanconiaceae. To a large
extent the species are parasitic, causing the type of plant disease known
as anthracnose. The three form genera, Gloeosporium (Fig. 193A), Col-
letotrichum (Fig. 194A), and Myxosporium (Hyalosporae), contain some
of the most destructive parasites of cultivated plants. They differ by
arbitrary characters. Colletotrichum produces stiff colorless or colored
bristles (setae) around the acervulus while these are lacking in the other
two. Gloeosporium occurs on herbaceous host structures while Myxo-
sporium occurs on woody stems. How artificial these distinctions are may
be seen in the case of the fungus called Colletotrichum gloeosporioides Penz.,
a common parasite of the leaves, young twigs, and fruits of species of
Citrus and many other genera. When inoculated upon the mango (Man-
gifera indica L.) setae are produced on some of the acervuli on the twigs

• r^r

Fig. 193. Melanconiales, Family Melanconiaceae. (A) Gloesporium populi-albae
Desm.; section through acervulus. (B, C) Septogloeum mori (Lev.) Briosi & Cav.
(B) Section through acervulus. (C) Spores. (A, after Briosi and Cavara: Fascicle VI,
No. 147. B-C, ibid., Fascicle I, No. 21.)

2 The figures for this and the following orders and families are taken from H. B.
Bender (thesis, 1931)

582 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

and leaves, but on the fruits the setae are lacking. George L. Fawcett,
then (1906-1908) a colleague of the author, grew this fungus on over 50
different hosts in Miami, Florida. These had been described in literature
under 25 or more species names in the two genera Gloeosporium and Col-
letotrichum. On many of these hosts the perfect stage was produced and
was found to be Glomerella cingulata (St.) Sp. & von S., of the Gnomoni-
aceae. Since it grows upon twigs as well as upon the leaves and fruits the
distinction between Myxosyorium and Gloeosporium breaks down. The
acervuli begin at first as tangled subepidermal masses of hyphae from
which arise numerous closely packed conidiophores which partially raise
the epidermis. From the apex of each conidiophore are developed one or
more conidia, embedded in gum. When moisture is abundant this gum
swells and the epidermis is burst open and the spores are exuded in a
sticky mass. Insects or other objects coming into contact with these
spores distribute them, as does the rain, whose drops striking such a spore
mass are broken up into smaller wind-borne droplets, each carrying its
burden of spores. Gloeosporium ribis (Lib.) Mont. & Desm. is the imperfect
form oi Pseudopeziza ribis Kleb., in Family Mollisiaceae, Order Pezizales.
Some species of Myxosporium are imperfect stages of Diaporthe in the
Sphaeriales. Marssonina panattoniana (Berl.) Magn. (Hyalodidymae)
causes injury resulting in the formation of holes in leaves of lettuce
{Lactuca saliva L.) and is sometimes very destructive in greenhouses.
Septogloeum (Hyalophragmiae) is mostly parasitic on leaves. It resembles
Gloeosporium but the somewhat elongated spores are several times trans-
versely septate. (Fig. 193B-C.) Coryneum (Phaeophragmiae) has numer-
ous species. C. beijerinckii Oud. occurs in Europe and various parts of the
United States as the cause of a serious disease of the peach (Amygdalus
persica L.). It kills twigs or may attack the buds and merely kill these and
the surrounding tissues, or may cause injury to the fruit. The four- or
more-celled spores are dark-colored and long stalked. Pestalotia (Pes-
talozzia) also has quite similar spores but the apical cell bears one to
three, rarely more, bristles. The terminal cells are lighter colored than the
intervening ones. (Fig. 194B.) These are saprophytes on many hosts but
some species have been supposed to be harmful parasites. The perfect
stages of these two genera are unknown. Cylindrosporiimi (Scolecosporae)
causes leaf spots and leaf fall on many plants. The acervuli resemble those
of Gloeosporium but the spores are long and slender. Perfect stages have
been demonstrated for a few species. Those attacking the genus Prunus
have as their perfect stage apothecial fungi belonging, according to
Higgins (1914), to the genus Coccomyces, of the Family Phacidiaceae; but
according to Nannfeldt (1932) they do not belong to that genus and
family but to the Mollisiaceae, and a genus to which he gave the name
Higginsia. This name, however, is preoccupied and must be replaced if
the genus is held distinct from Coccomyces.

ORDER MONILIALES (hYPHOMYCETEAe)

583

Sphaceloma (Hyalosporae) was long confused with Gloeosporium. Like
the latter its conidia are ellipsoid and produced on short conidiophores.
The acervulus differs considerably in forming a rather firm fungus cush-
ion, in some cases almost intermediate between the sporodochium of
Family Tuberculariaceae and a typical acervulus. On leaves it may cause
considerable malformation. It also attacks herbaceous stems, fruits, etc.

Fig. 194. Melanconiales, Family Melanconi-
aceae. (A) Colletotrichum malvarum (A. Br. &
Casp.) Southw. (B) Pestalotia versicolor Speg.
(A, after Southworth: J. Mycology, 6(2):45-50.
B, after Klebahn: Mycol. Cent, 3(3):97-115.)

The perfect stage of many species has been demonstrated to belong to
the genus Elsinoe of Order Myriangiales. Many species are destructive to
economic plants.

Order Moniliales (Hyphomyceteae). This order contains 651 genera
and toward 10,000 species divided into four families. The first two are
distinguished by the color of the mycelium and conidiophores. The older
distinction placed in the first family those genera with light-colored or
bright-colored mycelium and conidiophores and conidia. Forms with dark-
colored mycelium and conidiophores were placed in the second family,
whether the conidia were light or dark, as well as forms with light-
colored conidiophores but colored conidia. The author follows Bender in
making the distinction as follows:

Moniliaceae: mycelium and conidiophores hyaline or bright-colored (not brown,
smoky, or black), conidia hyaline or colored. These are formed on the ends
of short conidiophores not distinguishable from the other branches of the

584 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

mycelium or are terminal or lateral on distinct, unbranclied or branched,
separate conidiophores. In a few cases the mycelium breaks up into more
or less rounded conidia. Some species of Aspergillus with ochre-colored to
black conidia have conidiophores that are distinctly brown in their upper
portion. Because in their structure they correspond to the species of this
form genus that have colorless conidiophores they are not transferred to the
following family. Members of this family are saprophytes or parasites, includ-
ing some of the most ubiquitous molds and some very serious enemies of
economic plants. Bender recognizes 204 genera, and over 500 North American
species.

Dematiaceae: like the foregoing except that the mycelium and conidiophores
are dark. The conidia may be dark- or light-colored. There are 206 genera,
and over 1000 North American species.

Stilbellaceae^: in this family of 89 genera, and about 100 North American
species the mycelium spreads through the substratum in the usual manner
but the rather long conidiophores arise together in a more or less compact
column or synnema. At the top or down the sides the tips of the conidio-
phores spread apart and bear their spores. The degree of union is various
so that the structure varies from a very short column and bushy head to a
tall column with a small head.

Tuberculariaceae: in this family are 152 genera, and over 400 North American
species. The conidiophores arise more or less radially, packed close together
or separate, from the surface of a somewhat rounded mass of hyphae forming
a sort of cushion, or sporodochium. This may be hyphal in structure or
pseudoparenchymatous and may be waxy, gelatinous, or horny. The conidia
are produced terminally or laterally from the unbranched or branched
conidiophores. The sporodochium and conidia may be light-colored or
dark-colored.

Family Moniliaceae. In the Moniliaceae the classification is based
upon the number of cells in the conidia, whether these are borne upon
special conidiophores or are merely modified portions of the vegetative
mycelium, the number of conidia at the apex of the conidiophore and their
arrangement there, whether in a chain or a head of separate spores caught
in a mucilaginous drop, etc. The genus Oospora, as mostly interpreted by
mycologists, consists of a slender, branched or unbranched mycelium
which breaks up into ellipsoidal or spherical, hyaline or light-colored
conidia (often called "oidia")- There is no sharp line of distinction be-
tween the vegetative hyphae and those that are breaking up into conidia.
0. lactis (Fres.) Sacc. forms a thick wrinkled skin on the surface of sour
milk and of other liquids containing considerable organic matter. It is
quite similar to some of the asporogenous yeasts, many of which are the
causes of disease in Man and other animals, and which are discussed in

' Since the type species of the genus Slilhum, formerly assigned to this family, has
been determined to belong to Order Auriculariales (see p. 444) this name is not avail-
able for a genus in the Fungi Imperfecti and accordingly some mycologists have pro-
posed the name Stilbella for the imperfect forms, wherefore the family name becomes
Stilbellaceae instead of Stilbaceae.

ORDER MONILIALES (hYPHOMYCETEAE) 585

Chapter 11 under the Saccharomycetales. Many of the asporogenous
yeasts are truly Fungi Imperfecti, in that the ascogenous stage is un-
known. The plant-inhabiting species of Oospora such as 0. nicotianae
Splend. & Sacc. are probably not at all closely related to 0. lactis, but
since we are dealing with form genera the matter is not so important
unless good characters can be discovered that will permit the rather un-
wieldy genus to be broken up into several smaller genera.

The genus Actinomyces, in the older, wider sense of the name, repre-
sents beyond doubt a series of several closely related genera whose
relationships within the fungi are not agreed upon by various investi-
gators. The mycelium consists of branching, rarely septate hyphae of
great slenderness, approximately 1 m in thickness or more or less. These
may be aerobic or almost anaerobic in their mode of growth. The hyphae
submerged in the substratum eventually break up, by the formation of
numerous septa, into short, cylindrical spores. In the genus Streptomyces
there is formed a mat of external, branched, aerial conidiophores which
become septate basipetally into conidia which eventually break apart and
are oval or cylindrical or eUipsoidal. These germinate by 1 to 4 germ
tubes. The conidiophores are more often spirally wound. The presence of
true nuclei (Drechsler, 1919 and Newcomer and KenKnight, 1939) and
this mode of germination by germ tubes seem to justify Drechsler and
some other students in considering these genera as true fungi. Until more
is known as to any perfect stage they may well be placed in the Moni-
liaceae near Oospora. Perhaps Drechsler (1935a) is right in drawing atten-
tion to the close similarity of the vegetative structures and asexual mode
of reproduction between Streptomyces and some of the Zoopagaceae in
the Phycomycetes.

Waksman and Henrici (1943) place the genera related to Actinomyces
in two families, which they unite with the Mycobacteriaceae in a separate
order the Actinomycetales, locating it in a position intermediate between
the Bacteria and the Fungi Imperfecti. These two families are the Actino-
mycetaceae, with no aerial conidia, and with two genera Actinomyces,
anaerobic, and Nocardia (syn. Proactinomyces) , aerobic, and Strepto-
mycetaceae, with aerial conidia, in chains in Streptomyces and singly at
the tips of upright conidiophores in Micromonospora. The chief species of
Actinomyces is A. bovis Harz, the organism causing the disease of animals
(especially cattle) known as "lumpy jaw." The genus Streptomyces con-
tains many soil inhabitants feeding upon dead vegetable matter. S. scabies
(Thaxt.) Waks. & Henrici is one of the organisms causing the scab of
potato tubers and other subterranean organs of living plants. For the use
of Streptomyces sp. in the production of antibiotics, see p. 603.

In Cephalosporium (Hyalosporae) the conidiophores produce the
spherical or ellipsoidal conidia successively at the apex, each new conid-

586 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

ium pushing aside the one last formed and all of these adhering by
means of a small amount of slime and forming a head at the apex of the
conidiophore. Miss Pinkerton (1936) has shown that these conidia are
produced endogenously, each one pushing out the previously formed one
into the drop of shme extruded at the tip of the conidiophore. Some
species are parasitic on fruiting bodies of Polyporus. The microspores of
some species of Fusarium are of this type. Haplotrichum is essentially a
Cephalosporium with larger, upright, unbranched conidiophores. Other
genera in which conidia are produced on heads but not in chains are
Oedocephalum, Rhopalomyces, Sigmoideomyces, in all of which the conidia
arise on the swollen tips of the conidiophores. Some of these are probably
in reality Mucorales, but until the sexual reproductive stages are dis-
covered it is unwise to say that of all. B. 0. Dodge (1937) definitely proved
that one species of Oedocephalum is the conidial stage of Peziza pustulata
Pers. {Aleuria umbrina Gill.).

There is a large series of genera in which the conidia are produced in
basigenous chains. In many of these the ultimate, spore-bearing element
of the conidiophore is more or less flask- or bottle-shaped, producing the
spores successively at the neck of the bottle. These ultimate segments are
called phialides, or sterigmata. In Aspergillus the perfect stage is known
for a number of species, and it is probable that many more where this
stage is not now known will be found to possess it. Since it is only rarely
or never found in most of the species the genus is to be sought not only in
the Order Aspergillales of the Ascomyceteae but also in the Fungi Im-
perfecta The conidiophores are upright with a swollen head from which
arise at all sides or only on the upper portion numerous phialides or
sterigmata. At the tips of these are produced successively the conidia
which often form very long chains, the terminal conidium being the oldest.
Although classed in the Family Moniliaceae because of similarity of
structure, the conidia of the various species may be hyaline, yellow, green,
ochre, or even black, and the upper portion of the conidiophore may be
dark. In some species when grown with an abundance of nutrients the
primary sterigmata bear at their apices groups of secondary sterigmata
which produce the conidial chains. Such species have been segregated as
the genus Sterigmatocysiis, but since the distinguishing character is
largely dependent upon the nutrition and other environmental conditions
it does not seem to be of great validity. Closely related to Aspergillus is
the genus Penicillium. Here the conidiophore produces two to several
branches which grow more or less parallel to it and at about the same
level produce at their tips several sterigmata, each bearing a chain of
conidia. The branches may arise in a whorl and these themselves bear
whorled branches or some of the branches may arise singly at the level
below a whorl, thus producing an asymmetrical penicillus or brush of

ORDER MONILIALES (hYPHOMYCETEAE)

587

sterigmata and supporting branches. These substerigmatal branches are
the metulae. As in Aspergillus the conidia vary in color among the differ-
ent species but none are as dark as those of Aspergillus niger van Tiegh.
Both of these genera are ubiquitous molds, mostly saprophytes, although
one or two species of Aspergillus have been described as parasitic in the
human ear. Animals eating moldy hay heavily infested with these molds
sometimes inhale so many spores that fatal mycosis of the lungs ensues.
Penicillium notatum Westling and P. chrysogenum Thom are of great
economic importance because of their production of antibiotic substances
(see closing paragraphs of this chapter). Resembling Penicillium in the
general plan of its conidiophore is the genus Gliocladium which differs in
the secretion of a mucilaginous drop at the top of the penicillus in which
the successively formed conidia are held. The perfect stage of G. penicil-
lioides Corda is Lilliputia, which belongs to the Order Aspergillales.
Matruchot (1895) considered that it belonged to the Perisporiales (Ery-
siphales) and Winter (1873) named it Eurotium insigne and placed it in
the Aspergillales. (Fig. 195A-D.)

Another genus with catenulate conidia borne on phialides is Spicaria.
In this the conidiophore bears definite whorls of phialides and does not

■.^ K^ ygi :'-'■: v- te

ll©_-fc .f

- Conidia-

-SterigmofQ

Branches- ="=

- - Conidiophore

Fig. 195. Moniliales, Family Moniliar-eae. (A) Penicillium frequentans Westling,
illustrating simple type of conidiophore. (B) Penicillium expansum Link, illustrating
branched type of conidiophore. (C) Aspergillus niveo-gluucus Thom and Raper, illus-
trating conidiophore with one series of sterigmata. (D) A. versicolor (Vuill.) Tirab.,
showing conidiophore with two series of sterigmata. (A-B, courtesy, Raper and Thom:
A Manual of the Penicillia, Baltimore, Williams & Wilkins Go. C-D, courtesy, Thom
and Raper: A Manual of the Aspergilli, Baltimore, Wilhams & Wilkins Co.)

588

FUNGI IMPERFECTi: THE IMPERFECT FUNGI

Fig. 196. Moniliales, Family
Moniliaceae. (A) Verticillium
albo-atrum Reinke & Berthold.
(B) Gonatorrhodiella parasitica
Thaxt. (A, after Lindau, in
Engler und Prantl: Die Natlir-
lichen Pflanzenfamilien, Leipzig,
W. Engelmann. B, after Thax-
ter: Botan. Gaz., 16(6):201-215,
Univ. Chicago Press.)

form a brush as in PeniciUium. Closelj^ related to Gliocladium is Clo-
nostachys in which the successive whorls of branches of first and second
rank are more symmetrical and the conidia are in elongated mucilaginous
masses.

Among the Moniliales there are several genera in which the conidia
arise from intercalary enlarged cells in the upright conidiophores. In some

ORDER MONILIALES (hYPHOMYCETEAE) 589

cases well-developed phialides are present but in other cases the conidia
appear to be more properly "radulaspores." They may be single or in
chains, one-celled or septate, hyaline, or brown. Probably the fungi with
this type of conidiophore are not closely related. Among such fungi are
Gonatorrhodiella, Arikrobotrys, Gonatobotryum, Gonatorrhoduni, Gony-
trichum. (Fig. 196B.)

The genus Verticillium is named because of the whorled arrangement
of the branches of the conidiophore. The conidia arise singly at the tips of
phialide-like branchlets. V. alboatrum Reinke & Berth, is the cause of a
disease called "hadromycosis," producing death in very many plants,
woody and herbaceous, e.g., species of Acer, Solanum tuberosum L.,
Dahlia, etc. No perfect stage is known. (Fig. 196A.) Botrytis has branched
conidiophores, but these are not verticillate, nor swollen at the ends.
Various serious diseases of plants are caused by species of this genus.
Some species form sclerotia in the host tissues. Some of the Sclerotiniaceae
(Ascomyceteae) have a Botrytis stage, but for the majority of the species
of Botrytis no perfect stage is known. The genus Phymatotrichum differs
from the foregoing in that the terminal spore-bearing portions of the
branches of the conidiophore are somewhat swollen. P. oinnivorum (Shear)
Duggar is the conidial stage of the fungus causing the root rot of cotton
and many other plants in Texas and some adjacent states. Closely related
is the genus Oidium, more often known as Rhinotrichum. (Fig. 197A.) All
of the foregoing genera are hyalosporous, i.e., have one-celled conidia
which are colorless or light-colored (except as noted for Aspergillus).
Among the Hyalodidymae, Trichothecium, and Cephalothecium both pro-
duce somewhat pear-shaped hyaline or pale pink spores that are two-
celled, the basal cell being smaller than the terminal one. They are mostly
saprophytic although C. roseum Corda enters the fruit of the apple
through the lesions caused by the scab organism {Venturia inaequalis
(Cke.) Winter) and causes rot. In Cephalothecium the conidia are produced
in heads while in Trichothecium they are single at the tip of the conidio-
phore. Arthrobotrys bears similar conidia but they are clustered at various
swollen intercalary cells of the unbranched conidiophore. Some species
are saprophytic, but Drechsler (1937) described six species parasitic upon
terricolous nematodes which are caught in net-like structures of the
mycelium. Mycogone perniciosa Magn. is a dangerous pest in the com-
mercial culture of the common mushroom {Agaricus campestris L. ex Fr.).
Its short conidiophores, lateral to the hyphae of the mycelium, bear
terminally single conidia, two-celled and constricted between the upper
(larger) and lower cells. These conidia may be slightly roughened. They
are sometimes considered to be a form of chlamydospore since on slender
upright hyphae there sometimes occur verticillate branches bearing small
one-celled spores. Other Hyalodidymae growing upon fungi are Diplo-

590

FUNGI IMPERFECTI-, THE IMPERFECT FUNGI

-uTM^r^-^r^^ih^w^^^^

Fig. 197. Moniliales, Family Moniliaceae. (A) Oidiuvi (Bhinotrichum) aureum
Corda; conidiophores and conidia. (B) Piricularia oryzae Briosi & Cavara; conidio-
phores. (C) Ramularia rosea (Fckl.) Sacc; conidiophores with conidia. (A, courtesy,
Linder: Lloydia, 5(3):165-207. B, after Briosi and Cavara: Fascicle VIII, No. 188.
C, ibid.. Fascicle IV, No. 77.)

cladium and Didymocladium, both with creeping vegetative mycehum
and upright branched conidiophores, the former with single or a few not
catenulate conidia at each branch tip, and the latter with more strongly
verticillate branching and chains of conidia at the tips.

Among the Hyalophragmiae there are about a dozen unimportant
saprophytic genera and two that are serious parasites of cultivated plants.
These are Ramularia and Piricularia. In addition some species of Septo-
cylindrium are more or less parasitic. In Ramularia the conidiophores
emerge through the stomata of the host in bundles of two or three to half
a dozen. At first each conidiophore bears a single terminal conidium
which is ellipsoid and one-celled. As it enlarges it becomes elongated,

ORDER MONILIALES (hYPHOMYCETEAe) 591

ellipsoid and three- or more-celled. (Fig, 197C.) In some species at the
distal end of the conidium another conidium develops, and so on until a
short, acrogenously produced chain of conidia is formed. The conidiophore
produces sympodially a lateral projection on which another conidium
arises, and this may be repeated until the upper part of the conidiophore
is a little zigzag or shows several small teeth, each representing the posi-
tion where a conidium was borne. The perfect stage of a few species has
been shown to belong to the genus Mycosphaerella in the Sphaeriales.
Ramularia armoraciae Fckl. is the cause of the very abundant leaf spot of
horse radish {Radicula armoracia (L.) B. L. Rob.). The perfect stage of the
strawberry leaf spot {Mycosphaerella fragariae (Tul.) Lind.) develops only
on the dead overwintered leaves, but the conidial stage, Ramularia tu-
las7iei Sacc, is sometimes very harmful to the growing leaves of the host.
Piricularia causes severe damage to rice {Oryza sativa L.) and produces
leaf spots on many other grasses. Its conidiophores are simple, emerging
from the host's surface and bearing narrowly pyriform, three- to several-
celled hyahne conidia. (Fig. 197B.) In Septocylindrium from short conidio-
phores arise long, sometimes branched, chains of elhpsoidal, three- to
several-celled conidia. Perfect stage unknown.

The genus Dactylella has unbranched conidiophores with the conidia
single at the apex while Dactylaria has the conidia in clusters at the apex.
Many of the species have been described as saprophytic but Drechsler
(1935b, 1937) described some species parasitic on terricolous Amoebae
and nematodes. (Fig. 198C-H.)

The genus Helicomyces is saprophytic. It must be mentioned as one of
the series of probably related genera which have been distributed, because
of the spore color or arrangement of the vegetative hyphae and conidio-
phores among several groups of the four form families of the Moniliales.
In all of these the conidia are two- to many-celled and spirally rolled. In
some the spiral is in one plane, in others it is drawn out somewhat like a
screw. The color of the conidiophores is hyaline in Helicomyces, Hobsonia,
and some others, but in Helicoma and Helicoceras they are dark-colored.
For a fuller understanding of this group the reader should consult Linder
(1929, 1931a and b). (Fig. 199.)

In the Dematiaceae many genera parallel closely those in the pre-
ceding family, differing in the dark color of the mycelium and conidio-
phores. More often the conidia are dark, also. Haplographium (Phaeo-
sporae), except for its dark-colored conidiophores and conidia, closely
resembles Penicillium. Its species are probably all saprophytic. The genus
Coniosporium corresponds closely to Chromosporium in the Moniliaceae.
The round or ellipsoid dark conidia arise on very short stalks from the
scanty, dark, mostly saprophytic mycelium. Corresponding to Oospora in
the Moniliaceae are Torula and Hormiscium in which portions of or the

592

FUNGI IMPEBFECTi: THE IMPERFECT FUNGI

;:^^'

Fig. 198. Moniliales, Family Moniliaceae. Forms adapted to capturing and de-
stroying microscopic animals. (A, B) Arthrobotrys conoides Drechsler. (A) Creeping
hypha and conidiophore. (B) Net in whose meshes nematodes become entangled.
(C, D) Dadylella ellipsospora Grove. (C) Conidiophore. (D) Portion of hypha with

{Continued on facing page.)

OEDER MONILIALES (hYPHOMYCETEAE)

593

Fig. 199. Moniliales, Families Moniliaceae and Dematiaceae. Various helico-
sporous members of these families. (A) Helicomyces scanderis Morgan. (B) Helicoma
perelegans Thaxt. (C) Helicoon auratum (Ellis) Morgan. (D) Hobsonia mirabilis (Peck)
Linder. (Courtesy, Linder: Ann. Missouri Botan. Garden, 16(3):227-388.)

Fig. 198 — (Continued)
sticky knobs on short lateral branches. (E, F) Dactylaria brochophaga Drechsler.
(E) Conidiophore. (F) Nematode-capturing rings, the two upper ones not constricted,
the lower one constricted by contact with a nematode which has escaped. (G, H)
Dactylella tylopaga Drechsler, a fungus parasitic upon Amoeba sp. (G) External
mycelium with conidiophores. (H) An infected animal, with an external hypha bearing
three sticky knobs. (A-F, courtesy, Drechsler: Mycologia, 29(4):447-552. G-H, ibid.,
27(2):216-223.)

594

FUNGI IMPERFECTi: THE IMPERFECT FUNGI

whole mycelium become transformed into chains of individual spherical,
dark-colored conidia. Some of the species form the mildews that are
destructive to cloth, paper, etc. Periconia with its head of dark rounded
or oval spores at the top of a tall conidiophore reminds one of the Moni-
liaceous genera centered around Oedocephalum. Other dark, round-spored
genera are Hadrotrichum and Nigrospora. (Fig. 200A.) In both of these the
conidiophore is of moderate length and dark-colored. In the former the
conidium narrows at the point of attachment. In Nigrospora there is a
sort of vesicle, rather light in color, at the apex of the conidiophore, and
on this lies the very dark-colored spore. Apparently some of the species
are parasitic, mainly upon Monocotyledoneae. N. oryzae (B. & Br.) Fetch
{Basisporium gallarum Moll.) causes injury to the nodes of the stalks and
to the ears and especially to the pointed bases of the grains of Zea mays L.
The genus Zygosporium is peculiar in the manner of bearing the hyaline or
almost hyaline, spherical conidia. These are produced singly at the tips of
two or three hyaline phialides produced near the apex, on the convex
surface of a peculiar dark-colored, curved, and often pointed structure
called by Mason (1941) a falx. This was interpreted by Giesenhagen
(1892) as a basidium and the fungus placed under the name Urohasidium
in the Exobasidiaceae, but Mason's studies show that this interpretation
was erroneous. (Fig. 200B-C.)

Fig. 200. Moniliales. Some peculiar fungi. Family Dematiaceae. (A) Nigrospora
panici Zimmermann {Basisporium of some authors). (B) Zygosporium echiiiosporum
Bunting & Mason, showing the characteristic falces each bearing three phiahdes with
spores. (C) Zygosporium oscheoides Mont, with two phialides on each falx. (D) Lateral
view of a falx showing one phialide with spore. (A, after Zimmermann: Cenir. Bakt.
Parasitenk., Zweite Abt., 8(7):216-221. B-D, courtesy, Mason: Annotated account of
fungi received at the Imperial Mycological Institute, List II, Fascicle 3 (Special
Part), pp. 134-144.)

ORDER MONILIALES (hYPHOMYCETEAE)

595

Fig. 201. Moniliales,
Family Dematiaceae. (A)
Cercospora zeae-maydis Tehon
& Daniels. (B) Cladosporiurn
fulvum Cke. (A, after Tehon
and Daniels: Mycologia,
17(6):240-249. B, after Pril-
lieux and Delacroix, Bull,
soc. rnycol. France, 7(1) :19-
21.)

Polythrincium (Phaeodidymae) produces its dark-colored wavy conid-
lophores in tufts emerging through the epidermis of the host leaf. They
bear apically the colored two-celled conidia. P. trifolii Schm. & Kze, is
parasitic on the leaves of clover {Trifolium). Wolf (1935) has proved that
the perfect stage of this fungus is Cymadothea trifolii (Pers.) Wolf, so that
really this species should no longer be given consideration under the
Fungi Imperfecti but under Order Dothideales in the Ascomyceteae. The
genera Hormodendron and Clados'porium although maintained separately
in the reference books are scarcely worthy of distinction. In both' of them
the conidiophores are colored, septate, and variously branched. Near the
tip the branches are more numerous and bear acrogenously produced
chains, often branching also, of conidia. In Hormodendron these are one-
celled, in Cladosporium the younger spores are one-celled and those
further down in the chain may become two-celled and sometimes three-
to four-celled. (Fig. 20 IB.) Some species of the former represent the stage
in culture on artificial media of certain of the pathogens of Man that
cause a serious disease of the skin known as dermatitis verrucosa (see
Emmons and Carrion, 1937). The genus Cladosporium with over 160
described species is an assemblage of species representing many different
and probably not properly congeneric types. C. herbarum Link ex Fr. is
found the world over on dead organic material and apparently occurs
sometimes as a plant parasite. It probably does not represent a single
species but a host of closely similar and very variable species. The perfect
stage of one form has been described as Mycosphaerella tulasnei Jancz.,
parasitic on various cereals. Another very dissimilar parasitic species is
C. cucumerinum Ell. & Arth., with rather short, unbranched conidio-

596 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

phores, causing spots on leaves and decayed spots on the fruits of the
cucumber, Cucumis sativus L. Upon the leaves of the tomato one often
finds brown- or violet-colored, velvety patches sometimes quite large.
These are the conidia and conidiophores of C. fulvum Cke. In this species
the conidiophores are but little branched. In both the latter species the
conidia rarely if at all appear in chains and are always once septate.

Among the phragmosporous Dematiaceae the genus Helmintho-
sporium includes several species that are harmful to cereal crops. The
upright, unbranched or branched, septate conidiophores arise usually
several from a stoma and bear terminally, and less often laterally as well,
the more or less cylindrical or obclavate, multiseptate, brown conidia.
The infection may be systemic, the mycehum pervading almost the whole
plant, or the infected areas may be limited, forming rounded or elongated
leaf spots. For most of the species no perfect stage is known. The perfect
stage of H. teres Sacc, the cause of the net blotch of barley, is Pyrenophora
teres (Died.) Drechs., one of the Pseudosphaeriales. (Fig. 96 A.) The genus
Heterosporium has shorter conidiophores and ellipsoid, roughened spores,
usually single at the apex of the conidiophore but sometimes in short
chains. The perfect stage of H. gracile (Wallr.) Sacc, cause of leaf spots
and killing of leaves of Iris, is Didymellma iridis (Desm.) v. Hohn., also
of the same order as the foregoing. The perfect stages of the remaining 35
or 40 species of Heterosporium are unknown. There are many other genera
of the phragmosporous Dematiaceae, e.g., Dendryphium, with dendroi-
dally branched conidiophores and the conidia single or in short chains,
Napicladium, with short, weak conidiophores and large terminal conidia,
some species parasitic and others not.

The dictyosporous genera of this family are largely saprophytes. Their
conidia are septate transversely and to a certain degree longitudinally
as well, and occur singly or in chains on short or long, unbranched or
branched, stiff or weak conidiophores. In Alternaria the conidiophore is
rarely branched and the conidia are produced in acrogenously developing
chains or singly. The apical portion of each conidium is narrowed and
often much elongated, bearing at its tip the next narrowly or broadly
ovoid, tapering conidium. In the older literature a distinction was made
between Alternaria and M acrosporium with conidia single at the apex of
the conidiophore. It was shown, however, by Wiltshire (1933, 1938),
Groves and Skolko (1944a and b) and by Neergaard (1945) that the
latter name was first used for species with catenulate spores and only
later for those with spores i)roduced singly. Therefore they proposed that
the name be dropped and Stemphylium be used for some species called in
literature by the name Macrosporium. Those forms with spores tapering
at the upper end should be called Alternaria, even when they may usually
fail to produce chains. Stemphylium has its spores rounded at both ends

ORDER MONILIALES (hYPHOMYCETEAE) 597

-X f"^^~SZ: Fig. 202. Moniliales,

Family Dematiaceae. (A)

Stemphylium sarcinaeforme

(Cav.) Wilts. (B) Alternaria

e tenuis Auct. (sensu Wilt-

^ ^ shire 1933). (A, courtesy,

Groves and Skolko: Can. J.
Research, 22(4):190-199. B,
ibid., 22(5) -.217-234.)

■\ '}

and in some species has a constriction at the median transverse septum.
The conidiophore may be shghtly swollen at its apex. The perfect stage of
S. botryosum Wallr. is Pleospora herharuni (Pers.) Rabenh., a fungus
belonging to the Sphaeriales or Pseudosphaeriales and widespread the
world over. It occurs, apparently, in many strains on all sorts of hosts,
probably developing mostly only in the saprophytic stage of growth. The
perfect stages of the other species of Stemphylium are not known. S.
sarcinaeforme (Cav.) Wilts, is a frequent cause of sometimes serious leaf
injury of various species of Trifolium. Its spores are smooth whereas those
of S. botryosum are roughened. The genus Sporodesmium, with often
roughened conidia occurring singly on rather short conidiophores is of
interest because it sometimes is the imperfect stage of species of Hy-
steriales. (Fig. 202.)

Two genera of great economic importance with long, slender spores
(Scolecosporae) are Cercospora and Cercosporella. These differ only in the
presence of a brown pigment in the conidiophores and sometimes the
spores of the former and its absence in the latter. Since, however, this
pigmentation varies in degree with the age of the fungus and external
conditions there is no real basis for their separation except convenience.
Cercospora is placed in the form family Dematiaceae and Cercosporella in
the Moniliaceae. The conidiophores emerge through the stomatal open-
ings in tufts of varying number, from a pseudoparenchymatous mass of
cells that lies beneath the stoma or sometimes projects through it a little.

598 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

They are stiff, and usually septate and bear at the apex a long, obclavate,
sometimes slenderly tapering conidium, which is usually multiseptate. As
in Ramularia as soon as the conidium is formed another forms closely
below it on a short branch which pushes up beyond the first scar of co-
nidial attachment, so that the apical portion of an old conidiophore is
crooked and marked by numerous scars. The over 500 species are with
few exceptions parasitic in the green tissues of the host plants on which
they produce characteristic leaf spots. Cercospora heticola Sacc. causes a
very destructive leaf spot disease of beets {Beta vulgaris L.), especially the
sugar beet. A leaf spot of cherry is caused by C. ceraseUa Sacc. which has
been shown to be the conidial stage of Mycosphaerella ceraseUa Aderh. C.
apii Fr. is the cause of the early blight of celery {Apium graveolens L.). Of
the many hundred described species of Cercospora and Cercosporella the
perfect stage is known for only a few. (Fig. 20 lA.)

Family Stilbellaceae. In many of the Moniliaceae, especially in the
genus Penicillium, under certain conditions of growth the conidiophores
will be massed together into columns, coremia, from whose upper portion
the conidiophores spread out and produce their conidia. What is here a
response to special conditions is the normal condition in the Stilbellaceae.
The coremium or synnema may be relatively short, the upper half or more
being covered by the spreading tips of the conidiophores, or these may
appear only at the top, the closely united hyphae producing no spore-
bearing branches except at their upper ends. In Isaria the upright, simple
or branched, colorless coremia are covered from near the base to the apex
by slender hyphae bearing terminally the single, small, spherical or ellip-
soidal, hyaline, one-celled conidia. Many of the species grow saprophyti-
cally on plant tissues but a few grow on insects, probably in many cases
parasitically. These latter may be the conidial stages of species of Cordy-
ceps of the Hypocreales. In Graphium the coremium is dark-colored and
the spore-bearing head is only at its top. The conidia are hyaline or almost
so, ovoid or ellipsoid, not in chains, the whole head l^eing enclosed in a
drop of slime. (Fig. 203C.) Such fungi are mostly insect distributed. Some
species are the conidial stages of Ophiostoma (Ceratostomella) of the Asco-
myceteae. Most destructive is G. ulmi Buis. whose perfect stage is 0. ulmi
(Buis.) Nannf., the cause of the so-called Dutch elm disease, so destruc-
tive to American elm (Ulmus americana L.) in America. For the majority
of species the perfect stage is unknown. In the genus Stysanus the co-
remial stalk is colored and as in Graphium the conidia are light-colored or
hyaline. They are borne in chains, covering the upper half or so of the
fruiting body. Most of them are saprophytic but some are suspected of
being weak plant parasites. Perfect stages are unknown. (Fig. 203B.)

Family Tuberculariaceae. The genera customarily assigned to this
family are almost certainly not closely related. The conidia are borne on

ORDER MONILIALES (hYPHOMYCETEAE)

599

Fig. 203. Moniliales. (A)
Family Tuberculariaceae. Epi-
coccum nigruyn Link. (B, C)
Family Stilbellaceae. (B) Sty-
sanus stemonites (Pers.) Corda.
(C) Graphium rigidimi (Pers.)
Sacc. (A, after Lindau, in
Engler and Prantl: Die Natiir-
lichen Pflanzenfamilien, Leipzig,
W. Engelmann. B, after Hassel-
bring: Bota7i. Gaz., 29(5) :312-
322, Univ. Chicago Press. C,
after Hedgcock : Missouri Botan.
Garden Ann. Rep., 17:59-114.)

short or rarely long conidiophores arising from a cushion of fungal tissue
(sporodochium). This varies in color and consistency and the conidia and
conidiophores also show parallelism to the Moniliaceae and Dematiaceae.
The artificiality of the group is seen in the fact that the production or
nonproduction of the sporodochium often depends upon the cultural con-
ditions. Thus the author has grown Colleiotrichum gloeosporioides Penz. so
that it produced its normal acervuli, but in older drier cultures true
sporodochia appeared. Later the latter enlarged and became thrown into

600 FUNGI IMPERFECT!: THE IMPERFECT FUNGI

folds and eventually became stromatic structures containing pycnidial
cavities. Montemartini (1899) reported similar observations in several
groups of Imperfect Fungi. Tuhercularia (Hyalosporae) forms rounded,
bright-colored cushions, mostly on wood or bark. They are covered by
fine, branching conidiophores bearing singly at the tips of the branches
the small, ellipsoidal hyaline conidia. T. vulgaris Tode is the imperfect
stage of Nectria (Creonedria) cinnabarina (Tode) Fr. Volutella (Hyalo-
sporae) produces its small, almost spherical sporodochia on the leaves or
stems of herbaceous plants. Each sporodochium is surrounded by a circle
of long bristles. The conidiophores are mostly unbranched. Several species
cause diseases of plants. V. dianthi (Hals.) Atk. causes cankers near the
base of the stem of carnation {Dianthus caryophyllinus L.) which ulti-
mately kill the plant.

Fusarium (Hyalophragmiae) produces its usually lunate conidia on
rather broad, indefinitely bordered sporodochia as well as singly on the
mycelium. The conidia are usually produced in a mass of slime and in mass
may be white, yellow, orange, or red in color. The many hundred forms
are distinguishable with difficulty, requiring to be cultured on a variety of
culture media, under standard conditions of environment. In addition to
this type of conidia (macrospores) there may be produced microspqres
which are rounded or short ellipsoid conidia, in heads (Cephalosporium
type), as w^ell as spores intermediate in character. Chlamydospores are
often produced in abundance in the mycelium. Sometimes one or two are
formed in a macrospore. Many species of Fusarium produce pigments
which are of assistance in distinguishing the species. Among the species of
this genus are many that cause wilt diseases. The nomenclature of the
different wilt-producers in this genus is very much in dispute. The fact
that on the one hand forms culturally and morphologically indistinguish-
able may be limited to distinct, unrelated hosts, and that on the other
hand wilt may be caused in one host by forms culturally and morphologi-
cally quite different makes the assignment of names to these fungi difficult.
All wilt-producing species of Fusaria are capable of growing saprophyti-
cally in the soil for many years, and from the same lot of soil may usually
be isolated several forms that are distinguishable in structure, habits in
culture, and pathogenicity. In some forms sporodochia appear only very
rarely if at all in culture. It is only a very artificial classification to place
the genus in the Tuberculariaceae. Wilt diseases are produced by various
strains oi Fusarium in tomato {Lycopersicon esculentum Mill.), flax {Linum
usitatissimum L.), cotton (various species of Gossypium), watermelon
(Citrullus vulgaris Schrad.), cowpea {Vigna sinensis Endl.), potato
(Solanum tuberosum L.), celery {Apium graveolens L.), and many other
important crops. Fusarium-Yike fungi are known as the imperfect stages
of Gibberella zeae (Schw.) Fetch, and of various other Hypocreales.

Among the dark-spored members of this form family is the genus

ORDER MYCELIA STERILIA 601

Epicoccum, which forms httle black dots on leaves and stems and other
plant parts. These consist of short, stout conidiophores radiating from
small black sporodochia. The dark conidia are spherical, and slightly
spiny or reticulately marked. In the latter case the spore is apparently
many-celled (Goidanich, 1938), the reticulate lines marking the edges of
the walls of the separate cells. E. oryzae Ito & Iwadara is reported by
Iwadara (1934) to be the cause of injury to rice grains, producing pink or
red lesions. Some of the other species are also parasitic but some seem to
be saprophytes. (Fig. 203A.) Exosporium (Phaeophragmiae) forms similar
sporodochia and very short conidiophores from which arise the obclavate
or ellipsoidal conidia each with several transverse septa. Some species ap-
pear to be parasitic and some merely saprophytic. Spegazzinia (Phaeo-
dictyae or perhaps more properly Staurosporae) occurs on leaves, stems,
and other plant debris and most species are probably saprophytic. The
author (1907) showed that two kinds of conidia are produced on the small
black sporodochia. The commoner kind is very long-stalked, the conidium
consisting of four somewhat rounded spiny cells united at a common
central point to which the conidiophore is attached. The second kind has
very short conidiophores and the conidia are square, except for rounded
corners, being divided into four cells by diagonal septa. They are smooth.
The conidiophore is attached to the edge of one of the triangular cells
making up the conidium and arises directly from the sporodochium and
not, as described by some authors, from the long-stalked conidia. The
spiny conidia of the long-stalked type germinate by bladder-like out-
growths from which radiate many germ tubes. The smaller type of
conidium gives rise to a single germ tube from each cell of the conidium.

Another fungus sometimes ascribed to this family and sometimes to
the Melanconiaceae is Hainesia lythri (Desm.) von Hohn. This is para-
sitic upon strawberry leaves, fruits, and roots (Fragaria sp.) and many
other plants. It has a perfect stage, Pezizella oenotherae (C. & E.) Sacc.
This is rarely observed. Much more abundant are the two fruiting stages:
Hainesia, usually assigned to this family, and Sderotiopsis concava (Dum.)
Shear & Dodge, belonging to Family Sphaeropsidaceae of the Imperfect
Fungi. According to Shear and Dodge (1921) the former has been de-
scribed under seven generic names in four families while the pycnidial
stage has been described under four genera in two families. Part of this
multiple naming has been due to incorrect identification but in part this
is the result of the different appearances of the fungus in various media
and stages of development.

Order Mycelia Sterilia. This order is made up of 20 genera and 400
or more species. This exceedingly heterogeneous group does not at all
consist of closely related species. Any nonsporiferous mycelial structure,
whether sclerotium, rhizomorph, dense or loose mass of mycelium, etc. is
placed here. The presence of clamp connections in some species shows

602 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

that those forms belong to Class Basidiomyceteae, their absence in very-
many cases may or may not indicate that they belong elsewhere. The form
genus Sclerotium includes species producing sclerotia with more or less
definite form, usually light-colored internally and Avith a brown or black
rind. For many forms only the sclerotia and associated mycelium are
known, but many similar sclerotia produce apothecia (Sclerotinia in the
Pezizales) or basidiomycetous spore fruits {Typhula in Family Clavari-
aceae). Pachyma is an enormous subterranean sclerotium, more or less
fibrous-fleshy internally and rough or irregular externally. P. cocos Fr., the
tuckahoe, sometimes larger than a man's head, was formerly used for food
by the Indians inhabiting the regions now comprised by Virginia and
adjacent states. Its perfect stage is a species oiPolyporus which grows out
of the sclerotium. Rhizoctonia consists of superficial, irregular, scale-like,
black sclerotia growing on the surface of the subterranean portions of the
host plant, these being preceded and accompanied by a superficial dark-
colored, short-celled, abundantly branching, rather stout mycelium. This
mycelium is entirely different in appearance from the slender, hyaline
mycelium growing within the host tissues. The perfect stage of Rhizoc-
tonia solani Kiihn is Pellicularia filamentosa (Pat.) Rogers, more com-
monly referred to as Corticium vagum var. solani Burt ex Rolfs, or C.
solani (Prill. & Del.) Bourd. & Galz., Family Thelephoraceae. It is a very
destructive enemy of many species of cultivated plants as well as of a
large number of wild plants. Rhizomorpha is the name given to strands of
mycelium, dark externally and usually white internally. R. suhcorticalis
Pers. consists of the rhizomorphs of Armillariella mellea (Dahl) Karst.,
Family Agaricaceae. Ozonium consists of loose masses of usually bright-
colored mycelium, frequently united into strands and again spreading
out. Some species are parasitic, others saprophytic. 0. omnivorum Shear,
destructive to cotton and many other cultivated as well as wild plants in
Texas and adjacent states, spreads through the soil and attacks the roots
of the host plants. It sometimes produces a conidial stage, Phymato-
trichum omnivorum (Shear) Duggar, mentioned earlier in this chapter.
Once a species of Hydnum was found associated with it (Shear, 1925), but
its connection with this fungus has not been demonstrated beyond
question.

Penicillin, Streptomycin, and Other Antibiotics

Bacteriologists and students of fungi have observed for years that
frequently in cultures of organisms certain contaminating species of
Penicillium, Aspergillus, and other molds would destroy the adjacent
portions of the colonies of bacteria or fungi. Dr. Alexander Fleming had
such an experience in 1928, but instead of discarding his culture of Staph-
ylococcus as spoiled and useless, he began a study of the contaminant and
of the antibiotic substance that it produced. It proved to be a species of

KEY TO THE ORDERS AND FAMILIES OF FUNGI IMPERFECTI 603

Penicillium, later determined to be P. riotatum Westl., and in 1929 he gave
the name penicilHn to the active substance. His experiments demonstrated
that it would destroy many species of bacteria in cultures to which it was
added, while many other kinds of bacteria were unharmed. Dr. Florey, of
Oxford, about ten years later, his interest having been aroused by Dr.
Fleming, tested the effect of the injection of penicillin into the blood
stream of laboratory animals which had been inoculated with pathogenic
organisms which had been found by Fleming to be destroyed by penicillin
in culture. The result was marvelous and the animals recovered. Then it
was tried on Man with promising results. The result is the widespread
commercial production of penicillin on a wholesale basis, using improved
strains of P. notatum as well as of P. chrysogenum Thom, in media and
under conditions that produce many times more of the penicillin than in
the original experiment. Aspergillus has been observed to produce anti-
biotic substances but up to the present the products have shown harmful
effects when injected into the blood stream of animals. Probably some
strain of this genus may be discovered eventually that will lack the harm-
ful factor. Noting that some of the forms related to Actinomyces, e.g.,
Streptomyces as delimited by Waksman and Henrici (1943), also seemed
promising in this regard various soil inhabiting species of this group were
studied and from one of them was selected a strain which produced the
substance to which the name streptomycin has been given. This also is of
great value in medicine since it destroys in the body many organisms
which are not affected by penicillin (see Fleming, 1946; Duemling et al.,
1946; Herrell, 1945).

What the role of these antibiotic substances is in the fungi that pro-
duce them is not yet clear. That these are in the nature of waste products
has been suggested. They reach their maximum production in relatively
young (4 to 7 days), actively growing cultures, well supplied with oxygen.
The production is greatly increased by the addition of certain organic
substances which possibly stimulate the growth of the fungi. This is a
matter that must be left for the researches by mycologic physiologists.

Key to the Orders and Families of Fixngi Imperfecti^

Asexual spores regularly produced.

Spores produced in pycnidia; other types of spores may sometimes be present.

Order Sphaeropsidales
Pycnidia complete, with or without an ostiole, sometimes representing
merely pycnidial cavities in a stroma.
Pycnidia (or stroma) dark-colored, usually rather hard.

Family Sphaeropsidaceae
Pycnidia (or stroma) bright-colored, usually fleshy or leathery.

Family Zythiaceae

■* Note that these groups are with few exceptions artificiarand do|not represent
phylogenetic relationships.

G04 FUNGI IMPEKFECTi: THE IMPERFECT FUNGI

Pycnidia with well-developed roof but basal portions poorly differentiated.

Family Leptostromataceae
Pycnidia with well-developed base, soon opening wide to form a cup-like or
saucer-like structure. Excipulaceae

Spores produced in an acervulus; often other spore types present.

Order Melanconiales
Single family. Family Melanconiaceae

Spores produced neither in pycnidia nor in acervuli; but on or within free sporo-
phores free from one another or grouped in various ways.

Order Moniliales
Conidiophores or spore-bearing hyphae more or less separate, not united into
synnemata nor on sporodochia (cushion-Uke structures).
Conidia and conidiophores hyaline or bright-colored, not brown or black.

Family Moniliaceae
Conidiophores, and usually but not always the conidia, brown or black.

Family Dematiaceae
Conidiophores united into synnemata, the conidia arising on the spreading
tips of the conidiophores in a head at the top or all up and down the
sides. Family Stilbellaceae

Conidiophores usually rather short, arising more or less radially from cushion-
like sporodochia. Family Tuberculariaceae
No asexual spore forms known. Order Mycelia Sterilia

Keys to the More Important Genera of Fungi Imperfecti

Key to the More Important Genera of Family Sphaeropsidaceae
(Based largely upon Lindau in Engler and Prantl, 1899-1900)

Conidia one-celled, spherical, ellipsoid, or oval. Amerosporae
Conidia hyaline. Hyalosporae

Stroma lacking.

Pycnidia smooth, without appendages.
Conidia not produced in chains.
Conidia without appendages.

Pycnidia not surrounded by a subiculum; free or sunk in the sub-
stratum.
Pycnidia with regular ostiole or papillate.
Conidiophores simple or but little branched.

Pycnidia at first covered by the epidermis, later becoming
free, at least around the ostiole.
Conidia less than 15 ju long.

Forming definitely margined spots on leaves.

Phyllostida
On any part of the host, but if on leaves not forming
definitely delimited spots.

Phoma

(see also Fhonwpsis)
Conidia over 15 ju long. Macrophoma

Pycnidia free from the first, free on wood or bark.

Aposphaeria
Conidiophores dendroidally or verticillately branched.

Dendrophoma

KEYS TO THE MORE IMPORTANT GENERA OF FUNGI IMPERFECTI 605

Pycnidia without ostiole or with irregular opening.
Conidia oblong, pointed at each end.

Sclerotiopsis
Conidia oblong, ends rounded. Plenodomus
Pycnidia with long beak. Sphaeronetna^

Pycnidia in a radiating subiculum. Asterovia

Conidia with several early disappearing apical appendages.

Neottiospora
Conidia produced in chains. Sirococcus

Pycnidia with appendages or hairs.
Appendages unbranched.

Bristles all over the pycnidium; spores ellipsoid, curved.

Verinicularia
Bristles mostly around the ostiole; spores straight.

Pyrenochaeta
Appendages stellate or stellately branched at the tip.

Staurochaeta
Pycnidia (rarely) single or several in a stroma or frequently represented only

by pycnidial cavities in the stroma.
Pycnidium mostly single, often without definite, distinct stroma. Conidia

of two types, oval on long conidiophores and long
slender stylospores, usually curved like a hook at one
end, on short conidiophores.

Phomopsis
Pycnidial cavities several in a stroma, with separate ostioles.
Conidia large, more or less fusiform, straight.

Fusicoccum
Conidia cylindrical or ovoid, straight. Dothiorella

Conidia small, allantoid. Cytospora

Pycnidial cavities often with a common ostiole, conidia straight.

Ceuthospora
Conidia colored. Phaeosporae

No stroma, rarely with a subiculum.
Pycnidia glabrous, with regular ostiole.

Conidia oval or elongated, strikingly large. Sphaeropsis
Conidia spherical or ellipsoidal, very small.

Coniothyrium
Pycnidia glabrous, with pronounced beak. Naemosphaeria
Pycnidia glabrous, with lobed, irregular mouth.

Harknessia
Pycnidia with external hairs or bristles. Chaetomella

Stroma present; pycnidia superficial or at first in the stroma.

Haplosporella
Conidia two-ceUed. Didymosporae

Conidia hyaline, pale greenish in some species. Hyalodidymae
Pycnidia free, without subiculum or stroma.
Pycnidia glabrous.

Pycnidia without beak.

Pycnidia in definite spots on leaves or stems.

Spores without appendages. Ascochyta

^ Species of Ophiostoma with dissolved asci may be confused with this.

606 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

Spores with apical bristles. Robillardia

Pycnidia not in definite spots.

Spores without appendages; parasites or saprophytes on higher

plants, Diplodina

Spores with slimy or hair-like appendages at each end; parasitic
upon Uredinales. Darluca

Pycnidia with a beak. Rhynchophoma

Pycnidia with hairs or bristles. Vermiculariella

Pycnidia embedded in a radiate subiculum. Actinonema

Conidia colored. Phaeodidymae

Pycnidia free from one another, without stroma.

Pycnidia subepidermal or subcortical, then emerging.
Pycnidia glabrous.

Spores without outer slime layer. Diplodia^

Spores with external slime coat. Macrodiplodia^

Pycnidia with hairs or bristles. Chaetodiplodia^

Pycnidia from the beginning superficial, on wood.

Diplodiella^
Pycnidia crowded, with stroma. Botryodiplodia^

Conidia 3- or more-celled by transverse septa only.

Phragmosporae
Conidia hyaline. Hyalophragmiae

Conidia without appendages.

Pycnidia more or less globose or depressed globose, mostly remaining

covered, except the ostiolar area (practically a Hen-
dersonia with hyaline spores).

Stagonospora
Pycnidia vertically elongated, emerging in groups.

Mastomyces
Conidia with a bristle-like appendage at the apex.

Kellertnania
Conidia colored. Phaeophragmiae

Without stroma (sometimes a stromatic crust in Dilophospora).

Conidia without appendages, the terminal cells of the conidia hyaline in

some species. Hendersonia

Conidia with appendages.

Appendages a single fine thread at each end of the spore; no stroma.

Cryptostictis
Appendages a tuft of several, fine, branched threads at each end;
pycnidia sometimes with a stromatic crust; parasitic
on grass leaves. Dilophospora

Pycnidia embedded in a stroma; conidia not appendaged.

Hendersonula
Conidia several-celled by transverse and some longitudinal septa; colored.

Phaeodictyae
Without stroma; breaking out through the bark. Camarosporium
Without stroma; superficial on wood. Cytosporium

With pulvinate stroma. Dichomera

* These distinctions are very artificial, for under different cultural conditions any
of these may develop into almost any other form. Also when younger they may
represent Diplodina or Sphaeropsis stages.

KEYS TO THE MORE IMPORTANT GENERA OF FUNGI IMPERFECTI 607

Conidia slender, many times as long as thick, more often hyaline or only slightly

colored, with or without cross septa.

Scolecosporae
Without stroma.

Pycnidia tapering upward to a point; carbonaceous.

Conidia filiform, one-celled. Sphaerographium

Conidia several-celled, constricted at each septum.

Cornularia
Pycnidia globose to depressed globose, ostiole at most with a small papilla;

carbonaceous. *

Pycnidia opening by a rather small round ostiole.

Pycnidia more or less sunken in the host tissue, emergent somewhat at
maturity.
Pycnidia glabrous; parasitic in leaves or herbaceous stems.
Conidia hyaline or at most light-colored.

Septoria
Conidia distinctly brown. Phaeoseptoria

Pycnidia glabrous; growing on wood or bark.

Rhabdospora
Pycnidia hairy. Trichoseptoria

Pycnidia superficial; conidia straight, cylindrical, or narrowly fusoid.

Collonema
Pycnidia opening by a wide mouth; not completely developed.

Phleospora
Pycnidia opening by a narrow slit; not completely developed.

Phlyctaena
Pycnidia globose-conical; tough or leathery.

Pycnidia separate. Micula

Pycnidia crowded in heaps. Micropera

With stroma.

Conidia hyaline, filiform, curved, one-celled. Cytosporina
Conidia dark, several-celled. Septosporiella

Key to the Important Genera of Family Zythiaceae

Stroma lacking; pycnidia globose, with small ostiole, conidia ovoid or oblong,

resembling the perithecia of Nectria. Zythia

With a cushion-like stroma containing several pycnidial cavities; superficial,

parasitic in some cases on leaf-sucking insects. Aschersonia

Key to the More Important Genera of Family Leptostromataceae

(Including the Pycnothyriaceae and Rhizothyriaceae, Tehon, 1940)

Conidia arising from the roof of the external pycnidium.

(Order Pycnothyriales, Tehon, 1940)
Pycnidia ("pycnothyria") radial in structure, often connected with an external
mycelium or subiculum.

(Family Pycnothyriaceae, von Hohnel, 1910)^

' Tehon recognizes 15 genera, with probably not over 40 described species, mostly

tropical or subtropical. The following are included in Leptostromataceae by Lindau,

n Engler and Prantl: (Hyalosporae) Eriothyrium, Trichopeltulum; (Phaeosporae)

608 FUNGI IMPEKFECTi: THE IMPERFECT FUNGI

Pycnidia ("pycnothyria") radial in structure, mounted on columellae whose
bases are connected with the internal mycelium.

(Family Rhizothyriaceae, Tehon, 1940)^
Conidia arising from the floor of the dimidiate subcuticular pycnidium.

(Family Leptostromataceae)
Conidia one-celled, hyaline.
Stroma lacking.

Pycnidia shield-shaped, opening by a pore or short slit.

Leptothyrium
Pycnidia irregular in shape. Piggotia

Pycnidia mostly oblong; opening by a long slit,

Leptostroina
Stroma present (probably the conidial stage of Rhytisma).

Melasmia
Conidia phragmosporous, hyaline, with a hair at each end.

Discosia
Conidia hyaline, cruciform, with two large cells and usually two lateral cells,
all but the basal cell with a slender hair (conidial stage of Diplocarpon) .

Entomosporium
Conidia muriform, hyaline. Didyosporium

Conidia filiform, hyaline or nearly so. Leptosiromella

Key to the More Important Genera of Family Excipulaceae^

Conidia globose, ellipsoid, oblong subcylindrical or fusoid, hyaline or only faintly
colored.
Pycnidia without hairs or bristles.

Pycnidia eventually discoid; wall composed, at least in part, of modified host

tissue. Discula

Pycnidia eventually cup-shaped; walls of typical fungus tissue.
Pycnidia long buried, opening by laciniae. Sporonema
Pycnidia erumpent, opening by round pore. Excipula
Pycnidia erumpent, opening by a wide torn margin.

Dothichiza
Pycnidia with setae.

Conidia without appendages. Amerosporium

Conidia with a delicate bristle at each end. Dinemasporium

Conidia once septate; pycnidia opening by laciniae. Discella
Conidia oblong-fusoid, pluriseptate.

Conidia hyaline; not prolonged into a subulate beak, pycnidia without bristles,

Excipulina
Conidia colored; pycnidia with bristles. Excipularia

Conidia hyaline; prolonged into a subulate beak; pycnidia without bristles.

Heteropatella

Asterostromella; (Hyalodidymae) Lepiothyriella; (Phaeodidymae) Diplopeltis. Tehon
recognizes ten other genera.

* Tehon recognizes 6 genera of which Lindau, in Engler and Prantl includes in the
Leptostromataceae the following: (Hyalosporae) Aciinoihecium; (Phaeosporae) Piro-
stoma; (Scolecosporae) Actinothyrium. Besides these Tehon includes Piroslomella ,
Rhizothyrium, and Cylindrolhyrium.

' In some cases an old acervulus of Family Melanconiaceae may be mistaken for
the saucer-shaped pycnidium of the Excipulaceae.

KEYS TO THE MORE IMPORTANT GENERA OF FUNGI IMPERFECTI 609

Key to the More Important Genera of Family Melanconiaceae

Conidia nonseptate, hyaline, globose to ellipsoid to cylindrical, sometimes fusoid
or allantoid.
Acervulus immersed, but early erumpent, bright-colored, somewhat gelatinous;
some species, at least, parasitic upon Uredinales.

Hainesia
Acervulus immersed but remaining covered by the host epidermis for a con-
siderable time; pale or eventually dark-colored; spores surrounded by
slime; mostly parasitic or saprophytic on herbaceous parts of higher
plants, not on Uredinales. Setae wanting. Gloeosporium
Much like Gloeosporium but the very short conidiophores arising from a firm
base, almost stromatic, or even resembling a small sporodochium. Per-
fect stage Elsinoe. Sphaceloma
Similar to Gloeosporium but with pale or colored setae around or in the acervulus,
their presence often depending upon the substratum.

Colletotrichmn^^
Similar to Gloeosporium but growing only on branches of woody plants.
Conidia straight or almost so. Myxosporium

Conidia allantoid. Naemospora

Conidia nonseptate, globose to ovoid, etc., dark-colored.

Conidia more or less globose; mostly saprophytic on twigs of woody plants.

Melanconium
Conidia fusiform, often curved; mostly on herbaceous plants.

Cryptomela
Conidia hyaline, with one transverse septum.

Parasitic on leaves; conidia more or less oblong. Marssonina
Usually on twigs (rarely on leaves), conidia oblong to fusoid.

Septomyxa
(Rhynchosporium, with conidia strongly beaked, and growing mostly on grasses
is sometimes placed in this group.)
Conidia colored, with one transverse septum. Mostly saprophytic on twigs.

Didymosporium
Conidia hyaline, with two or more transverse septa; on leaves or twigs.

Septogloeum,
Conidia colored, with two or more transverse septa.

Conidia not beaked, emerging in a black drop or cirrhus.

Stilhospora
Conidia not beaked, not emerging in a drop or cirrhus.

Basal cell of conidium usually colorless; the elongated conidiophore persist-
ent on the conidium; walls of the cells thin and collapsing slightly on
drying.

Coryneopsis
Basal cell of conidium colored; walls of the cells of conidium usually much
thickened; conidiophores usually shorter than the spore.

Coryneum
Conidia with colorless upper cell forming a sort of beak.

Conidia three to five septate, arcuate, only the two middle cells colored.

Toxosporium

1" Some species may be confused with Vermiadaria or with Volutella.

610 FUNGI IMPEKFECTi: THE IMPERFECT FUNGI

Conidia five to eleven septate, straight or only slightly curved, all the cells
colored except the basal cell and the one- to three-celled beak.

Scolecosporium
Conidia with filiform appendages.
Appendages only from the apical cell.

Appendage single. Monochaetia

Appendages several. Pestalotia^^

Appendages from both apical and basal cells.

Appendages single. ' Hxjaloceras

Appendages two. Diploceras

Conidia colored, with transverse and some longitudinal septa.

Steganosporium
Conidia colorless, filiform or rod-shaped, many times as long as thick.
Parasitic in leaves or fruits. Cylindrosporium

Saprophytic (or sometimes parasitic?) on twigs.

Conidia filiform, narrow, more or less curved; spore mass yellow or reddish.

Libertella
Conidia cylindric or fusoid, broader; spore mass whitish.

Cryptosporium.

Keys to Special Groups of Order Moniliales; Endosporeae^^

Endoconidiophores forming a layer on the surface of a sporodochium ; endo-
conidia globose, colored, embedded in a slimy layer (Tuberculariaceae).

Hymenella
Endoconidiophores not borne on a sporodochium, endoconidia one-celled.

Endoconidia one-celled, hyaline, pushed out in a chain from the spreading
opened apex of the conidiophore; "macroconidia" in chains, colored. Para-
sites in roots. Thielaviopsis

Endoconidia as in the foregoing, but the apex of the conidiophore not flaring:
"macroconidia" single, sessile or short-stalked. Parasites in roots and upper
portions of plants. Chalaropsis

Endoconidia hyaline, produced in chains in the more or less flask-shaped dark-
walled conidiophores which do not flare at the top. Parasites and saprophytes.

Chalara

Endoconidia mostly becoming brown-walled with age; pushed out successively
into the flaring mouth of the conidiophore and gathering there into a slimy
ball. Saprophytic in woody material, or parasitic on leaves, one species
pathogenic to Man, causing chromoblastomycosis. The saprophytic species
are mostly called Cadophora. Phialophora

Endoconidiophores with flaring collars, producing a short chain of colored dry
conidia, and then proliferating so that a chain of conidiophores is formed,
each with its flaring collar. Catenularia

" More often written by the later name Peslalozzia.

1" In this group of probably not all closely related fungi some of the conidia are
produced in tubular conidiophores from whose apical end the spores are pushed out
successively. These special conidiophores are perliaps to be considered as especially
modified phialides. In addition to the endoconidia other usually larger conidia of the
more usual types ("macroconidia") may occur. By the system of classification ordi-
narily used for Order Moniliales these genera would be distributed among three
different families: Moniliaceae, Dematiacoae, Tuberculariaceae, and perhaps, under
certain conditions of growth, Stilbellaceae.

KEYS TO THE MORE IMPORTANT GENERA OF FUNGI IMPERFECTI 611

Endoconidiophores not on a sporodochium; endoconidia colored, cylindrical,
several-celled, pushed out of the not flaring opening.

Sporoschisma
The following genera with endosporous conidia have only one or two species each
and are not well understood: Sporendonema, Malbranchea, Glycophila. The
spores produced on the phialides of Cephalosporium, Gliocladium, and some
other genera have been shown by Miss Pinkerton (1936) to have an endog-
enous origin.

Key to the Helicosporous Genera of Order Moniliales^^

(Based upon Linder, 1929, 1931a, 1931b)

Conidiophores forming a loose arachnoid, cottony or velvety colony or else appar-
ently obsolete; not forming a compact fruiting body (Moniliaceae
and Dematiaceae).
Conidia coiled in three planes to form a cylindrical or barrel-shaped spore body.
Conidia in chains. Helicodendron

Conidia not in chains. Helicoon

Conidia coiled in two planes (sometimes but slightly inclined to coiling) or if in
three planes not as above.
Parasitic on vascular plants; conidiophores obsolete or as swellings of the
vegetative hyphae; spores toruloid, often nearly straight.

Helicoceras (Gyroceras)
Saprophytic or some doubtfully parasitic on other fungi; conidiophores
present, in some not conspicuous, but then the spores not toruloid.
Conidial filaments thick in proportion to their length, not hygroscopic.
Conidia in chains. Helicodendron

Conidia not in chains.

Conidia longitudinally and transversely septate.

Xenosporella
Conidia only transversely septate. Helicoma

Conidial filaments thin in proportion to their length, hygroscopic.

Conidiophores and conidia hyaline, the conidiophores as teeth on, or
short erect branches from, the creeping vegetative mycelium.

Helicomyces
Conidiophores or conidia, or both, fuscous in shade; conidiophores
conspicuous. Helicosporium

Conidiophores aggregated to form a stele upon which the spores are borne acrog-

enously (Stilbellaceae) . Helicostilbe

Conidiophores aggregated to form a flattened pulvinate or irregularly globose
sporodochium (Tuberculariaceae) .
Sporodochia effuse-pulvinate, at first covered by the epidermis of the host, then
erumpent; conidia once coiled, with thick hyaline walls.

Drepanoconis
Sporodochia pulvinate to irregularly globose, dry, horny, or gelatinous; conidia
without a conspicuously thickened wall.
Conidia coiled in three planes to form a conical or oblong-ellipsoidal spore
body. Troposporium

'3 The genera here assembled in one key represent members that are usually dis-
tributed in the form families Moniliaceae, Dematiaceae, Tuberculariaceae, and
Stilbellaceae.

612 FUNGI IMPERFBCTi: THE IMPERFECT FUNGI

Conidia not coiled in three planes, or if so then the filaments irregularly-
twisted and contorted.
Conidia once coiled, one to three septate; fructifications gelatinous.

Delortia
Conidia not one to three septate; or if so, then sporodochia not gelatinous.
Conidial filaments 7 ju or more in width; conidia coiled in three planes,

twisted and contorted. Hohsonia

Conidial filaments less than 7 /x in width; conidia not coiled in three
planes.
Conidiophores slender, even; fructifications horny when dry.

Everhartia
Conidiophores moniliform; fructifications not as above.

Troposporella

Key to the More Important Amerosporous Genera of Family Moniliaceae

(Based chiefly on Lindau, in Engler and Prantl, 1899-1900)

No conspicuous conidiophores, the globose conidia forming a pulverulent layer
on the surface of the scanty mycelium; saprophytes.

Chromosporiwn
Conidia produced in chains by the basipetally progressing segmentation of the
vegetative hyphae or conidiophores.
Mycelium very slender (< 2 n), mostly nonseptate, producing aerial, more
often coiled conidiophores breaking up into chains of cylin-
drical or ellipsoidal conidia. Sometimes still included in the
genus Actinomyces. Soil organisms, parasitic or saprophytic on
plant material. Streptomyces

Mycelium larger than the foregoing, septate.
Conidia ovoid or globose.

Mycelium forming spreading or cushion-like masses on the substratum.
On plant or animal debris, occasionally parasitic.

Oospora
Mycelium extensive in the substratum, producing tufts of external hyphae
which break up into chains of moderately large to large conidia.
Parasitic or saprophytic on plant material. Some species are
the imperfect stage of Sclerotiniaceae.

Monilia
Mycelium mostly external, with haustoria penetrating the epidermal cells
of the host, and forming upright chains of basigenous conidia.
Asexual stage of various Erysiphaceae.

Oidium^*
Conidia fusiform. Fusidium

Conidiophores simple or only slightly branched; conidia not catenulate, grouped
in heads.
Conidiophores mostly stout, swollen at the top into a distinct sphere or disk,
with radiating globose or fusiform conidia. Some, perhaps all,
of the genera falling into'this category really belong to the
Mucorales.

^* Commonly, but erroneously so called (see Linder, 1942).

KEYS TO THE MORE IMPORTANT GENERA OF FUNGI IMPERFECTI G13

Conidiophores straight, unbranched.
Conidiophores ending in a globose head.

Surface of the head not, or only slightly, areolate.

Oedocephalum
Surface of the head plainly hexagonally areolate.

Rhopalomyces
Conidiophores ending in a lobed disk. Kickxella

(sometimes called Coronella)
Conidiophores repeatedly curved, in S-form; conidial heads lateral.

Sigmoideoniyces
Conidiophores slender, not, or only slightly, enlarged at the apex.

Conidia issuing one at a time from the apex of the conidiophore and remain-
ing embedded in a slime drop. Conidiophores short and straight
from the mycelium. Saprophytic on plant remains, and some
species found in skin lesions in Man.

Cephalosporium
Conidiophores branched; mycelium forming cushions on the surface of the
substratum; conidia becoming green. Saprophytic on plant
material. Trichoderma

The genus Coemansiella placed here by Lindau belongs in Kickxella in the
Mucorales.
Conidia in basigenous chains, grouped in heads.

Conidiophores enlarged at the apex, covered by the sterigmata (phialides)
which bear at the apex the chains of conidia: sometimes pri-
mary and secondary sterigmata present. ^^

Aspergilhis
The genus Dispira (including Dimargaris) of the Mucorales might be keyed
here.
Conidiophore not markedly enlarged at the apex, but branching in regular or
irregular whorls of branches which terminate in two or more
sterigmata.
Conidia and upper portion of the "penicillus" embedded in a drop of slime.

Gliocladium

Slime drop none or small; conidia thin-walled, without ring or collar at base.

Sterigmata irregularly produced, partly variously arranged on fertile

branches, partly in verticils; mostly tapering to long slender

points which are commonly curved or bent from the main axis

of the sterigmata; conidial areas never green.

Paecilomyces
Sterigmata in characteristic verticils with the conidium-producing points
or tubes straight; conidial areas commonly some shade of
green, blue-green or gray-green during their growing period;
white, yellow, or reddish forms occasional, but few.

Penicillium}^
Slime drop lacking, conidia rather thick-walled, with a thickened ring or
collar at the truncate base. Scopulariopsis

Conidiophores unbranched, bearing a tuft of conidial chains without sterigmata.

Briarea

1^ The species with two series of sterigmata have been called Sierigmatocystis.
" The genus Citromyces represents only citric acid producing species of this genus
and is not morphologically distinguishable from Penicillium.

614 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

Conidiophores more or less richly branched, rarely simple.

Conidia arising from especially differentiated intercalary cells of conidiophore.
Conidia produced singly.

Fertile cells of the conidiophore as well as the intervening sterile cells

cylindrical. Gonatobotrys

Fertile cells spherical, intervening cells bone-shaped.

N eviatogonimn
Conidia produced in chains on the globose intercalary or terminal fertile cells.

Gonatorrhodiella
Conidia not arising from especially differentiated intercalary cells.

Branching of the conidiophore very various but never purely verticillate.
Conidia typically pleurogenous on the conidiophore, never terminal.
Conidiophore forked two to several times. Haplaria
Conidiophores unbranched; conidia globose or ellipsoid.

Acladium
Conidia pleurogenous and acrogenous.
Conidiophores typically unbranched.

Parasitic within leaf tissues, clusters of conidiophores emerging from
the stomata, bearing at their tips several (usually 6) conidia.

Microstroma
Mycelium creeping, producing numerous upright unbranched conidio-
phores each with a single hyaline or brightly colored conidium.

Acremonium
Conidiophores with short teeth along the enlarged or not enlarged
upper end; conidia single or in chains from these teeth, large in
proportion to the diameter of the conidiophore, globose to
elhpsoid. Mainly saprophytic (including Olpitrichuin and
Physospora). Oidium

(syn., Rhinotr'ichum)
Resembling the foregoing but parasitic on leaves; conidia hyaline,
globose or oval, single, rarely in short chains.

Ovularia
Conidiophores almost always branched.

Mycehum creeping, conidiophores not upright.

Sporotrichum
Conidiophores always upright.

Conidia single at the tips of the branches.

Conidiophores dendroidally branched; conidia single at the tips
of the branches, hyaline or bright-colored, oval or globose.

Mo7iosporiuni
Conidia clustered near the ends of the usually branched conidio-
phores.
Branches of the conidiophores slender, almost uniform. Conidia

forming loose groups at the tips. Botrytis
Mostly thickei' than the foregoing; the conidia on small sterig-
mata; sclerotium formation frequent.

Polyactis^''
The tips of the rather slender branches somewhat enlarged and
bearing the conidia on distinct sterigmata.

Phymatotrichum ^ ^

" Frequently included in Botrytis.

KEYS TO THE MORE IMPORTANT GENERA OF FUNGI IMPERFECTI 615

Conidia borne along the upper or the lower side of lateral, somewhat
curved, several-celled branches. Martensella and Coemansia

(Mucorales)
Branching of the conidiophores verticillate.

Conidia in chains. Spicaria

Conidia not in chains.

Conidiophores verticillately branched, more often in threes and twos;
conidia hyahne or light-colored, globose or oval, soon falling off.
Ultimate branches straight, conidia mostly terminal.

Verticillium
Ultimate branches subulate, zigzag toward the tip, bearing a conidium
at each angle, this portion with the adhering conidia resembling
the rachis of a head of wheat with attached spikelets.

Tritirachium

(Limber, 1940)
Like Verticillium, but the conidia cylindric.

A crocylindrium

Key to the More Important Genera of the Hyalodidymous Moniliaceae

Conidia not produced in chains.
Conidia smooth.

Conidiophores unbranched; conidia oval or pyriform, the upper cell often
larger than the basal cell; hyaline or bright-colored; mostly saprophytes.
Conidium single, at the apex of the conidiophore.

Trichothecium
Conidia forming a head at the apex of the conidiophore.

Cephalotheci'um
Conidia clustered on somewhat enlarged, intercalary swellings on the
conidiophore. Arthrobotrys

Conidiophores always branched; mostly saprophytes.

Conidiophores verticillately branched; growing saprophytically on fungi.

Diplocladium
Branching of conidiophores irregular, conidia single at the branch tips.

Diplosporium
Conidiophores mostly unbranched, parasitic on leaves.
Conidiophores straight. Didymaria

Conidiophores spirally curved. Bostrichonema

Conidia warty, parasitic on fleshy fungi. Upper cell of conidium larger than the
lower one. Mycogone

Conidia in chains.

Conidiophore branches verticillate. On decaying fungi.

Didtjniocladitim

Key to the More Important Genera of the Phragmosporous Moniliaceae

Conidiophores not much developed; conidia borne singly, curved, fusoid; para-
sites. Fusoma
Conidiophores well developed.

Conidiophores unbranched; not parasitic on vascular plants.
Conidia single on conidiophores. Dactylella

Conidia several in a head on the conidiophore. Dactylaria
Conidiophores unbranched, parasitic on vascular plants.

616 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

Conidia cylindric-oval, single or in short chains, growing out of the stomata

of the host plants. Ramularia

Conidia pyriform; single; mostly parasitic on grasses, sometimes saprophytic
and aquatic. Piricularia

Conidiophores simple or shghtly branched, bearing the conidia in chains. Sapro-
phytic or some species parasitic on vascular plants.

Septocylindrium

Key to the Scolecosporous Moniliaceae

Parasitic on vascular plants, mostly producing leaf spots.

Only genus. Cercosporella

Key to the More Important Staurosporous Moniliales

Conidia hyaline, borne successively on definite hyaline phialides.

Conidia when mature consisting of four slender, ultimately septate, widely
diverging arms, attached to the phialide at their point of divergence ;
conidiophore long and slender, with two to eight phialides near the
apex; aquatic. Lemonniera

Conidia consisting of a main curved axis attached at one end to the phialide and
with two, nearly opposite, lateral, nonseptate branches produced at
the high point of the curve; conidiophores long, with one to four
phialides. Aquatic. Alatospora

Conidia consisting of a clavate, main axis which may be one to three septate
or nonseptate, bearing at the broader, upper end three slender, non-
septate arms or only short divergent processes (the spore then resem-
bling a clove) ; conidiophore long, unbranched or branched, bearing at
the apex one to four phialides. Mostly aquatic; the divergent arms
sometimes lacking on conidia produced in the air; sporodochia some-
times produced. Heliscus
Conidia (aleuriospores or radulaspores) not borne on phialides.
Conidia and conidiophores hyaline.

Aquatic saprophytes, conidia produced submersed. ^^

Conidia with a main cylindrical, or more often clavate to pyriform, axis at
or near whose upper end usually three divergent branches arise
successively.
Main axis of conidium narrowly pyriform or broadly clavate, once
septate; on a long slender conidiophore; the three branches from the
upper cell diverging at angles of about 120° and at about right angles
to the main axis of the spore. In cultures pycnidia sometimes formed.

Clavariopsis
Main axis of conidium narrowly clavate, eventually once or twice
septate, giving rise to three unequal, divergent tapering branches with
a knob or thick finger-like process on the upper side, near the base,
of the first two branches produced. Tetracladium

Main axis of the conidium and the three somewhat longer branches
about equal in thickness, nearly cylindrical, one to three septate at
maturity. Articulosjwra

Conidia with a straight or curved, septate, main axis from which arise
laterally, near together or from separate cells, the strongly divergent
septate branches which may, in their turn, bear lateral branches.

i» For details of these aquatic genera, see Ingold (1942, 1943, 1944).

KEYS TO THE MORE IMPORTANT GENERA OF FUNGI IMPERFECTI 617

Main axis of conidium slender, with two lateral branches diverging at
the same level from the convex side of the curved axis, the two
branches and the lower and upper portion of the axis forming a four-
armed spore; a short "separating cell" present at the upper end of
the conidiophore. Tetrachaetum

Main axis of the conidium stout, septate, curved, the usually two eventu-
ally septate branches arising from adjacent cells of the axis or at
opposite ends of the same long cell. Tricladium

Main axis of the conidium septate, curved, bearing from separate cells
on the convex side several branches approximately in the same plane;
these may also produce lateral branches, the whole spore resembling a
character of Chinese writing; conidiophore slender, bearing one to
several conidia on its distal quarter. Varicosporium

Main axis of conidium long and slender, much septate, straight, bearing
from the cells of the lower half one or more septate, lateral branches,
which in similar manner bear shorter lateral branches; main axis of
spore a continuation of the long, slender, septate conidiophore.

Dendrospora

Not all aquatic, producing the conidia in the air; saprophytes or parasites.

Conidiophore slender, nonseptate, . bearing a single conidium with three

cylindrical, septate arms, one of which is attached by its lower end to

the conidiophore. Trinacrvum

Conidiophore slender, nonseptate; conidia four- to five-armed, nonseptate,

all but one or two tapering into slender bristles; parasitic on fungi.

Titaea
Conidiophore long and slender, hyahne, septate, sometimes branched;
the single hyaline conidium inversely pyramidal, the lower part sep-
tate and tapering to the conidiophore, the broad upper portion with
four one- to two-ceUed conical arms. Parasitic on terricolous nema-
todes.

Triposporina
Conidia colored.

Conidia practically sessile, dictyosporous, at the apex with four long, septate,
cylindrical, spreading extensions. Saprophytic on herbaceous stems.

Tetraploa
Conidia sessile, three to several radiate, the arms obclavate and septate,

saprophytic on woody substrata. Ceratosporium

Conidia sessile, horseshoe-shaped, therefore two-armed, multiseptate; para-
sites. Hirudinaria
(The systematic placing of the three foregoing genera is very uncertain.)
Conidiophores elongated, upright, septate; the single terminal conidium with
a short axial cell from whose top radiate horizontally three septate
arms tapering to a blunt apex. Saprophytic on woody or herbaceous
stems or parasitic on leaves. Triposporium
Conidia of two kinds, usually arising from a small dark-colored sporo-
dochium: (1) long-stalked conidiophore bearing at its apex four spiny
cells, the attachment at the center where the cells meet; (2) four-
celled, smooth spores, attached by the edge of one cell to the short
conidiophore. The latter kind of spore and the sporodochium are
apparently lacking in some species. Saprophytic on dead herbaceous
leaves or stems, one species possibly parasitic on leaf fungi.

Spegazzinia

618 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

Key to the More Important Anierosporous Genera of Family Dematiaceae

Conidia sessile or very short-stalked, vegetative mycelium not extensive.
Conidia round, oval, or discoid. Coniosporium

Conidia elongated, almost fusiform. Fusella

Whole mycelium or the more or less elongated branches developing into chains of
dark-colored conidia. ^^
Conidia falling apart easily; spherical or elongate. Torula
Conidia remaining attached and separating with difficulty.

Horniiscium
Clusters of spreading ovoid or limoniform conidia at the tips of the short conidio-

phores. Echinohotrtjum

Conidiophores well developed; vegetative hyphae abundant.

Conidia dark-colored (except some species of Acrotheca and Trichosporium) .
Conidia not in chains.

Conidia terminal in heads.

Conidia curved or unsymmetrical on unbranched conidiophores which
are hyaline but with black rings at each septum ; conidia dark.

Camptoum
Conidia spherical or ovoid, dark.

With sterigmata. ' Stachybotrys

Practically sessile. Periconia

Conidia fusiform, dark to hyaline, practically sessile.

Acrotheca
Conidia lateral, mostly in whorls.
Conidia smooth, not angular.

Conidiophores dark, unbranched or dichotomously branched.

Gonatobotryum
Conidiophores hyahne, with thick dark septa, mostly unbranched.

Arthrinium
Conidia angular, opposite or in whorls. Goniosporium

Conidia terminal on branched or inflated conidiophores.

Conidiophores distinct from the mycelium, simple or forked, straight.

Virgaria
Conidiophores arising on short lateral upright twigs from the creeping
mycelium.
Conidia sessile. Trichosporium

Conidia on sterigmata. Rhinocladium

Conidiophores branched, the branches curved or hooked; conidia crowded
near the ends of the twigs, colored or almost liyaline.

Canipsotrichum
Conidia terminal and sometimes also lateral on unbranched conidiophores

fin the sense of Saccardo). Monotoxpora

Conidiophores short, unbranched, in a crowded clustei' with a single dark
conidium at the apex, narrowed at point of attachment.

Hadrotrichum
Conidio])hores unbranclied or branched, the black conidium sessile on the
vesicle-like enlargement at the apex of the conidiophoric branch.

Nigrospora

19 Forms with bent or somewliat spirally wound rows of spores should be sought in
the Helicosporae under the genus Helicoceras {Gyroceras).

KEYS TO THE MORE IMPORTANT GENERA OF FUNGI IMPERFECTI 619

Conidia formed in chains, mostly colored.

Conidiophore branched at the summit in a manner resembling Penicillium.

Haplographmyn
Conidiophore bearing a few lateral branches with chains of conidia.

Dernatium
Conidiophore dendroidally much branched, the ends of the branches
bearing acrogenously produced chains of conidia, resembling some
species of Cladosporium but conidia not septate.

Hormodendron
Conidiophore nodulose, with clusters of phialides at the nodes.

Gonatorrhodium
Conidia hyahne or almost so; conidiophores dark, or hyaline and then dark
sterile hyphae present.
Conidiophores unbranched, straight, usually hyaline, upright, close together,
among tall usually dark sterile hyphae, sometimes growing from the
foot of such hyphae.
Conidia ellipsoid or allantoid.

Sterile hyphae wavy, unbranched. Sarcopodiwn

Sterile hyphae circinate, imbranched. Helicotrichum,

Sterile hyphae repeatedly forked, with circinately curved branches.

Circiiiotrichum
Conidia fusiform, curved. EUisiella

Conidiophores bearing the conidia on lateral branches or sessile on the sides.
Conidiophores unbranched, conidia sessile near the upper end.

Chloridium
Conidia arising on lateral branches from the middle portion of the conidio-
phore, cylindrical. Chaetopsis
Fertile branches of the conidiophore not confined to the middle portion.
Branches verticillately borne; conidia globose or ovoid.

Verticicladium
Branches irregular, conidia fusiform. Menispora

Conidiophores branched, bearing clusters of numerous sterigmata at the

joints. Gonytrichum

Conidia arising on (two to three) phialides on the convex side of a dark,
curved, pyriform "falx," which may arise directly from a brown
"foot cell" in the hyaline mycelium or on a short or elongated, or
even branched conidiophore (or "falciphore"); conidia hyaline,
spherical. Zygosporium

Conidia arising in heads of slime at tips of whorled branches of the upright,
dark conidiophore. Stachylidium

Key to the More Im,porlant Didymosporous Genera of Family Dematiaceae

Conidiophores very short or scarcely different in appearance from the dark-
colored conidia.
Conidia not in chains; conidiophores very short. Dicoccum,
Conidia in chains; conidiophores short. Bispora

Conidiophores distinctly different from the mycelium, mostl}^ erect.
Conidia not in chains.

Conidia exclusively apical on conidiophores.
Conidiophores not twisted or swollen.

Conidiophores quite long, with several septa.

Passalora

620 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

Conidiophores short, with only one or two septa.

Fusicladium
Conidiophores regularly twisted or swollen. Polythrincium
Conidiophores short, mostly clustered; conidia terminal and lateral, conidio-

phore not branched. Scolecolrichum

Conidiophores longer, branched. Cladosporium

(some species)
Conidia in mostly acrogenously developed chains.
Conidiophores showing no swelling.

Conidia in short, branched clusters, at first nonseptate but mostly some of
the older conidia once septate (or even twice septate).

Cladosporium
Conidial chains clustered nearer the apex of the long upright conidiophore;
conidia once septate. Diplococcimn

Conidiophores branched, .septate, the individual cells swollen above; conidia
in acrogenously produced chains, early becoming once septate.

Cladotrichum

Key to the More Important Phragmosporous Genera of Family Dematiaceae

Conidiophores very short or scarcely different in appearance from the dark-
colored conidia.
Conidia single, not in chains.

Conidia not drawn out into a long tail.

Conidiophores here and there on the creeping mycelium.

Clasterosp orium
Conidiophores standing close together. Stigmina

Conidia drawn out into a long, pale, often curved tail.

Ceratophorum
Conidia produced in chains. Septonema

Conidiophores well developed.

Conidia not in chains, nor in whorls, nor clustered in a head.
Conidia smooth.
Conidiophores firm.

Conidia elongate. Helniinthosporium'^'^

Conidia ovoid. Brachysporium-^

Conidiophores weak. N apidadium'^'^

Conidia rough. Heterosporium

Conidia formed in whorls, laterally, on the conidiophores.

Spon dylocladium
Conidia clustered in a head at the apex of the unbranched conidiophore.

Acrothecium
Conidia in chains at the tips of the much-branched conidiophore.'^^

Dendryphium

Key to the More Important Dictyosporous Genera of Family Dematiaceae

Conidiophores very short or lacking; conidia mostly sitting directly on the
mycelium.

2° These three genera grade into one another and have no sharp distinguishing
characters.

21 In some species the spore chains are short or lacking.

KEYS TO THE MORE IMPORTANT GENERA OF FUNGI IMPERFECTI G21

Conidia not in chains.

Cells of the conidia not arranged in regular vertical rows.

Conidia of irregular shape with irregular divisions; mostly rather short

sessile. Coniothecium

Conidia elongated, sessile or nearly so, on scanty mycelium, rather uniform

in size and in manner of septation. Sporodesmium

Conidia more or less ovoid, muriform, in little epiphyllous dense heaps.

Stigmella
Cells of the conidium arranged in regular vertical rows which may separate
and spread at maturity. Speira

Conidia in chains. Sirodesmium

Conidiophores distinct, mostly erect.

Spores obclavate, or attenuate at the distal end, borne singly or under favorable

circumstances forming acrogenous chains. Alternaria^^
Spores rounded at both ends, often sarciniform in appearance.

Stemphylium'^'^

Key to the Scolecosporous Dematiaceae

Conidiophores colored ; conidia lightly colored to almost hyaline ; never occurring
in chains; parasitic. Cercospora

(closely related to Cercosporella)
Single genus.

Key to the More Important Genera of Family Stilhellaceae^^

Conidia and conidiophores hyaline.

Conidia one-celled, not in chains; synnema with a capitate sporogenous por-
tion, conidia embedded in a slime drop. Stilbella

(Stilbum of many authors)
Conidia one-celled, not in chains; sporogenous portion of synnema cylindrical
or clavate; conidia not embedded in a slime drop.

Isaria
Conidia one-celled; not in chains; synnema covered by numerous lateral
conidial heads; parasitic on insects (probably not properly placed in this
form family). Gibellula

Conidia one-celled, produced in chains on more or less verticillate sterigmata
(phialides) ; in many cases merely cultural forms of Penicillium whose
conidiophores often are united into synnemata.

Coremium
Conidia phragmosporous, straight, not in chains. Arthrosporium
Conidia phragmosporous, slender falcate, not in chains.

Atradiujn
Conidia or conidiophores, or both, dark-colored.
Conidia one-celled.

Conidia not in chains, globose, ellipsoid, or oval.

2^ In mycological and pathological literature the name Alternaria is often reserved
for species occurring in chains and the genus Macrosporium for species never pro-
ducing chains. Sometimes the name Stemphylium is used in a manner synonymous
with Macrosporium.

^' For helicosporous Stilbellaceae, see key on p. 611 for helicosporous Moniliales.
Many Moniliaceae and Dematiaceae under special cultural conditions will produce
coremia, e.g., Aspergillus, Penicillium, etc.

622 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

Conidia hyaline or pale, mostly embedded in slime.

Gra'phium

Conidia dark-colored. _ Sporocybe

Conidia not in chains, more or less falcate, hyaline.

Harpographium
Conidia hyaline, in chains; spore-bearing portion of synnema globose or

elongated, without hairs. Stysanus

Conidia in chains; but spore-bearing portion of synnema with long black
hairs. Trichurus

Conidia phragmosporous.

Spore-bearing portion covering the whole synnema; conidia smoke-colored.

Podosporium
Spore-bearing portion only at the upper end of the synnema.

Synnema of loose hyphae; spore-bearing head loose; conidia hyaline or

pale. Isariopsis

Synnema of stiff, firmly united hyphae; spore-bearing head distinct;
conidia pale or smoky. Arthrobotryum

Key to the More Important Genera of Family Tuber culariaceae

(After Lindau in Engler and Prantl, 1900)

Conidia and conidiophores and sporodochium hyaline or the last bright-colored

(not dark).
Conidia amerosporous.

Sporodochia without hairs or bristles.
Conidia single, not occurring in chains.
Conidia not embedded in slime.

Sporodochia of various shapes but never cup-shaped with definite
margin.
Conidiophores unbranched or more rarely a little branched.
Sporodochia almost spherical, superficial.

Aegerita
Sporodochia cushion-shaped or tuberculate or indefinite in form.
Conidia globose; sporodochia very small, later becoming hard.

Tuberculina
Conidia ovoid; sporodochia disk-like, larger.

Hymenula
Conidiophores always branched, mostly considerably.
Conidia ovoid or elongate.

Conidiophores not verticillately branched.

Tubercularia
Conidiophores verticillately branched.

De7idrodochium
Conidia fusiform to falcate or cylindrical.

Fusicolla
Sporodochia cup-shaped, with definite margin.

Patellina
Conidia embedded in slime; sporodochia disk-shaped, gelatinous- waxy.

lUosporium
Conidia occurring in chains.

Sporodochia more or less globose, sometimes stalked.

Sphaeridium

KEYS TO THE MOEE IMPORTANT GENERA OF FUNGI IMPEFECTI 623

Sporodochia more or less disk-shaped, not stalked.

CylindrocoUa
Sporodochia sessile or short-stalked ; at the margin with bristles.

Volutella^*
Conidia phragmosporous.

Conidia straight, at the upper end somewhat thickened and angular.

Heliscus
Conidia straight, elongate-cylindrical, not thickened at upper end.

Bactridium
Conidia falcate-fusiform.

Sporodochia conical or cushion-like, delicate; often parasitic upon insects.

Microcera
Sporodochia extended, or large and cushion-formed, more often lacking,
the spores then arising from the mycelium.

Fusarium
Conidia brown, rarely almost hyaline; sporodochia and usually the conidiophores

brown.
Conidia one-celled.

Sporodochia without hairs or bristles.

Sporodochia cushion-like or tuberculate or almost spherical, black.

Conidiophores short; conidia globose; parasites, or more often, sapro-
phytes. Epicoccum
Conidiophores longer, mostly branched; conidia ovoid to elongate, some-
times curved. Saprophytes. Strumella
Conidiophores slender or clavate at the apex; conidia ovoid, elongate, or
pyriform, borne singly or in chains.

Epidochium
Sporodochia flat, not convex; conidiophores cylindrical, conidia ovoid,
elongated, or rod-shaped. Hymenopsis

Sporodochia with hairs or bristles.

Hairs black, marginal. Chaetostromn

Hairs hyaline, marginal. Myrothecium

Conidia phragmosporous.

Conidia single on conidiophores. Exosporium

Conidia in chains on conidiophores. Trimmatostroma

Conidia spherical, reticulately marked, each areola representing a distinct cell.

Epicoccum

(some species)

Key to a Few Genera of Order Mycelia Sterilia

Sclerotium-like bodies produced.

Sclerotia flattened, usually on surface of host, connected by brown fibrillose
hyphae. Imperfect forms of Basidiomyceteae. Rhizoctonia

Sclerotia round, ellipsoid or elongated, often more or less flattened on lower
surface, often several growing together into a compound structure; internally
firm and usually white, with a definite, black, brown, or light brown cortical
layer. Many species are imperfect forms of Aseomyceteae, others of Basidio-
myceteae. Sclerotium

^* Parasitic species when young may be confused with CoUetotrichum.

624 FUNGI IMPEEFECTi: THE IMPERFECT FUNGI

Sclerotium subterranean, consisting of a large mass of tangled hypliae enclosing
particles of soil, plant debris, etc., the outer surface not forming a hard
cortical layer; giving rise to spore fruits of Polyporaceae.

Pachyma
Root-like, branching, growing through the soil or between wood and bark or in
timber; hard, white within, with black cortex. Usually representing a stage of
some Basidiomycete, most frequently Armillariella mellea.

Rhizomorpha
Coarsely interwoven hyphae; mostly parasitic on roots of vascular plants.

Ozonium^^

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Some Hyphomycetes that prey on free-living terricolous nematodes, ibid.,

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Fitzgerald, L. R.: Phoma stenobothri, a fungus parasite of the grasshopper.

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25 Besides Ozonium many names have been given to various hyphal forms.

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Botan. Ztg., 76:130-156. PZs. 12-13. 1892.
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626 FUNGI IMPERFECTi: THE IMPERFECT FUNGI

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THE PHYLOGENY OF THE FUNGI

IT IS a commonly accepted axiom that evolution in both plants and
animals generally proceeds from the simpler to the more complex
structures, although it is recognized that in many cases a retrogression
from complex to simpler structures may occur. Therefore it is natural
that from the time that the ideas of evolution began to be given considera-
tion in the attempts to develop taxonomy along supposed phylogenetic
lines search was made for simple organisms as the probable primitive
types from which the complex phylogenetic trees of plants and animals
had developed.

Since animals lack the power of synthesizing organic food stuffs out of
inorganic materials it has usually been assumed that the predecessors of
the animal kingdom must have been organisms that did possess that
power. The same assumption is necessary for the establishment of a
logical phylogenetic arrangement of the fungi and other more or less
plant-like organisms that are not autotrophic. There are a number of
groups of microscopic organisms ordinarily classed among the Bacteria in
which the energy necessary for the synthesis of organic materials is ob-
tained by various types of chemical reaction independent of light. Thus
the sulphur bacteria by the oxidation of sulphur or of HoS are able to gain
the necessary energy for their life processes. The same is true of those
bacteria which oxidize iron. There are several other types of energy-
obtaining processes. It is, however, only in those plants which develop
chlorophyll-like substances able to react with light to ol:)tain the energy
necessary for synthesis that a comparatively large amount of energy
became available and that production of large amounts of food occurred.
Probably not until then did the evolution of larger, more complex organ-
isms begin and not till then was the available food supply great enough to
permit the extended evolution of the various orders of animals and of

fungi.

From the fact that most of the simpler animals, green plants, and
fungi for the major portion of their life cycle consist of single, uninucleate

628

THE PHYLOGENY OP THE FUNGI 629

cells with flagella or cilia or produce such cells for the purpose of multi-
plication it seems logical to conclude that the ancestors of these groups
were unicellular, motile organisms. They must have been already highly
developed in their cell structure for we find in the green algae, lower fungi,
and animals essentially the same basic nuclear structure, sexual repro-
duction whose fundamental phenomena are the union of two haploid
nuclei into a diploid nucleus followed sooner or later by meiosis, and
similar laws of heredity. Therefore we must believe that the forms an-
cestral to these three lines had already reached the height of development
attained by the unicellular, motile or zoospore-producing algae. The
sulphur or iron bacteria mentioned above lack such definite nuclei and are
much too simple in structure to have been the immediate ancestors
sought.

It appears to the author that probably the lines of evolution after the
invention of chlorophyll progressed rapidly in various directions. The
known Myxophyceae have not reached the type of cell structure that
could have given rise immediately to the unicellular, flagellate green algae
or to the lower fungi or animals. It is to the organisms more like the simple
chlorophyll-bearing flagellate algae that we must turn in our search. In
such algae we have a constant production of organic material, by means of
photosynthesis, and thus chlorophyll-lacking organisms could develop,
depending upon them either directly or indirectly for their food. We find
that the motile organs of such simple alga-like organisms as we now know
are usually one to four in number, connected to little organelles, the
blepharoplasts, at or just below the plasma membrane, these usually being
then connected to the nucleus. Evolution led to the production of two
main types of flagella. In one type there is a firmer outer tubular envelope
surrounding a softer more flexible portion which extends beyond this
outer portion and is capable of very active lashing motion. These are the
"whiplash" flagella. Another type consists of a much more flexible axis
from whose sides arise at right angles numerous very fine cilioles which are
capable of movement. This is the "tinsel type" of flageflum. Some of the
single-celled plants possess one or more of the one type or of the other
type, but in one group of algae one flagellum of each type occurs. This is
the group called Heterocontae or Xanthophyceae. Here the flagella may
be equal in size but usually with the tinsel flagellum directed anteriorly
and the other turned back. The latter is frequently shorter and in some
cases appears to be lacking.

Another characteristic of the unicellular algae is the fact that the
motile cells do not normally produce a true cell wall, the outer layer being
simply a plasma membrane. This becomes encysted with a true cell wall
which serves as a protection. In the further evolution of the various
groups of algae the encysted stage becomes the normal vegetative condi-

630 THE PHYLOGENY OF THE FUNGI

tion, the naked, swimming cells being produced for multiplication. We
find this same trend of evolution in the fungi that we speak of as the
Lower Fungi. In the line of evolution of animals the contrary feature is
emphasized. The motile stage becomes more and more the prevailing one,
the encystment occurring only at the reproductive stage and finally dis-
appearing altogether. In some of the simple animals the plasma membrane
has become less firm and amoeboid motion is possible, together with the
amoeboid engulfment of food particles, yet even here encysted stages
may occur.

We may perhaps assume that some of the chlorophyll-containing, one-
celled, naked, swimming organisms, whether we call them plants or not,
and which had possibly not yet reached the level of the present simplest
green algae, lost their power of producing chlorophyll and so had to
become parasitic in the tissues of animals or plants or saprophytic, feeding
upon the organic products of plant-like organisms. Among the group com-
monly called the Flagellata we find numerous examples of this type in
which the lack of chlorophyll is the chief distinguishing character. Also
such examples of lost chlorophyll occur among the true green algae, as for
example in Rhodochytrium, in which the carotinoid pigments are still pro-
duced although the chlorophyll has disappeared. This plant lives para-
sitically in the tissues of higher plants. Intermediate steps are found in
certain unicellular algae that still retain their chlorophyll but which are
obligate endophytes in aquatic higher plants. Examples of these are
Chlorochytrium and Eremosphaera.

Phylogeny of Mycetozoa

The Mycetozoa are characterized by their naked plasmodial vege-
tative stage and by the ultimate rounding up and encystment of the cells
as "spores," the only stage (with minor exceptions, such as the stalk cells
of Acrasiales) in which cell walls are found. With the exception of the few
described Acrasiales and Labyrinthulales, the encysted spores germinate
by production of anteriorly biflagellate swarm cells or sometimes one
flagellum seems to be lacking although two blepharoplasts are present.
These two fiagella are both of the whiplash type. The presence of flagel-
late, amoeboid planospores and inclination toward the formation of
Plasmodia or pseudoplasmodia is found also in some of the Protozoa
included in the Rhizopoda or Sarcodina and therefore it seems to the
author only reasonable to conclude that the Mycetozoa (in the broader
sense of this term) are derived from those organisms. Where they came
from remains to be determined but possibly the amoeboid Protozoa are
further developments of chlorophyll-less Flagellata. The Plasmodiopho-
rales are classified by Sparrow (1943) and some others in the Biflagellatae,
a group which includes also the Saprolegniales, Leptomitales, Lage-

PHTLOGENY OF CHYTRIDIALES AND VEGETATIVELT SIMILAR FORMS G31

nidiales, and Peronosporales. Since, however, Ellison (1945) has shown
that the two flagella oi Plasmodiophora are both of the whiplash type while
those of the other groups assigned to the Biflagellatae have one each of the
tinsel and whiplash type, the relationship of Plasmodiophora appears more
likely to be with the biflagellate slime molds whose flagella are both of the
whiplash type and whose plasmodial structure is more nearly similar.
Whether the other genera at present assigned to the Plasmodiophorales
really belong there needs further study, especially of their flagellar
structure.

The nonflagellate order Acrasiales is possibly closely related to the
true slime molds. Besides the loss of fiagellum production there is no
definite peridium nor is a capillitium produced unless possibly the slime
in which the spores are embedded and which helps to make up the growing
stalk are modifications of these structures. Furthermore the production of
true Plasmodia is lost except possibly just before spore formation, but
sexuality appears to be present (Skupienski, 1920). As for the Labyrinth-
ulales it can only be surmised that they possibly are related to the Myce-
tozoa because of their naked cells, which form a sort of pseudoplas-
modium, and the encystment of their spores. Their life history is so
incompletely known that their relationship cannot be determined more
closely at present.

Phylogeny of Chytridiales and Vegetatively Similar Forms

The whole group of chiefly aquatic fungi formerly grouped together
under the name Chytridiales agrees in the production of flagellate naked
swarm cells which eventually encyst and enlarge and become the spo-
rangia or give rise to the sporangia within which the swarm cells are pro-
duced. The study of the number, location and structure of the flagella has
shown that this group must be divided into at least three orders : Chytri-
diales (in the more limited sense), with one posterior whiplash fiagellum;
Hj^phochytriales, with one anterior tinsel type fiagellum; and certain
families of the Lagenidiales, with two anteriorly or laterally attached
flagella, one of the tinsel type (directed forward if the flagella are laterally
attached) and the other of the whiplash type (directed posteriorly in most
cases). Correlated with these flagellar characters are chemical differences
in the composition of the cell wall. In the Lagenidiales this gives the
cellulose test readily with chloriodide of zinc, in the Hyphochytriales this
reaction may or may not appear and may depend upon the age of the
organism, while in the Chytridiales only rarely is cellulose revealed by
this test and in some cases fungus chitin occurs.

In 1942, the author suggested that these three orders which vege-
tatively and in their mode of reproduction are so closely parallel, might
have had a common origin in some of the green, heterocont, one-celled,

G32

THE PHYLOGENY OF THE FUNGI

motile, anteriorly biflagellate algae (Heterocontae or Xanthophyceae). It
is necessary to make the assumption that the loss of chlorophyll and the
adoption of a saprophytic or parasitic mode of life occurred early in their
evolution. The simpler Lagenidiales represent forms in which both flagella
are retained, as well as the cellulose composition of the cell walls. For the
Hyphochytriales we must assume a gradual modification of the hetero-
contal habit until finally (as sometimes occurs in the algae) the shorter,
posteriorly inclined flagellum entirely disappeared, leaving only the single
anterior tinsel type flagellum. At the same time the cellulose became in
part replaced by or concealed by an admixture of fungus chitin. For the
Chytridiales (in the narrower use of the term) the anterior (tinsel type)
flagellum has disappeared leaving the single, posteriorly directed whiplash
flagellum. Concurrently the cellulose reaction of the cell walls has become
less and less marked, appearing only in a minority of the species so far
studied. (Fig. 204.)

These three groups have undergone parallel evolutionary develop-

ENTOMOPHTHORALES

MUCORALES

PERONOSPORACEAE

MONOBLEPHARIDACEAE

BLASTOCLADIACEAE

SAPROLEGNIACEAE

LEPTOMITACEAE

ALBUGINACEAE

PYTHIACEAE

CLADOCHYTRIACEAE

RHIZIDIACEAE

SYNCHYTRIACEAE

OLPIDIACEAE

ANTERIOR FLAGELLUM LOST

LAGENIDIACEAE

HYPHOGHYTRIACEAE

POSTERIOR FLAGELLUM LOST

OLPIDIOPSIDACEAE

,BOTH FLAGELLA RETAINED

HETEROCONT UNICELLULAR ALGAE

Fio. 204. Suggested lines of evolution of the Pliycomyreteae, based upon the idea of
their origin from unicellular algae. (After Bcssey: M ycologia, 34(4):355-379.)

PHYLOGENY OP CHYTRIDIALES AND VEGETATIVELY SIMILAR FORMS 633

SAPROLEGNIACEAE

ments from holocarpic to eucarpic monocentric forms and finally to
eucarpic polycentric forms. So far as known the Hyphochytriales have
not given rise to further advanced, hyphal forms of a truly mycelial
nature. The Chytridiales, on the contrary, are certainly closely related to
the Blastocladiales and Monoblepharidales, culminating in the latter in
an oosporic mode of sexual reproduction. This line seems to end blindly.
From the Lagenidiales we can draw lines of ascent to the Saprolegniales
and Peronosporales.

PERONOSPORACEAE

ALBUGINACEAE

ENTOMOPHTHORALES

I
PYTHIACEAE MUCORALES

LEPTOMITACEAE

LAGENIDIACEAE

I
OLPIDIOPSIDAGEAE / VAUCHERIACEAE

HIGHER /si PHONALES

Fig. 205. Suggested lines of evolution of
the biflagellate Phycomyceteae, based upon
Sachs's and Mez's idea of the origin of the
Saprolegniales from the Siphonales. (After
Bessey: Mycologia, 34(4):355-379.)

On the contrary Mez (1929), de Bary (1884), and many others have
suggested that the evolution may have been regressive from the Sapro-
legniales or from the Pythiaceae, to the Lagenidiales, by a process of
simplification. From the simple, holocarpic Olpidiopsidaceae, by the loss
of the posterior or anterior flagellum could have arisen the Hyphochy-
triales and Chytridiales respectively. No forms are known from which the
Monoblepharidales and Blastocladiales might have arisen and then by
regression led to the production of the Chytridiales. The Saprolegniales
are assumed by Mez and by Sachs (1874) under this hypothesis to have
arisen by loss of chlorophyll from some alga similar to Vaucheria in the
Siphonales, at a point in the evolution of this alga prior to the substitution
of a single compound zoospore for the many separate biflagellate zoo-

634 THE PHTLOGENY OF THE FUNGI

spores characteristic of most Siphonales and of the Saprolegniales. The
close similarity of the serum reactions by Saprolegnia and Vaucheria as
determined by Mez would seem to add force to this idea. (Fig. 205.)

Phylogeny of Higher Phycomyceteae

The phylogeny of the Mucorales and Entomophthorales is probably
properly tied up with that of the Zoopagales. Of the Eccrinales so much
still remains to be learned that their origin and their relationships to other
groups are very obscure. The fact that their cell walls respond to the
cellulose test positively with chloriodide of zinc is not sufficient to indicate
where their closest kinship lies.

The Mucorales and Entomophthorales have cell walls at maturity in
which cellulose is not readily demonstrated although in some Mucorales
it can be detected in younger mycelium by the use of suitable iodine-
containing reagents. As the mycelium becomes older fungus chitin makes
up more and more of the cell wall. The fact that cellulose is sufficiently
abundant in proportion to the chitin to be demonstrable in the younger
mycelium would seem to justify the suggestion of the possibility of an-
cestors with little or no chitin in their cell walls. Be it remembered that in
Pythium no fungus chitin is demonstrable (Thomas, 1942, 1943) while it
occurs in measurable amounts in the very closely related Phijtophthora.
The more typical Mucorales are usually considered to be those in whose
asexual reproduction the aerial hyphae terminate in sporangia within
which by cleavage are produced angular, naked cells which quickly round
up and become encysted. Upon their escape from the sporangium by the
dissolution or fragmentation of the membrane the aplanospores may be
distributed by water or by air currents or even by insects. Except for the
nonflagellate condition of the spores this type of asexual reproduction is
found in the Blastocladiales, Monoblepharidales, Saprolegniales, and
some of the terrestrial and aquatic Peronosporales. The fact that the
mycehum of the Mucorales is mostly stout and that large numbers of
species are soil inhabitants would seem to exclude many of the Pythiaceae
and Blastocladiales and Monoblepharidales from consideration. In the
Saprolegniaceae we find the genus Aplanes in which the spores produced
in the sporangium become aplanospores, without undergoing the swim-
ming stage. Thus, so far as the asexual mode of reproduction is concerned,
there is no serious barrier to the behef that the Mucorales may have
evolved from some Saprolegniaceous soil fungus with aplanospores. In
general, however, the conjugating gametangia in the Mucorales are almost
equal in size while in the Saprolegniales they consist of a small antherid
and a large oogone, the former usually producing a conjugation tube
which penetrates through the oogone wall and opens at its tip when
nearly or quite in contact with the egg. Yet in Brevilegnia diclina no such

ORIGIN OF ASCOMYCETEAE 635

conjugation tube is formed, there being developed simply an opening
through the antherid and oogone walls which permits the entry of the
male nucleus. However, the supposed isogamy in the Mucorales is more
apparent than real. Even where the gemetangia are equal in size the
nuclei and part of the cytoplasm of one pass through an opening in the
walls into the other so that we actually have a functioning antherid
although it is equal in size with the oogone. In Dicranophora one game-
tangium is very large and the other very small and the "zygospore" wall
includes the oogone alone or rarely part of the antherid also. So we can
perhaps with justification look to some soil-inhabiting member of the
Saprolegniales for the ancestral form of the Mucorales. It must be noted
that Jaczewski (1929-30) suggested that these latter fungi arose directly
from some of the Chy tridiales (in the wider sense) . Others have suggested
the Monoblepharidaceae or the Cladochytriaceae as the ancestral stock
of the Mucorales.

Within the Mucorales the evolution has apparently been in various
directions of modification of the sporangium, terminating in small in-
dehiscent sporangioles or portions of elongated sporangia. The Ento-
mophthorales appear to be terminal lines of evolution in which the
mycelium is much reduced, and the "conidia" represent usually violently
discharged, mostly indehiscent, sporangioles (but in Basidiobolus these
produce internal aplanospores). The sexual reproduction of the various
genera represents modifications of some of the different types found in
the Mucorales.

In the Zoopagales the sporangia are reduced to indehiscent sporan-
gioles (or "conidia") and the sexual reproduction is sometimes isogamous
and reminiscent of some of the Mucorales or more often heterogamous.
Not enough is known of their cytology, the chemical composition of the
cell walls etc., to permit more definite suggestions as to their phylogeny.

Origin of Ascomyceteae

There are two main schools of thought regarding the phylogenetic
origin of the Higher Fungi (Ascomyceteae, Basidiomyceteae, etc.). In the
one it is held that the fungi as a whole form a monophyletic series and,
consequently, it is believed that the Ascomyceteae and other higher fungi
arose from the Phycomyceteae. There is no general agreement as to the
definite paths along which such derivation occurred, nor whether the
higher fungi are monophyletic or polyphyletic in their origin from the
lower fungi. The other school holds that the fungi are not necessarily
monophyletic and that some, if not all, of the higher fungi arose from
algae that had some of the characteristics of the simpler Florideae. In
both theories the points of connection between the Ascomyceteae and the
Rusts and Smuts and the more characteristic Basidiomyceteae are not in

636 THE PHYLOGENY OF THE FUNGI

agreement, regardless of the ideas as to the ancestry of the Ascomyceteae.
The author follows the school of Sachs in the belief that the evidence is
stronger in favor of the Floridean ancestry of the Ascomyceteae, but he
will attempt to indicate the viewpoints of this hypothesis which are most
unsatisfactory and to exhibit the evidence that convinced de Bary and
many of his successors that the Phycomyceteae actually gave rise to the
higher fungi.

Among the points against the Floridean ancestry, the composition of
the cell wall has been emphasized. In the higher fungi as well as the lower
fungi the basic compounds making up the wall are various types of carbo-
hydrates, among which may be found cellulose, callose, various pectin
compounds, etc. Admixed with these, especially as the mycelium becomes
older, are various substances which, when in sufficiently large proportion,
prevent the reaction of the cell wall to the various cellulose tests, such as
chloriodide of zinc and certain stains and solvents. These admixtures are
sometimes fats, deposited in the surface layers of the wall, as is the case in
Monohlepharis, Blasiocladia, etc. In these, treatment with substances that
will saponify the fat (such as KOH solution, warm but not hot, followed
by thorough washing) will then permit the cellulose reaction to appear.
Far more often the hindering agent is a substance which, although it is
usually called chitin, has been shown to be sufficiently different from
animal chitin to warrant its designation as fungus chitin (see Chapter 1,
p. 3). This is found in some of the Chytridiales (e.g., Synchytrium) , to a
small extent in Phytophthora, but not enough to prevent the cellulose reac-
tion entirely, and in the Mucorales and Entomophthorales. Even in the
former it may not block the cellulose reaction in the younger hyphae but
usually does so in the more mature mycelium. The author's investigations
on Piloholus (1948) have shown that in P. kleinii van Tiegh. the cellulose
coloration by chloriodide of zinc is limited to the slightly thickened lip
surrounding the opening at the top of the subsporangial vesicle after the
sporangium has been discharged, while in P. longipcs van Tiegh. the lower
two-thirds or three-quarters of the sporangiophore and the trophocyst,
the basal swelling, and much of the vegetative mycelium show this reac-
tion. In the Entomophthorales the chitin appears to make up a large
portion of the cell wall but cellulose prevails in Basic/ iobolus (Couch,
1939). In the Ascomyceteae and other higher fungi, fungus chitin usually
blocks all cellulose tests although the presence of the latter may be shown
by chemical analysis. In the ascogenous hyphae of some lichens the cel-
lulose reaction shows up well. Some of the yeasts do not indicate the
presence of fungus chitin though cellulose seems to be replaced by other
carbohydrates. From the foregoing it seems that there is a growing trend
as we pass from the lower fungi to the higher fungi for an increased
amount of fungus chitin in proportion to the cellulose but in general the

ORIGIN OF ASCOMYCETEAE 637

higher fungi are much like the lower fungi in cell wall composition. In the
Florideae cellulose is present usually accompanied by a considerable
amount of pectic substances but there is no evidence of fungus chitin

The hypothesis of Floridean ancestry of the higher fungi requires that
the phylogenetic evolution of the latter must be assumed to progress from
complex forms (Pezizales, Sphaeriales, etc.) to simpler forms such as
Taphrina, Endomyces, Saccharomyces, etc., contrary to the usual direction
of evolution accepted for most groups of fungi. On the contrary from the
relatively simplj^-built Phycomyceteae the rather simple Endomycetaceae
may be assumed to have arisen without too great difficulty.

The chief arguments for a Floridean ancestry lie in the similarity
between the sexual reproduction in the simpler Florideae and in many of
the Ascomyceteae, including:

1. Nonmotile, naked or at most very thin-walled, spermatia, with a
relatively large nucleus, mostly produced one or more at a time endog-
enously in scattered or crowded antherids. These depend for distribution
in the Florideae upon water currents and in the Ascomyceteae upon
streaming surface layers of water or upon insects.

2. The production of a receptive filament (trichogyne) projecting
from the oogone.

3. The multiplication within the oogone of the zygote nucleus (or of
the paired but not united gamete nuclei) and their passage through out-
growths from the oogone to terminal cells which become carpospores or
asei in the Florideae or Ascomyceteae respectively.

4. Usually, but not always, the formation of enclosing envelopes of
various types around the oogone, sometimes before but more often after
fertilization.

Aside from these reproductive similarities the vegetative structure
shows similarities.

5. The filaments (hyphae) of both groups of organisms mostly grow in
length by the elongation and division of the terminal cells, which however
occurs also in the Phycomyceteae.

6. The septa which arise by circular shelf-like growth from the lateral
wall of the cell do not entirely close in the Ascomyceteae as they do in
the Phycomyceteae (according to Buller, 1933) but leave a central pore
through which the protoplasmic continuity of adjacent cells is main-
tained, a condition which also occurs in the Florideae.

In the Phycomyceteae in general each sexual act results in the produc-
tion of a single zygote: zygospore or oospore (e.g., Monohlepharis, Sapro-
legnia, Peronospora, Mucor, Entomophthora, Zoopage, etc.) but in the
Ascomyceteae in which sexual reproduction occurs by the union of non-
motile sperm cells with a trychogyne the one sexual act leads to the pro-
duction of many asci borne on ascogenous hyphae and usually surrounded

638 THE PHYLOGENY OF THE FUNGI

and protected by a structure made up of vegetative hyphae (apothecium,
perithecium, etc.)- The same is true of the Florideae where the union of
one sperm cell with the trichogyne will bring about the formation of a
spore fruit (as Sachs called it) with many carpospores, either surrounded
by a protective envelop or not. It should be noted that in the very simple
Endomycetaceae and Saccharomycetaceae, considered by the author to
be extreme simplifications from very complex ancestors, a single ascus is
produced by each sexual act. The proponents of the hypothesis that the
Ascomyceteae have arisen directly from the Phycomyceteae naturally
consider the foregoing fungi to represent connecting hnks between the

two groups.

A further argument against the phycomycetous origin of the Asco-
myceteae is the type of mycelium, which is prevaihngly coenocytic in the
former and cellular in the latter. To be sure cross walls occur frequently
in some Phycomyceteae but they are mainly (1) to set off reproductive
organs from the main mycelium, (2) to fence off emptied portions of the
mycelium from those regions in which living protoplasm is still present,
and (3) to wall off injured regions. Furthermore the production of the
septa in the coenocyte is entirely independent of any immediately pre-
ceding nuclear division. In the higher fungi the mycelium is usually
cellular and the cell walls separate uninucleate or binucleate cells. Cell
division and septum formation are the immediate consequences of the
division of the nucleus or of the two nuclei of the cell. It cannot be denied
that the mycelium of many Ascomyceteae is composed of coenocytic seg-
ments separated by cross walls, but these forms are relatively few in
number. On the other hand the somewhat anomalous Basidioholus in the
Entomophthoraceae has its mycelium made up of uninucleate cells.

If we accept Sachs's suggestion as to the possible origin of Asco-
myceteae from the Florideae we must prepare a phylogenetic tree that
will include as more primitive those forms whose characters include the
greater number of those found in the Florideae. Therefore we must give
first place to those families or orders in w^hich receptive hyphae (tricho-
gynes) are fertilized by nonmotile, naked or thin-walled, usually endog-
enously produced sperms and from whose oogones arise several or many
asci or ascogenous hyphae bearing the asci. Thus we should place first in
the Ascomyceteae the Laboulbeniales, Lecanorales, Sphaeriales, and some
Pezizales, granting that many of the last two orders accomplish their
sexual reproduction by direct contact of the antherids with trichogynes
or oogones. All of these forms are rather complex in their structure, far
more so than the very simple yeasts and Taphrinales, which are placed at
the base of the class by so many mycologists. On the same grounds
of reasoning the Uredinales also, must be considered to proceed from
near the orders believed by the author to be primitive in the Asco-

ORIGIN OF ASCOMYCETEAE G39

myceteae. H. S. Jackson (1944) gives very convincing arguments for the
hypothesis that the closely parallel life histories of the Uredinales and
Florideae indicate something more than a mere convergence, probably
true relationship.

From the Pezizales it is usually considered that the Tuberales have
arisen and possibly the Hysteriales. The fructifications of the Asco-
corticiaceae may be looked upon as representing much reduced apothecia,
thin and without a definite margin. They still retain the characteristic
hook or crozier mode of formation of the asci. The family Taphrinaceae
might be looked upon as producing a much further simplified apothecium
in which the hook or crozier are lacking and where the whole vegetative
mycelium is dicaryotic, perhaps representing the ascogenous hyphae of
the usual types of Pezizales.

The Sphaeriales show in the genus Mycosphaerella numerous cases of
the production of rounded oogones with unbranched, nonseptate, apical,
usually uninucleate trichogynes which are fertilized by sperm cells pro-
duced internally (sometimes in fours) in the spermogonial structure. In
this order as in the Pezizales the union of oogone and antherid may take
the place of spermatization of a trichogyne. The Erysiphales represent
mostly Sphaeriaceous forms with usually superficial, nonostiolate peri-
thecia. Probably the Pseudosphaeriaceae arose from typical Sphaeriaceae
as also did the Aspergillales and Myriangiales. The Gymnoascaceae
represent a development in the Aspergillales toward the loosening up of
the perithecial wall which is entirely lacking in the Endomycetaceae as
are the ascogenous hyphae, so that a single sexual act produces but one
ascus. (Fig. 206.)

B. O. Dodge (1914) gave a very scholarly exposition of the similarities
in the sexual reproductive structures of the Ascomyceteae and Florideae
with special reference to the probable origin of the former from the latter.
This should be read in connection with the excellent explanation (by
Atkinson, 1914) of the origin of the simpler Ascomyceteae from the
Phycomyceteae.

For those who look to the Phycomyceteae as the possible source of
the higher fungi several hypotheses have been proposed. For many years
that of Brefeld (1889) was the prevalent one. He started from the assump-
tion that sexuality among the fungi was confined to the Phycomyceteae
and was entirely lacking in the Ascomyceteae and Basidiomyceteae. He
assumed that the ascus is a sporangium homologous to that of the Muco-
rales. Accordingly he considered the polysporous asci to represent a more
primitive stage and the fungi with the standard eight-spored asci as more
advanced. The basidium he considered to be a conidiophore bearing a not
standardized number of conidia in the more primitive forms (the Usti-
laginales), the number finally becoming fixed at four per conidiophore

640 THE PHYLOGENY OF THE FUNGI

OTHER EUBASIDIAE SACCHAROMYCETACEAE

HETEROBASIDIAE CORTICIUM ASCOIDEACEAE

ENDOMYCETACEAE
GYMNOASCACEAE

MYRIANGIACEAE

ASPERGILLACEAE
PSEUDOSPHAERIALES, ETC. / ERYSIPHALES

HYSTERIALES

PEZIZALES a LEGANORALES,

SPHAERIALES S HYPOCREALES
'\
\
\
\
\

-^ IN THESE GROUPS OCCUR
FORMS WITH FREE
AND TRICH0GYNE5

LABOULBENIALES_---^''*^^ ™^''^ ^'^" '^'^^^ ^''"^^

ANCESTRAL RED SEAWEEDS

Fig. 206. Suggested lines of evolution within the Ascomyceteae and origin of the
Basidiomyceteae, based upon Sachs's idea of their derivation from the Florideae.
(After Bessey: Mycologia, 34(4):355-379.)

(i.e., the typical i'our-spored basidium). The "conidial" Phycomyceteae
(some Mucorales, the Entomophthorales, etc.) were looked upon as the
origin of the Basidiomycetous hne. Although this Brefeldian hypothesis
has become discredited with the demonstration of the actual occurrence
of sexuality in the higher fungi and by the general acceptance of the
homologies of ascus and basidium and of the crozier and clamp connec-
tions his theory still exerts a strong influence in most systems of classifica-
tion of these fungi.

With the acceptance of true sexuality in the higher fungi various
mycologists have turned to the morphologically simple families of the
Ascomyceteae as probably representing the intermediate stages between
the Phycomyceteae and the Ascomyceteae. The discovery by Lagerheim
(1892) of Dipodascus albidus and the careful working out by Juel (1902)
of the (;ytology of its sexual reproduction have revealed a form that lends
itself well to consideration as such a link. In this fungus the mycelium is
septate, each segment being coenocytic. Usually adjacent segments pro-
duce upright, multinucleate gumetangia which unite near their apices. A
"privileged nucleus" of each gametangium unites with the corresponding
one from the other to form a large zygote nucleus. From the junction of

ORIGIN OF ASCOMYCETEAE 641

the gametangia or from one of the latter the long ascus develops, tapering
toward its apex. In it the zygote nucleus divides repeatedly and finally
around each nucleus is formed an ascospore. Numerous nuclei degenerate
without acting as centers for spore formation. It is assumed that these
were the supernumerary nuclei of the gametangia which did not become
the privileged nuclei. The interpretation by Dangeard (1907), Atkinson,
Gaumann (1926), and others is that the gametangia are homologous to
those of the Mucorales and that the ascus represents the zygospore of the
latter and the sporangium which it usually produces upon germination
after a period of rest. In Dipodascus, however, there is no resting period
and the zygote develops immediately to become the ascus. In D. uni-
nucleatus Biggs, the mycelium is cellular, consisting of uninucleate cells.
The gametangia are also uninucleate and there are no supernumerary
nuclei. The ascus and ascospores develop as in D. albidus, the number of
ascospores being typically large but often small in poorly developed asci.
This species forms a close connection to Endomyces and Eremascus in the
Endomycetaceae in which single asci, of eight ascospores or less, arise as
the result of the sexual union of the two uninucleate gametangia. By in-
crease of the tendency toward the budding habit of vegetative growth we
can derive the Saccharomycetaceae. It is assumed that in some forms
similar to Dipodascus the elongated multinucleate ascus gave rise to a
series of branching hyphae containing the as yet unfused gamete nuclei
and their products, and producing terminal cells where the pairs of nuclei
unite and undergo meiotic division. Thus is interpreted the origin of the
dicaryon ascogenous hyphae and asci. By the growth of loose protective
vegetative hyphae between and around the ascogenous hyphae it is
assumed that such a loose perithecium is developed as we find in the
Gymnoascaceae. By the development of a firmer cortex the Aspergillaceae
were derived. From these organisms one can imagine the development of
Erysiphales, Myriangiales, Pseudosphaeriales, Sphaeriales, Pezizales, etc.,
a complete reversal of the phylogenetic tree based upon the idea of the
origin of the Ascomyceteae from Florideae. (Fig. 207.)

Whichever of the two phylogenetic series mentioned above is con-
sidered to be nearer the truth there are certain observed facts or interpre-
tations of facts that disagree. In both hypotheses the ascus is postulated
as being derived from a sporangium: for the Floridean ancestry it must
come from a tetrasporangium and for the Phycomycetous derivation from
a sporangium more or less on the plan of that of the Alucorales. In either
case the spores arise by the complete segmentation of the protoplasm,
with no enucleate epiplasm left over. Yet, in the ascus, part of the cyto-
plasm gathers around the nuclei and becomes encysted, leaving a portion
without nuclei, the epiplasm, surrounding the ascospores. Whether or not
this difference in manner of spore formation is so fundamental as many

642

THE PHYLOGENY OF THE FUNGI

have considered, it must be settled by further studies upon the various
types of sporangia in the Phycomyceteae and different groups of algae.
In the Phycomycetous hypothesis of the origin of the Ascomyceteae
there is presented the difficulty of accounting for the production of the
nonmotile sperms and of the trichogynes. Apparently very early in the
development of the Phycomyceteae sexual reproduction took place by the

IN THESE THREE GROUPS OCCUR FORMS WITH

FREE SPERMS AND TRICHOGYNES

\ \ "LECANORALES

PEZIZALES.ETC.

(

ERYSIPHALES

MONASCACEAE

ASPERGILLACEAE

GYMNOASCACEAE

ENDOMYCETACEAE

ASCOIDEACEAE

MUCORALES

SPHAERIALES, PSEUDO-
SPHAERIALES, ETC.

MYRIANGIACEAE

ELAPHOMYCETACEAE. ETC.

SACCHAROMYCETACEAE

Fig. 207. Suggested lines of evolution within the
Ascomyceteae, following in the main the ideas of
Dangeard, Atkinson, Gaumann, Mez, et al. (After
Bessey: Mycologia, 34(4) -.355-379.)

union of two equal-sized motile gametes, as in some of the Chytridiales
and Blastocladiales, by the union of two motile gametes of unequal size, as
in some species of Allomyces, or by the union of a flagellate sperm with a
nonflagellate egg, as in Monohlepharis. In all the other Phycomyceteae in
which sexual reproduction is known, this occurs by the union of two non-
motile cells (Olpidiopsis) or by the passage of nuclei and cytoplasm from
one gametangium to another one through a simple opening or through a
conjugation tube. In Dipodascus, Endomyccs, and most of the forms con-

ORIGIN OP ASCOMYCETEAE 643

sidered to be the primitive Ascomyceteae, if their Phycomycetous origin
is accepted, we have this same general method of sexual reproduction, but
in the forms considered under that hypothesis to be the furthest advanced
we find separate nonmotile sperms uniting with special receptive organs
(trichogynes). Whence have these been derived? They represent an
entirely new development. Attempts have been made to consider the
spermatia to be specialized conidia that have assumed the sexual function.
That might possibly be so but it is far easier to surmise that they are the
reproductive structures inherited from the Floridean ancestors where
that is the standard mode of sexual reproduction.

The case of Liagora tetrasporifera Borgesen (1927) indicates how a
short-cycled red seaweed has developed a life history very closely parallel
to that of such an Ascomycete as some species of Mycosphaerella. In this
representative of the Order Nemalionales, after spermatization of the
trichogyne there soon grow out from the oogone a number of gonimo-
blasts whose terminal cells instead of becoming carpospores, as is the
usual case in red seaweeds, become tetrasporangia which produce four
tetraspores each. Although in this particular case the cytological behavior
of the nuclei in the oogone after fertilization and in the tetrasporangium
has not been studied it may well be assumed, from the analogy of other
Nemalionales and other orders of the Florideae, that the gamete nuclei
united normally in the oogone but did not undergo immediate meiosis, the
result being that the nuclei in the gonimoblasts remained diploid. The
terminal cells instead of becoming carpospores became tetrasporangia in
which meiosis occurred and tetraspores with haploid nuclei were pro-
duced. Compare this with Mycosphaerella. A uninucleate sperm cell
spermatizes a one-celled unbranched trichogyne and passes down into the
oogone where it associates itself with, but does not unite with, the egg
nucleus. The two nuclei then divide simultaneously by "conjugate divi-
sion" until many pairs of nuclei are produced which pass out into the
ascogenous hyphae. At or near the tip a pair of nuclei passes into a ter-
minal or subterminal cell where they unite. Then this diploid nucleus
undergoes meiosis to produce four haploid nuclei and, after the fashion of
the majority of the Ascomyceteae, a further mitotic division occurs, so
that eight ascospores arise in each ascus. It should be remembered that in
many Ascomyceteae where typical sexuality is known only four nuclei
and therefore not over four ascospores are produced in each ascus. Al-
though one cannot assume that Liagora is a direct ancestor of the Asco-
myceteae this shows that in the red seaweeds the conditions exist now
that might have given rise ages ago to this group of fungi.

Possibly the acquisition of the land habit might have led to the pro-
duction of ascospores in an ascus where the epiplasm is of importance in
the scattering of the spores, whereas the tetraspores which depend upon

644 THE PHYLOGENY OF THE FUNGI

the water currents for their distribution sHp out from the tetrasporangium
as naked cells.

Origin of Rusts and Smuts and Heterobasidiae

As mentioned above, Jackson (1944) has emphasized the parallelism
of the life history types in the Uredinales and the Florideae. He has
demonstrated beyond chance of contradiction that the various types of
life history, such as alternation of haploid and diploid generations and
shortened life cycles occur in both groups of organisms, together with a
number of structural similarities (spermatia, receptive hyphae, etc.)-

In view of the extreme, obligate type of parasitism of the rusts and the
usual nonparasitic mode of life of the red seaweeds, and of the type of
teliospores with promycelium — characters not indicated in the latter
group — it appears evident that the relationship is not necessarily very
close. Indeed the structure and function of these organs seem to indicate
relationship with the Basidiomyceteae under which they are more often
now classified. Bound up with the Uredinales is the question as to the
relationship of the Ustilaginales with which they have very often been
associated in classification. There is no denying the fact that in very
many regards the latter order is far less specialized in structure and life
history than the former. Yet there are a number of characters common to
both : (1) a high degree of intercellular parasitism confined to hosts among
the vascular plants; (2) a more or less well-marked distinction of two
types of mycelium, sometimes associated with special types of spores and
hosts, the monocaryon and dicaryon types of mycelium; (3) the produc-
tion on the dicaryon mycelium of binucleate cells, usually with colored
and thickened walls, in which the two nuclei unite ; (4) the outgrowth from
this teliospore of a thin-walled hypha (promycelium) of limited growth
within which meiosis occurs and upon which are then borne the sporidia
w^hich are of two or four sexual phases; (5) the production by the latter
of the monocaryon phase of mycelial growth; (6) the possession of various
methods by which this mycelium becomes diploidized to form the di-
caryon mycelial phase which produces the teliospores. Both orders give
rise to repeating spores, i.e., asexual spores which repeat the mycelial
phase upon which they are borne. In the Uredinales these are known onl,y
upon the dicaryon mycelium (being known as urediospores) but in some
of the Ustilaginales they occur on both monocaryon and dicaryon types
of mycelium.

In some regards the Ustilaginales appear the more primitive in that
their dicaryon mycelium often has typical clamp connections, while these
have been definitely discovered in only a few cases in the Uredinales. In
the latter order there are present typically definite spermogonia which
produce spermatia capable of spermatizing receptive hyphae (tricho-

ORIGIN OF RUSTS AND SMUTS AND HETEROBASIDIAE 645

gynes) and thus diploidizing special cells from which are produced typical
dicaryon spores (aeciospores). In the Ustilaginales the diploidization may
occur in various ways but does not include spermatia and receptive
hyphae.

If, then, the two orders Ustilaginales and Uredinales are considered to
be related we must conclude that they have arisen probably (1) from a
group of fungi in which clamp connections occurred or their homologues,
the croziers, characteristic of ascus formation; (2) from fungi in which
spermatia were produced in well-organized spermogonia; (3) from fungi
in which sexual reproduction was initiated by the spermatization of recep-
tive hyphae ; and (4) from fungi in which the monocaryotic and dicaryotic
mycelial phases were both present. Clamp connections are found in
most groups of the Basidiomyceteae and in some lichens while their
homologues, the croziers, occur in Lecanorales, Pezizales, Tuberales,
Sphaeriales, Aspergillales, and in the Ascocorticiaceae and a few other
forms. They are absent in the Laboulbeniales, Erysiphales, Saccharo-
mycetales, and a number of other groups. Spermogonia are found in
Laboulbeniales, Lecanorales, Sphaeriales, Pyrenulales, Dothideales, and
some Pezizales. Special receptive hyphae (trichogynes) are found in
Lecanorales, Pezizales, Laboulbeniales, Sphaeriales, Hypocreales, Py-
renulales, Dothideales, and a few other groups. The two mycelial phases
are absent in the Saccharomycetales and Taphrinaceae. We must there-
fore consider that the Rusts and Smuts may have been derived from some
Ascomyceteae possibly before the Pezizales, Sphaeriales, and Dothideales
had become strongly distinguished from one another. Possibly these early
ancestral Ascomyceteae if they were derived from Florideae that were
diplobiontic might have had their two mycelial phases much more
strongly marked than in the forms now known, in which the dicaryon
phase has been reduced to or has not yet proceeded beyond the formation
of the ascogenous hyphae.

Linder (1940), in a very thought-provoking discussion, has sought to
derive the LTredinales from forms near the Dothideales. He assumed that
the disappearance of clamp connections or croziers may have been con-
nected with the parasitism entirely within the host tissues. He believed
that the Ustilaginales (which still retain clamp connections) were derived
from the Uredinales with reduced life cycles in which loss of spermatia and
receptive hyphae had been recompensed by the ability of any two cells of
opposite sexual phase to fuse and thus initiate the dicaryotic type of
mycelium. It must be remembered that even in the Uredinales there
appears to develop mutual diploidization of mycelia of opposite sexual
phase when they meet in the tissues of the host, even though spermatiza-
tion of receptive hyphae is prevented.

Linder's view of the origin of the teliospore and promycelium from the

646

THE PHYLOGENY OF THE FUNGI

Fig. 208. Diagram showing Linder's suggestion as to the origin of the teliospore
and promyceUum of Uredinales from an ascus. (Courtesy, Linder: Mycologia, 32(4):
419-447.)

ascus assumes that the type of ascus was that with a two-layered wall,
such as occurs in Pleospora, Leptosphaeria, etc. The outer layer is firm and
the inner layer thin and elastic. Upon the softening of the apex of the
ascus where there is a germ pore the inner layer expands so that the endo-
ascus and contained ascospores project some distance beyond the outer
layer and eventually ruptures to set the spores free. He assumes that such
an ascus became more thick-walled so as to become a resting spore but
with the oncoming of favorable conditions the inner portion pushed out as
described above, the meiotic division occurring usually within this pro-
jection. The contents of this extruded portion (promycelium or basidium)
divide to form four spores by simple cross walls instead of by free spore
formation as is usual in the Ascomyceteae. These four spores remaining
attached to each other permit their contents to escape by forming ex-
ternal sterigmata from which secondary spores (sporidia or basidiospores)
are set free. (Fig. 208.)

Accompanying the gradual loss of obligate parasitism and simplifica-
tion of the life cycle the Uredinales according to Linder led to the Auricu-
lariales, probably to those forms in which a "probasidium" is present
such as Septohasidium and Helicohasidium. With the complete loss of the
formation of a distinct probasidium, forms such as the entirely sapro-
phytic Auhcularia could be reached. In other more or less similar ways he
postulates the origin of the Tremellales and from them the Autobasidio-
myceteae (Eubasidiae).

ORIGIN OF RUSTS AND SMUTS AND HETEROBASIDIAE 647

Another view as to the origin of the Heterobasidiae (excluding the
Uredinales and Ustilaginales) is that they may have arisen from some
Ascomycetous form somewhat hke Ascocorticium. In this fungus the asci
are developed with the formation of croziers. There are no conspicuous
paraphyses and between the wood or bark on which the fructification is
produced there are only a few layers of hyphae running parallel to the
surface. From the outer hyphae arise the asci. There is no definite limiting
margin to this structure which may be looked upon as a much simplified
apothecium. It is not known whether the subhymenial hyphae are di-
caryotic or monocaryotic or a mixture of the two or whether the mycelium
within the substratum is one or the other. Also, it is not known whether
spermatia and receptive hyphae are present or how the dicaryophase
arises. Therefore we do not have much on which to base a suggestion that
a fungus of this structure is a possible forerunner of the Corticium-Yike
Basidiomyceteae. If we do make this assumption, we must account for
the origin of the basidium (holobasidium in this case) from the ascus. It
is known that in all fungi with holobasidia the meiotic division of the
diploid nucleus of the young basidium occurs within the latter until four
nuclei, or very often eight through a subsequent mitotic division, are
produced. Then there arise slender outgrowths from the upper portion of
the basidium and these sterigmata enlarge terminally. At this point the
nuclei, usually one to each, squeeze through the slender sterigmata and
round up in their enlarged ends. Then, and not until then, is another wall
laid down within this "spore." This wall is always visible separating the
opening of the sterigma from the spore and frequently can be traced all
over the inner surface of the outer spore wall, sometimes not tightly
appressed to it. In other words, after the nucleus enters the terminal
swelling of the sterigma it proceeds to induce the formation of a wall
around the nucleus and accompanying cytoplasm. We may then interpret
such a basidium as an ascus on which arise evaginations or external
pockets within which the ascospores are produced. The so-called basidio-
spores therefore may be looked upon as ascospores enclosed within the
evaginated ascus pockets. This is somewhat different from the suggestion
of Linder by whom the septate promycelium of the teliospore was looked
upon as a row of four ascospores, separated by simple septa, not rounded
up as occurs in a normal ascus. From each such ascospore a short stalk
(sterigma) produces a basidiospore which must then be looked upon as a
secondary spore, budded off from the ascospore. Rogers (1932, 1934) looks
upon Tulasnella as perhaps derived from an Ascocorticmm-like ancestor.
He interprets the four large pockets at the top of the basidium as ascus
pockets, each containing the homologue of an ascospore. The spore pro-
duced on the slender sterigma from such a pocket is then, as in Linder's
hypothesis, not homologous to an ascospore but, as Linder intimated, a
secondary spore budded off from the sterigma sent out by the ascospore.

648 THE PHYLOGENY OF THE FUNGI

The fact that the asymmetrical manner of attachment of the basidio-
spore to the sterigma is found throughout the Heterobasidiae (except for
a few "angiocarpous" forms) and in the Polyporales and Agaricales
probably indicates that they all have a common origin, for it is unlikely
that so intricate a mechanism for spore discharge has been polyphyletic
in origin. Since the same structure occurs in the Uredinales and in the
Tilletiaceae these too must be descended from the same ancestors. There
is considerable evidence for the surmise that the Basidiomyceteae with
symmetrically attached basidiospores which are not shot off from the
tips of the sterigmata are secondary modifications from the asymmet-
rical types, associated with the angiocarpous development of the spore
fruit (but see below for discussion of the Agaricales-Gasteromyceteae
relationships).

If then the Uredinales (and Ustilaginales) are of common phylogenetic
origin with the Heterobasideae and the Eubasidial Hymenomyceteae we
must decide whether the heterobasidium or eubasidium is the more primi-
tive. Linder's suggestion has been mentioned above that from the Ured-
inales arose all other types of Basidiomycetous fungi. Rogers, on the
contrary, would derive the Uredinales from Tulasnella, Tremella, and
Auriculariales, while deriving the Eubasidiae in another direction from
the same primitive genera. If either of these hypotheses is correct then
the objects that we call basidiospores (or sporidia) are not homologous
with ascospores but are secondary spores produced from them to provide
for aerial distribution, the homologues of the ascospores being the indi-
vidual cells of the transversely or longitudinally septate basidium. If
Rogers is right in postulating the change in position of the four septa in
Tulasnella so as to divide the ''probasidium" or "hypobasidium" ver-
tically as in Tremella and that then by a further shifting of the septa
(intermediate forms seem to occur) they become transverse, as in Auricu-
laria, we still must insist upon the homology of the external basidiospores
in all these cases. If the Uredinales arose from the Auriculariales by the
emphasis on the development of the "probasidium" into a thick-walled
resting spore, as occurs in many species of Septobasidium, the sporidia of
these too are homologous to the foregoing basidiospores, i.e., are second-
ary spores. The close relationship of the Auriculariales, Tremellales,
Dacrymycetales, Tulasnellales, Uredinales, and Ustilaginales would seem
to be emphasized by the wide-spread occurrence of the production of
secondary spores from the basidiospores (sporidia) by budding. The
germination of these spores directly by single germ tubes may occur but
in water or damp air they may give rise to yeast-like or to allantoid buds,
often accompanied by transverse divisions of the basidiospore. Even in
the Uredinales (e.g., as observed by the author in Kunkelia nitens) the

ORIGIN OF RUSTS AND SMUTS AND HETEROBASIDIAE 649

sporidia in damp air while still attached to the promycelium may produce
successively secondary, tertiary, and further sporidia, all obliquely
perched on short sterigmata from the sporidia below. In addition there is
a great tendency toward the production of spore fruits whose cell walls
swell and become gelatinous when w^et, although this is not universal-. In
the Uredinales we find this in Gymrwsporangium, Coleosporium, Uropyxis,
etc., chiefly in the teliospores and their pedicels. In the Auriculariales,
Tremellales, Dacrymycetales, and some Tulasnellales so many genera have
this character throughout the w^hole spore fruit that they are often
grouped under the common name "Jelly Fungi." Both of the foregoing
characters are rare in the Eubasidiae. However in Exohasidium the
basidiospores may divide transversely and produce external buds but this
is scarcely known in any other Eubasidial genera. Scattered here and
there are occasional species with gelatinous cell walls but these are
marked exceptions.

If the Thelephoraceae are to be considered as derived from near
Tulasnella it must be by the elimination of the cross walls between the
main body of the basidium and the four large spore-bearing pockets, ac-
companied by a reduction in size of these pockets until finally in most
cases these have entirely disappeared leaving merely their apical sterig-
mata. In that case the homologues of the ascospores have, as it were,
drawn back into the body of the basidium and entirely disappeared. If
this is the way that evolution occurred we must still regard the basidio-
spores as secondary spores from hypothetical ascospores, as in the Tulas-
nellales, etc.

Let us consider the case if the holobasidium is the more primitive type,
as it is beyond controversy the most freciuent in the whole class. We can
still derive it theoretically from one of the Pezizales somewhat similar to
Ascocorticium, although probably spermatia were still produced, perhaps
even in spermogonia. We can imagine that the basidiospores, as suggested
by the author on page 647, are ascospores, in some cases provided with
true ascospore walls, inside the everted pockets. The sterigmatal ap-
paratus for their discharge rnay have to be looked upon as a new develop-
ment to permit their being discharged violently, although perhaps this is
a modification of the discharge mechanism found in most asci in which
four or more spores contained in the ascus are discharged simultaneously
or in quick succession through the single opening at the apex of the ascus.
This is brought about by the increase of the osmotically produced tension
of the ascus wall until it gives way at the apex and the epiplasm and
spores are violently ejected. In the new modification as found in the
basidium the discharge affects only one spore at a time at the apex of each
sterigma. This new habit is clearly not confined to the primary spore for

650 THE PHYLOGENY OF THE FUNGI

in the case of Kunkelia mentioned above the secondary and tertiary spores
are similarly discharged and in the Tilletiaceae this seems to extend to
secondary sporidia and even to conidia in some species. So we might
imagine that this might have been a habit acquired for asexual spores
that has been extended to the sporidium and basidiospore as well.

If the holobasidium was a primary step in the evolution from the
Ascocoriicium-like Ascomyceteae we can assume that in some cases the
normally slender sterigmata as in Corticium became stouter, bearing at
their tips the points on which the basidiospores perch and finally attaining
the separable condition found in Tulasnella. The basidium itself may be
considered to have formed vertical walls as in Tremella and finally trans-
versely septate walls would bring about a structure like the basidium of
Auricularia. From Ceratohasidium which has stout sterigmata, only two
in number in some species, it is an easy step to Dacrymyces.

From the simple, resupinate forms like Corticium, Tomentella, etc. it
is easy to make surmises as to the modifications which increase the hy-
menial surface so as to derive the Hydnaceae, Clavariaceae, Polyporaceae,
and eventually Agaricaceae.

In the immediately preceding paragraphs we have left the Uredinales
hanging without a connection. They cannot be derived from any Auricu-
lariales such as we now know, for well organized spermogonia and recep-
tive hyphae are lacking in the latter. We can only guess that the trans-
formation from ascus to holobasidium occurred deeper down in the
Ascomyceteae than the present Ascocorticium and that the development
of holobasidium to phragmobasidium took place perhaps at several levels
or when once accomplished proceeded in various directions. The earlier
Uredinales must have come from the newly developed holobasidiomycete
while the latter still retained the primitive characters of its ancestors,
including spermogonia, receptive cells, diplobiontic life cycle, croziers or
clamp connections, and probably well pronounced parasitism. We must
assume that at an early stage of development along this line thick-walled
resting probasidia began to develop. This would be the beginning of
teliospore formation. On resuming their growth these probasidia would
grow out as thin-walled promycelia except in such cases as Coleosporium
where the teliospore wall was thin enough to permit of stretching as a
whole, thus producing a so-called "internal promycelium" instead of the
usual external one characteristic of the order. We need not homologize the
separate cells of the promycelium whether external or internal with
ascospores but consider these cells as parts of a phragmobasidium. Under
this assumption the sporidia may be considered as ascospores in external
pockets not as secondary spores. It must be admitted that this whole
suggestion stands on shaky foundations and the whole matter needs
further study. (Fig. 209.)

PHYLOGENY OF HYMENOMYCETEAE AND GASTEROMYCETEAE

651

SEPTOBASIDIUM
\

AURICULARIA

AURICULARIALES

TREMELLALES

TULASNELLA

DACRYMYCETACEAE

CERATOBASIDIUM

CLAVARIACEAE

HYDNACEAE c..

LEVEL OF BASIDIUM ORIGIN

DISCOMYCETEAE

FLORIDEAE

Fig. 209. Diagram showing the author's ideas as to the possible origin of and
relationships between the chief families of Subclasses Teliosporeae, Heterobasidiae,
and Hymenomycetous Eubasidiae.

Phylogeny of Hymenomyceteae and Gasteromyceteae

Another very uncertain phylogenetic tree is that concerned with the
relationship of the Gasteromyceteae with the Hymenomyceteae. The
former, be it remembered, differ from the latter by the fact that the spore
fruits do not open and permit the escape of the spores until after the
basidia and basidiospores have reached maturity, while in the latter the
hymenium becomes exposed to the air before the basidia have produced
their spores. Associated with this Gasteromycetous structure of the spore
fruit is the fact that the basidiospores are not shot off from the sterigmata

652 THE PHYLOGENY OF THE FUNGI

at whose tips they stand symmetrically perched. Frequently the sterigma
breaks at some distance below the spore and remains attached to it like a
handle. The spore fruits vary in simplicity or complexity of structure from
very small bodies with a single closed hymenial cavity as in Protogaster up
to the very complex structures with a columella which may be extended
into a stalk as in Secotium, Podaxis, Battarrea, etc. Some remain closed
until some accident (decay, attack by insects, etc.) breaks them open and
permits the spores to escape, others provide definite modes of spore-
escape, e.g., Lycoperdaceae, Sphaerobolaceae, Phallaceae, etc. Yet in all
cases the spores are mature before they can escape and are symmetrically
attached to the sterigmata. Yet aside from these differences there are
many points of structural similarity, as pointed out in Chapter 15,
between some Gasteromyceteae and some Agaricaceae. Russula and
Lactarius have anatomical features and spore markings and chemical re-
actions very much like Elasmomyces, so that their close relationship can
be assumed unless we surmise this to be a case of convergent evolution.

Origin of Agaricales

Singer (1950), in a chapter entitled " Phylogenetic theories concerning
the origin of the Agaricales," discusses three hypotheses that have been
proposed to explain the origin of this order: (1) they are directly and
wholly derived from the Aphyllophorales (Polyporales) ; (2) they are
directly and wholly derived from the Gasteromyceteae, probably along
several lines; (3) they are partly derived from the Polyporales and partly
from the Gasteromyceteae. He, himself, is in favor of the second hypoth-
esis but his explanations of the other two are good. Taking these hypoth-
eses in turn the author will give a resume of the facts that stand in favor
of and against each one.

Singer^ s Hypothesis No. 1

Agaricales Derived Wholly from Polyporales. This is perhaps the
oldest theory since the time that the idea of evolution began to affect the
classification of fungi. Briefly sketched, it assumes that from a simple
resupinate spore fruit with a smooth, or possibly rather loose, cottony,
hymenium, evolution progressed toward more compact structures some
of which assumed a lateral position or even became stipitate. These forms
with flat hymenium made up the old family Thelephoraceae. The amount
of hymenial surface in proportion to the size of the spore fruit was limited,
so long as the hymenium was a plane surface. This surface became in-
creased by the formation of hymenium-covered pegs or teeth projecting
from the surface, leading to the Hydnaccae. In a slightly different manner
the surface became increased by a reticulate pattern of outgrowth so that
eventually there were very numerous pits or pores of various depths, lined

ORIGIN OF AGARICALES C53

by hymenium, the Polyporaceae. By the radial elongation of these pores
so that they extended from the point of attachment (more often a central
stipe) to the edge of the spore fruit a system of gills or lamellae was
developed, the Agaricaceae. The lateral branches producing the Exo-
basidiaceae and Clavariaceae were not in the main line of upward evolu-
tion and need not be given further consideration in this hasty review. Up
to the Agaricaceae and the closely related Boletaceae practically all the
foregoing fungi were gymnocarpous in their ontogeny. The hymenium
w^as developed externally over the whole surface of the spore fruit or over
part of the surface beginning as a plane which in the more advanced forms
became thrown into teeth, pores, or gills. Naturally, therefore, those
Agaricaceae were considered the most primitive in which this gymno-
carpous ontogeny continued. Upon study of the development of the
fruiting body from very early stages of growth it has been shown that in
many Agaricaceae and in the Boletaceae the primordium of the hymenium
is laid down gymnocarpously, but that by the outward spreading and
downward and inward curving of the edge of the pileus this hymenial
surface is finally enclosed in a circular tunnel surrounding the top of the
stipe and sealed from the exterior by the inturned edge of the pileus which
has grown fast to the stipe. Into this circular space the gills grow from the
pileus and the basidia begin to form on them but before they have ma-
tured sufficiently to produce their spores the edge of the pileus breaks
loose from the stipe and pulls away from it so that the gills at their
maturity are exposed to the exterior. This is called the pseudoangio-
carpous mode of development. Beyond this type of development we find
many Agaricaceae in which the hymenial primordium originates in a
circular layer entirely closed from its first inception within the tissues of
the spore fruit. Not until the gills have reached almost their full develop-
ment does the growth of the fruiting body bring about a circular rupture
when the expanding pileus pulls away from the stipe, so that finally the
gills are exposed to the air and the basidia shed their spores. This angio-
carpous mode of development is found especially in those genera with a
volva or with a well-developed annulus or cortina or a universal veil, even
though this may be rather thin. Since the Gasteromyceteae have a perid-
ium that in some forms, at least, seems homologous to the universal veil
and since, as mentioned above, other Agaric-like structures are present in
some members of this group it has been suggested that these represent
further stages of evolution from the Agaricaceae. Thus Heim (1937, 1948)
suggested that the Asterosporales are a closely related group, containing
the genera Russula and Lactarius of the Agaricales and Arcangeliella,
Elasmomyces, Maccagnia, and some others usually included in the Gas-
teromyceteae. He indicates his belief that these last mentioned genera
have been derived from Russula and Lactarius.

654 THE PHYLOGENY OF THE FUNGI

The chief merit of this suggested system is that the Agaricales appear
to be an extension, as it were, of the Polyporales with a gradual increase
in complexity of ontogeny from strict gymnocarpy to pseudoangiocarpy
and finally to complete angiocarpy. From the latter it would seem logical
to extend this development to the Gasteromyceteae. Here, where the
spores are set free from the basidia before the spore fruit opens we find
that they are symmetrically attached to the sterigmata or even practically
sessile on the surface of the basidium.

There are two obstacles to this system. In the first place the type of
the trama and character of the basidia and cystidia are in the main quite
different in the Agaricales and Polyporales. So that even where, exter-
nally, similarities and apparent transitions seem to occur careful study of
the anatomy and chemical reaction of the hymenium and trama show that
these are perhaps better explained as morphological convergences but not
true relationships. Furthermore it would seem more reasonable to expect
that the Agaricaceae which possess a well-developed universal veil, as
revealed by the production of volva, annulus, etc., and which have an
angiocarpic mode of development, would be far more likely to have arisen
from the mostly strictly angiocarpic Gasteromyceteae and that the reduc-
tion and final disappearance of a universal veil and the appearance of
pseudoangiocarpy and finally of gymnocarpy would indicate a gradual
degeneration from the complex to simplified forms, now that the firm
universal veil (peridium) had become no longer necessary. A third and
very strong objection is that the Gasteromyceteae that show the closest
similarity to the Agaricaceae lead by gradually simplified structures to
the forms assumed by almost all mycologists to be the simplest ones, the
forms that well may have developed from organisms related to Aleuro-
discus, Corticium, Tomentella, etc.

Singer^s Hypothesis No. 2

Agaricales Derived Wholly from Gasteromyceteae. This is the hypoth-
esis, favored by Singer and at least in part by Bucholtz (1903), that the
Polyporales do not give rise to the Agaricales; the latter arise from the
Gasteromyceteae and at some points, mostly by convergent evolution,
assume forms externally similar to some of those of the Polyporales. Its
chief virtue is that it explains the presence of the universal veil and of
angiocarpy in some of the Agaricales and allows a logical explanation of
their disappearance as the rather definite affinities to the Gasteromyceteae
become more distant. The undeniable close similarities of some Agaricales
to some Polyporales is difficult to explain away. To the author the most
outstanding objection to this hypothesis is the necessity of assuming the
evolution anew of the sterigmatal function of spore discharge at several
points, for if this hypothesis is adopted the Agaricaceae probably arose

ORIGIN OF AGARICALES 655

from the Gasteromyceteae at several different places. In this viewpoint
the simplest organisms of this group are supposed to have originated from
simple Thelephoraceae, which already had their spores perched asym-
metrically on the tips of the sterigmata from which they are discharged.
Several cases are known where near relatives of the Tremellales have
adopted the angiocarpous structure and with it have lost this peculiarity
of sterigmatal discharge. It is difficult however to understand how so
complex a structure could have been lost at the beginning of the Gastero-
myceteae and regained several times independently as some Agaricaceae
developed from them.

Singer's Hypothesis No. 3

Agaricales Derived Both from Polyporales and from Gasteromyceteae.

This is really a sort of compromise. Its chief objection is that we have to
find a line of separation within the Agaricaceae between those descended
from the Polyporales and from the Gasteromyceteae. This is perhaps
even more difficult than the first or second hypothesis.

The relationships within the Gasteromyceteae as well as their origin
are very uncertain. To be sure Eduard Fischer (1933 and earlier) proposed
a system that is fairly logical, progressing from simpler to more complex
structures, and this has been the basis for much of the work in this group
for the last fifty years. The characters that have been considered as the
more important are the modes of development of the gleba, the progres-
sive development of the columella (and its downward extension, the stipe)
and the tendency to progress from a coralloid to a multipilar and eventu-
ally unipilar structure. Other important characters are the development
of the capillitium, the formation of definite hymenial cavities or their
obliteration by ingrowing hyphae (plectobasidial structure), the auto-
digestion of the gleba to an evil smelling m9,ss attractive to insects,
angiocarpous or pseudoangiocarpous development, etc.

One very important point, in the author's opinion, that has been sub-
ordinated to a secondary position is the character of the spores. It appears
to him that there are several types of spores that cut across the family
and ordinal boundaries as customarily recognized. These, perhaps, are of
much greater importance and it may be that they indicate closer relation-
ships than have been recognized. Besides spore structure a much more
extensive study of the very early stages of the formation of the spore
fruits is absolutely essential before a well-grounded system can be
established.

In the genera G aster ella, Hymenog aster, and Gasterellopsis we have,
respectively, very small, unicameral spore fruits, a moderate sized organ-
ism with very numerous hymenial chambers with a more or less coralloid
arrangement, and a small, at first unicameral, structure with a percurrent

656 THE PHYLOGENY OF THE FUNGI

columella and gill-like outgrowths from the top of the chamber which may
divide it into several radial cavities. The spores in all three are ovoid or
limoniform, dark-colored and symmetrically perched on the sterigmata.
Many of the spores when set free bear at their base a piece of the upper
end of the sterigma. In their spore character they are practically identical.
The first two could find a place in the same family, Hymenogastraceae,
but the third with its percurrent columella and radial hymenial chambers
and basally circumscissile dehiscence shows some characters that might
be considered as belonging to the Secotiaceae. This is merely an example
to show how, possibly, the current classifications are faulty.

Phylogenetic System of Gasteromyceteae

In default of further studies on the anatomy and ontogeny of the very
young spore fruits and of intensive comparison of spore types throughout
the whole group the following may be suggested as a tentative phylo-
genetic system of the Gasteromyceteae. The primitive (or the simplest)
forms are minute, with one closed hymenial chamber. This might be
supposed to have arisen from a small Aleurodiscus-like fungus in which
the upward curvature at the edges continued until a closed cavity was
produced, lined by the hymenium. As seems to be the case when angio-
carpy develops the basidia come to bear sterigmata with symmetrically
attached spores. The genus Protogaster represents one of these small,
unicameral fungi. Much like this, but with an entirely different type of
spores, is Gasterella. In the spore fruit of this a somewhat arched hymenial
primordium arises angiocarpously and gradually a basidial layer appears
with the basidia directed downwards to line a shallow cavity which soon
becomes larger and nearly spherical. In the more vigorous specimens the
roof of this cavity may be thrown into folds and convolutions which do
not reach to the base so that although the hymenial surface is increased
the cavity is not divided. At this stage it resembles closely the early
developmental stages of Hymenogaster as described by Rehsteiner (see
Chapter 15). Probably the next step beyond the unicameral condition
arose by the increase in the folds and convolutions to form many cavities
lined by hymenium instead of just the one. In the Ilysterangiaceae the
coralloid development is much more marked along with the production of
a more or less pronounced columella. The cartilaginous and gelatinous
character of the gleba is possibly a modification to permit the dissolution
of the spore fruit at maturity. In the Lycoperdaceae the glebal tissues
enclosing the numerous hymenial cavities dissolve after the spores have
become mature, leaving one large cavity filled with the spores entangled
in the filamentous capillitium. By the flaking off of the outer layer of the
peridium and the formation of an ostiole in the inner peridium or some
other mode of dehiscence the spores are enabled to escape and be scat-
tered by air currents.

PHYLOGENETIC SYSTEM OF GASTEROMYCETEAE

657

AGARICACEAE
j?

SECOTIACEAE

PODAXACEAE

TULOSTOMATACEAE

EVOLUTION UPWARD TO AGARICACEAE

DOWNWARD TO PROTOGASTER

AND GASTERELLA

HYDNANGIACEAE

GEASTRACEAE

LYCO PER DACE AE

MELANOGASTRAGEAE

SPHAEROBOLACEAE PHALLACEAE
NIDULARIACEAE

CLATHRACEAE
HYSTERANGIACEAE

SCLERODERMATACEAE

RHIZOPOGON
HEMIGASTER / HYMENOGASTER

GASTERELLA

PROTO-
GASTER

GASTEROMYCETEAE

Fig. 210. Diagram showing the possible relationships within the Gasteromyceteae,
based largely upon Fischer (1933), with some additions.

The plectobasidial structure of some Gasteromyceteae appears, in the
author's opinion, to have arisen at many points in evolution by the failure
of the hymenium to form a distinct layer bounding each hymenial cavity
when produced. In its place the basidiogenous hyphae grow unequally
and fill these cavities with a loose mass of hyphae bearing scattered
basidia. Since this appears in one or two species of Sphaerobolaceae while
other species have normal hymenial cavities, in Astraeus while the rest of
the Geastraceae have typical glebal structure, and in some genera of
Tulostomataceae while some genera are not plectobasidial, it seems more
reasonable to distribute such plectobasidial forms among genera with
normal hymenial cavities with which they show the closest resemblance
instead of putting them all together in one series.

The coralloid structure by emphasis on a number of main branches

658 THE PHYLOGENY OF THE FUNGI

which reached to the cortex produced the multipilar structure. By further
emphasis upon the axial branch and reduction of the lateral branches the
unipileate condition doubtless arose. These two types of structure are
present in the Phallales which doubtless have close kinship with the
simpler, multipilar Hysterangiaceae. The Tulostomataceae perhaps have
their origin near those Lycoperdaceae in which a stipe-like base shows
itself and where a columella is sometimes clearly beginning to develop.

All the foregoing brings us to Elasmomyces, with its close kinship to
Russula and Ladarius and to Secotium which has many analogies with
other Agaricaceae. This leads to the consideration of the possibility that
these higher Gasteromyceteae arose from the Agaricaceae and that the
evolution has been from these, by simplification and reduction, until
ultimately the minute, one-chambered forms may be considered to have
reached the furthest point in evolution away from the ancestral higher
Hymenomyceteae instead of representing forms that have changed only
a little from lower Hymenomyceteae. (Fig. 210.)

Holm (1949) suggests that the Gasteromyceteae may be polyphyletic,
with part of the group, e.g. Hymenogastrales, derived from the Tuberales
and part, such as some of the forms like Elasnio7nyces, etc., from the
Agaricales. He emphasizes the similarity of ontogeny of the spore fruits of
Hymenogaster with that of some species of Tuber. In the latter, according
to Greis (1938) typical clamp connections are found early in the course of
development.

Literature Cited

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MycoL, 7(5) :441-472. Figs. 1-20. 1909.
: Phylogeny and relationships in the Ascomycetes, Ann. Missouri Botan.

Garden, 2(1-2) :315-376. Figs. 1-10. 1914.
de Bary, Anton: Vergleichende Morphologic und Biologic der Pilze, Mycetezoen

und Bacterien, xvi + 588 pp. 198 figs. Leipzig, Wilhelm Engelmann, 1884.
Bessey, Ernst A.: Some problems in fungus phylogeny, Mtjcologia, 34(4) :355-

379. Figs. 1-5. 1942.
: Studies on Pilobolus: P. kleinii and P. longipes, Papers Mich. Acad. Sci.,

32:15-25. Pis. 1-3. 1946 (1948).
Borgesen, F.: Marine algae from the Canary Islands: III. Rhodophyceae, 1.

Bangiales and Nemalionales, Kgl. Danske Videnskab. Selskab Biol. Medd. VI,

6:1-97. Kobenhavn. 1927.
Brefeld, Oscar: Basidiomyceten: III. Autobasidiomyceten und die Begrundung

des natiirlichen Systems der Pilze, Untersuchungen aus dem Gesammt-

gebiete der Mykologie, Heft. 8, pp. 1-305. P/s. 1-12. Leipzig, Arthur Felix,

1889.
BucHOLTZ, F.: Zur Morphologic und Systematik der Fungi hypogaei, Ann.

MycoL, 1(2) :152-174. Pis. 4-5. 1903.
Buller, a. H. Reginald: Researches on Fungi, vol. 5, pp. i-xiii, 1-416. Figs.

1-174. London, Longmans, Green and Co., 1933. (Especially p. 154, Fig. 78.)

LITERATURE CITED 659

Couch, John N.: A new Conidiobolus with sexual reproduction, Am. J. Bot.,

26(3):119-130. Figs. 1-53. 1939.
Dangeard, p. a.: L'origine du p^rithece chez les Ascomycetes, Le Botaniste,

10:1-385. P/s. 1-91. 1907.
Dodge, B. 0.: The morphological relationships of the Florideae and the Asco-
mycetes, Bull. Torrey Botan. Club, 41(3): 157-202. Figs. 1-13. 1914.
Ellison, Bernard R.: Flagellar studies on zoospores of some members of the

Mycetozoa, Plasmodiophorales and Chytridiales, Mycologia, 37(4) :444-459.

Figs. 1-4. 1945.
Fischer, Eduard: Gastromyceteae, in A. Engler und K. Prantl: Die Na-

tiirlichen Pfianzenfamilien, Zweite Auflage, vol. 7a, pp. 1-122. Figs. 1-91.

Leipzig, Wilhelm Engelmann, 1933.
Gaumann, Ernst Albert: Vergleichende Morphologie der Pilze, pp. 1-626. Figs.

1-398. Jena, Gustav Fischer, 1926.
Greis, Hans : Die Sexualvorgange bei Tuber aestivum und Tuber brumale, Biol.

Zentralb., 58(11-12) :617-631. Pz^s. 1-3. 1938.
Heim, Roger: Les Lactario-Russules du Domaine Orientale de Madagascar.

Essai sur la classification et la phylogenie des Ast^rosporales, pp. 1-196. 8

pis. (4 colored). 59 figs. 2 phylogenetic diagrams. Paris, Laboratoire de Crypto-

gamie du Museum National d'Histoire Naturelle, 1937 (1938).
: Phylogeny and natural classification of macro-fungi, Brit. My col. Soc.

Trans., 30 :1Q1-17 8. Figs. 1-19. 1948.
Holm, Lennart: Some aspects on the origin of the Gastromycetes, Svensk

Botanisk Tidskrift, 43(1):65-71. 1949.
Jackson, H. S.: Life cycles and phylogeny in the higher fungi. Presidential

address. Trans. Roy. Soc. Can., Ser. 3, V, 38:1-32. Figs. 1-5. 1944.
von Jaczewski, a. a.: Zur Phylogenie der Pilze, Phijtopath. Z., 1(2):117-150.

1929-30.
JuEL, H. 0.: tJber Zellinhalt, Befruchtung und Sporenbildung bei Dipodascus,

Flora Oder Allgemeine Botan. Ztg., 91:47-55. Pis. 7-8. 1902.
DE Lagerheim, G.: Dipodascus albidus, eine neue geschlechtliche Hemiascee,

Jahrb. wiss. Botan., 24:549-565. Pis. 24-26. 1892.
LiNDER, David H. : Evolution of the Basidiomycetes and its relation to the ter-
minology of the basidium, Mycologia, 32(4):419-447. Figs. 1-6. 1940.
Mez, Carl: Versuch einer Stammesgeschichte des Pilzreiches, Schriften konigs-

berg. gelehrten Ges. Naturw. Klasse, 6:1-58. 1 fig. 1929.
Rogers, Donald P.: A eytological study of Tulasnella, Botan. Gaz., 94(1) :86-105.

Figs. 1-79. 1932.
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1934.
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pp. 492 ^^s. Leipzig, Wilhelm Engelmann, 1874.
Singer, Rolf: Phylogenetic theories concerning the origin of the Agaricales, a

chapter in "The Agaricales." Waltham, Mass., Chronica Botanica Co.,

Publishers, 1950. (In press.)
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pp. 2 pis. 2 figs. Paris, Imprimerie M. Flinikowski, 1920.
Sparrow Jr., Frederick K.: Aquatic Phycomycetes Exclusive of the Sapro-

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Press, 1943.
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Sci., 42:60-62, 1942; IV. Phytophthora, ibid., 43:135-138, 1943.

GUIDE TO THE LITERATURE FOR THE
IDENTIFICATION OF FUNGI

THE vast number of fungi makes a knowledge of even a considerable
part of the species beyond the ability of all but a few specialists. To
be able to identify a given specimen requires that the necessary literature
be available. Before 1880 the great Italian mycologist, P. A. Saccardo,
began to work on a compilation of all species of fungi described up to that
time. These descriptions were brought together in a compendious work
entitled "Sylloge Fungorum," the first volume of which appeared in 1882.
The descriptions, in Latin, were arranged in accordance with the system-
atic classification of fungi then recognized by the author. He completed
the work with Volume 8 in 1889, but in the meantime, such was the
stimulus afforded by this great work bringing together in one place de-
scriptions of all known species of fungi, that thousands of additional species
had been recognized and described. Thus it became necessary to pubhsh
supplementary volumes, the last of which. Volume 25, appeared in 1931.
For many years species of fungi have been described at the rate of 1500 to
2500 species a year. Thus it is inevitable that such a work must be from
two to five years behindhand. A further difficulty is the language of the
description, Latin, a knowledge of which is unfortunately all too meager
among the later generation of botanists. Furthermore the lack of illustra-
tions and the necessary scattering of the descriptions among the original
volumes and the many supplements make the work difficult to use.
Besides this, the necessary high cost precludes its purchase by most
botanists so that they must depend upon copies owned by libraries.

To obviate these difficulties local fungus floras have been issued in
various countries or even subdivisions of countries. These pubhcations
are usually in the language of the country and are furthermore smaller,
inasmuch as only the species occurring in the limited areas concerned are
included.

Still another type of publication is the monograph, or intensive study

660

GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI 661

of some smaller group of fungi, such as a family or a genus or even a sec-
tion of a genus. By virture of the limitation of effort to this relatively
small number of species it is possible for such a monograph to devote
larger space to the description of the individual species and to make their
identification easier.

In the following pages the more general works are first listed; then
under different headings are included the publications that are confined
more to special groups. These are arranged systematically in accordance
with the classification of the fungi. It is the aim of the author to list the
more recent publications of this nature from all parts of the world, but it
is certain that there are many omissions, especially for parts of the world
whose mycological literature is not so widely available in the United
States as is the literature of this country and of the larger European
countries. Even for the latter the large number of publications of some-
what limited scope from botanical or mycological societies makes a com-
plete list difficult to obtain as in no one library will even a majority of
such works be found. The disruption of communication and the destruc-
tion due to World War II and the subsequent disturbed political and
economic conditions have made it increasingly difficult to keep in touch
with the mycological work of other regions of the world.

The student is advised to turn first to the portion of the list where
these papers of monographic nature are to be found. In case no such paper
is listed for the fungus he has, he must turn to the more general lists.

Because of the various systems of classification used by different
authors the arrangements of the items in the following lists do not follow
any one system in all its details. Cross references are necessary in the
cases where one work includes groups now segregated although formerly
united.

It must be remembered that certainty of identification depends not
only upon the availability of the necessary literature but also upon the
fullness of the knowledge of the structure, development, etc., of the fungus
in question. The literature should not be consulted until the main morpho-
logic and anatomic details have been ascertained, including measurements
of spores, sporophores, etc. The fuller the knowledge of details as to the
substratum on which the fungus grows, its habitat, color, appearance
when fresh, appearance at different stages of development, etc., the more
easily will the identification be effected.

662 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

List 1. General Works Covering the Whole Field of Systematic

Mycology

Saccardo, p. a. : Sylloge fungorum omnium hucusque cognitorum, 25 vols, issued
up to 1931. Pavia, Italy, published by the author.

1:1-768. 1882. Pyrenomyceteae.

2:1-959. 1883. Pyrenomyceteae (continued).

3:1-860. 1884. Sphaeropsideae and Melanconieae.

4:1-807. 1886. Hyphomyceteae.

5:1-1146. 1887. Hymenomyceteae : I. Agaricineae.

6:1-928. 1888. Hymenomyceteae: II. Polyporeae, Hydneae, Thelephoreae,
Clavarieae, Tremellineae.

7:1-941. 1888. Gasteromyceteae, Phycomyceteae, Myxomyceteae, Ustilag-
ineae, Uredineae.

8:1-1143. 1889. Discomyceteae, Phymatosphaeriaceae, Tuberaceae, Ela-
phomycetaceae, Onygenaceae, Laboulbeniaceae, Saccharomycetaceae,
Schizomycetaceae.

9:1-1141. 1891. Supplement 1: Hymenomyceteae, Gasteromyceteae, Hypo-
dermeae (Ustilaginaceae and Uredinaceae), Phycomyceteae, Pyrenomy-
ceteae, Laboulbeniaceae.

10:1-964. 1892. Supplement 2: Discomyceteae, Onygenaceae, Tuberoideae,
Myxomyceteae, Sphaeropsideae, Melanconieae, Hyphomyceteae, Fossil
Fungi.

11:1-753. 1895. Supplement 3: All groups of Fungi. Generic index to all
volumes.

12:1-1053. 1897. Index to vols. 1-11.

13:1-1340. 1898. Host index.

14:1-1316. 1899. Supplement 4: All groups of Fungi. Sterile Mycelia.

15:1-455. 1901. Supplement 5: Synonyms.

16:1-1291. 1902. Supplement 6: All groups of Fungi. Generic index to all
volumes.

17:1-991. 1905. Supplement 7: Hymenomyceteae, Gasteromyceteae, Uredi-
naceae, Ustilaginaceae, Phycomyceteae, Pyrenomyceteae, Laboulbenio-
myceteae.

18:1-838. 1906. Supplement 8: Discomyceteae (including Saccharomyce-
taceae, Exoascaceae, Gymnoascaceae, Tuberaceae, etc.), Myxomyceteae,
Deuteromyceteae (Fungi Imperfecti). Generic index to all volumes.

19:1-1158. 1910. Index of illustrations of Fungi, A-L.

20:1-1310. 1911. Index of illustrations of Fungi, M-Z.

21:1-928. 1912. Supplement 9: Hymenomyceteae, Gasteromyceteae, Usti-
laginaceae, Uredinaceae, Phycomyceteae.

22:1-Bl2. 1913. Supplement 10: Ascomyceteae, Deuteromyceteae, Sterile
Mycelia.

23:1-1026. 1925. Supplement 10 (continued): Hymenomyceteae, Usti-
laginales, Uredinales.

24(Section I):l-703. 1926. Supplement 10 (continued): Phycomyceteae,
Laboull)eniales, Pyrenomyceteae, in part.

24(Section II):704-1438. 1928. Supplement 10 (continued): Remainder of
Pyrenomyceteae, Discomyceteae; Appendix, consisting of additions to
vols. 23 and 24.

25:1-1093. 1931. Supplement 10 (continued): Myxomyceteae, Myxobac-
teriaceae, Deuteromyceteae, Mycelia Sterilia.

LIST 1. GENERAL WORKS 6G3

(Volumes 1, 10, and 17 contain bibliographies; Vol. 14 contains an explana-
tion of the arrangement of the genera by the spore form and color
scheme.)
Engler, a., und K. Prantl: Die nattirlichen Pflanzenfamilien, Leipzig, Wilhelm
Engelmann. The parts devoted to fungi (including the Lichens) are the
following:
Teil I, Abteilung 1 :1-513. Figs. 1-293. 1897. Myxomyceteae, Phycomyceteae,

Ascomyceteae.
Teil I, Abteilung l*:l-249. Figs. 1-125. 1907. Lichens.
Teil I, Abteilung l**:l-570. Figs. 1-263. 1900. Basidiomyceteae, including

Hemibasidii (Ustilaginales) and Uredinales. Fungi Imperfecta
(This work will enable one to determine the genus of almost any fungus but
not the species. It is very helpful because of the illustrations.)

, UND : Die nattirlichen Pflanzenfamilien, Zweite Auflage. Leipzig,

Wilhelm Engelmann.
2:304^339. Figs. 425-447. 1928. Mycetozoa.
5b:l-42. 22^1^5. 1938. Tuberineae.
6:1-290. Pis. 1-5. Figs. 1-157. 1928. Ustilaginales, Uredinales, Hymenomy-

ceteae.
7a:l-122. Figs. 1-91. 1933. Gastromyceteae.
8:1-270. Figs. 1-127. 1926. Lichens.
Clements, Frederick E., and Cornelius L. Shear: The Genera of Fungi,

iv + 496 pp. 58 pis. New York, H. W. Wilson Company, 1931.
Martin, G. W.: Outline of the fungi, Univ. Iowa Studies in Natural History,

18(supplement):l-64. Fi^s. 1-118. 1941.
R.VBENHORST, L. : Kryptogameu-Flora von Deutschland, Oesterreich und der
Schweiz, Zweite Auflage. Leipzig, Verlag von Eduard Kummer.
Band 1. Winter, Georg: Die Pilze. This "Volume" on Fungi is so extensive
that it is issued as ten separately bound "Abteilungen," as follows:
1:1-924. 1 pi. and numerous text figs. 1884. Schizomyceten, Saccharo-
myceten, und Basidiomyceten. By A. de Bary, H. Rehm, and Georg
Winter.
2:1-928 and Index, 1-112. Numerous text figs. 1887. Ascomyceten: Gym-
noasceen und Pyrenomyceten. By A. de Bary, H. Rehm, and Geo'rg
Winter.
3:1-1275 and Index, 115-169. Numerous text figs. 1896. Ascomyceten:

Hysteriaceen und Discomyceten. By H. Rehm.
4:1-505. Figs. 1-74. 1892. Phycomyceten. By Alfred Fischer.
5:1-131. Numerous text figs. 1897. Ascomyceten: Tuberaceen und Hemias-

ceen. By Eduard Fischer.
6:1-1016. Numerous text figs. 1901. Fungi Imperfecti: Hyalin-sporige

Sphaerioideen. By Andreas Allescher.
7:1-993 and Index to Abteilungen 6 and 7, pp. 995-1072. Numerous figs.
1903. Fungi Imperfecti: Gefarbt-sporige Sphaerioideen, sowie Nectri-
oideen, Leptostromaceen, Excipulaceen und Melanconieen. By Andreas
Allescher.
8:1-852. Numerous text figs. 1907. Fungi Imperfecti: Hyphomyceten :
Mucedinaceen und Dematiaceen (Phaeosporae und Phaeodidymae).
By G. Lindau.
9:1-983. Numerous text figs. 1910. Fungi Imperfecti: Dematiaceen (Phaeo-
phragmiae bis Phaeostaurosporae) , Stilbaceen und Tuberculariaceen.
By G. Lindau.

664 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

10:1-474. 182 ^grs. 1920. Myxogasteres (Myxomycetes, Mycetozoa). By
Hans Schinz.
Band 8. Keissler, Karl von: Die Flechtenparasiten, xi + 712 pp. ISbfigs.

1930.
Band 9. Zahlbruckner, Alexander: Die Flechten. This is divided into
Abteilungen, Teile und Lieferungen of which, so far as the records are
available, the following have appeared to date:

Abteilung I, Teil 1. von Keissler, Karl: Moriolaceae, pp. 1-43. Figs.
1-15. 1933. Zschacke, Hermann: Epigloeaceae, Verrucariaceae und
Dermatocarpaceae, pp. 44-695. Figs. 16-344. 1934. Teil 2. von Keiss-
ler, Karl: Pyrenulaceae, Trypetheliaceae, Pyrenidiaceae, Xantho-
pyreniaceae, Mycoporaceae und Coniocarpineae, pp. 1-846. Figs. 1-210.
1937-1938.

Abteilung II, Teil 1. Redinger, K.: Arthoniaceae, Graphidaceae, Chio-
dectonaceae, Dirinaceae, Rocellaceae, Lecanatidiaceae, Thelotremaceae,
Diploschistaceae, Gyalectaceae, Coenogoniaceae, pp. 1-404. 2 pis. Figs.
1-107. 1937-1938.

Abteilung III. Magnusson, A. K.: Lecideaceae. (The numbers of Liefer-
ungen and pages and dates were not available.)

Abteilung IV, Erste Halfte. Frey, Eduard: Cladoniaceae (unter Aus-
schluss der Gattung Cladonia), Umbilicariaceae, pp. 1-426. Pis. 1-8.
Figs. 1-64. 1933. Zweite Halfte. Sanstede, Heinrich: Die Gattung
Cladonia, pp. 1-531. Pis. 1-34. Figs. 1-8. 1931.

Abteilung V, Teil 1. Magnusson, A. K. : Acarosporaceae und Thelo-
carpaceae, pp. 1-320. Figs. 1-64. 1935. Ericksen, C. F. E.: Pertusari-
aceae, pp. 321-728. Figs. 1-74. 1935-36. Teil 3, Lieferungen 1-2. Hill-
mann, Johannes: Parmeliaceae, pp. 1-309. Register 1-10. Figs. 1-16.
1936.

Abteilung VI, Lieferung 1. Hillmann, Johannes: Teloschistaceae, pp.

1-36. Figs. 1-4. 1935. Lynge, Bernt: Physciaceae, pp. 37-188. Pis.

1-12. Figs. 1-10. 1935.

Kryptogamenflora der Mark Brandenburg, Leipzig, Gebrlider Borntraeger. This

work appears in nine or more volumes of which the following concern the

fungi:

5. 630 pp. 151 figs. 1915. Schizomycetes, by R. Kolkwitz; Myxobacteriales,
by E. Jahn; Chytridiineae, Ancylistineae, Monoblepharidineae, Sapro-
legniineae, by M. von Minden.

5a. 946 pp. 380 figs. 1914. Uredineen, by H. Klebahn; Ustilagineen, Auricu-
lariineen, Tremellineen, by G. Lindau.

6. Not complete. Issued so far only:

Heft 1:1-92. Many illustrations. 1910. By W. Herter. Autobasidio-
mycetes: Dacryomycetaceae, Exobasidiaceae, Tulasnellaceae, Corti-
ciaceae, Thelephoraceae, Cyphellaceae, Craterellaceae, Clavariaceae,
Sparassiaceae, Hydnaceae (incomplete).
6a. 264 pp. IIA figs. 1935. Mucorineae. By H. Zycha.

7. Not complete. Issued ^o far only:

Heft 1 :1-160. Many illustrations. 1905. Hemiasci, by G. Lindau; Saccharo-
mycetineae, by P. Lindner; Protoascuneae, by G. Lindau; Exoascaceae,
by F. Neger; Ascocorticiaceae and Gymnoascaceae, by G. Lindau;
Aspergillaceae, by F. Neger; Onygenaceae, Elaphomycetaceae and
Terfeziaceae, by P. Hennings; Erysiphaceae and Perisporiaceae, by
F. Neger; Tuberaceae (incomplete) by P. Hennings.

Heft 2:161-304. Many illustrations. 1911. Tuberaceae (completed), by

LIST 1. GENERAL WORKS 665

P. Hennings; various families of the Sphaeriales, by W. Kirch-
stein.
Heft 3:305-448, 1938. Ascomycetes (continued), by W. Kirchstein.

8. No parts yet issued.

9. 962 pp. SS9figs. 1915. Sphaeropsideen, Melanconien, by H. Diedicke.
Oudemans, C. a. J. A. : Revision des Champignons tant sup^rieurs qu'inf^rieurs

jusqu'a ce jour trouves dans les Pays-Bas.

1 : 1-638. Hymenomycetes, Gasteromycetes, Hypodermeae. Amsterdam, J.

Miiller, 1893.
2: 1-491. Pis. 1-14. Phycomycetes and Pyrenomycetes. Separate reprint from
Verh. Kon. Akad. Wet. AmMerdam 2. Ser II. 1897.
Schroeter, J.: Die Pilze Schlesiens, in Ferdinand Cohn: Kryptogamen-Flora
von Schlesien, vol. 3. Breslau, J. V. Kern's Verlag.

Erste Halfte: pp. 1-814, 1889. Myxomycetes, Schizomycetes, Chytridiei,
Zygomycetes, Oomycetes, Protomycetes, Ustilaginei, Uredinei, Auricu-
lariei, Basidiomycetes. Host index for this half volume.
Zweite Halfte: pp. 1-597. 1908. Ascomycetes and a small part of the Fungi
Imperfecta Host index for second half volume.
MiGULA, W.: Krytogamen-Flora von Deutschland, Deutsch-Osterreich und der
Schweiz, in D. W. Thom^: Flora von Deutschland etc., Zweite Auflage. Gera,
Friedrich von Zezschwitz.
Band VTeil 1. iv + 510 pp. 92 pis. 1910. Myxomycetes, Phycomycetes, Ba-
sidiomycetes (Ordnungen Ustilagineae und Uredineae).
Teil 2. iv + 814 pp. 304 pis. 1912. Basidiomycetes (completed).
Teil 3, Abteilung 1. iv + 1-684. Pis. 1-100. 1913. Hemiasci, Saccharo-
mycetineae, Protodiscineae, Plectascineae, Pyrenomycetes (Perisporiales
und Sphaeriales).
Teil 3, Abteilung 2. iv + 685-1404. Pis. 101-200. 1913. Dothideales, Hy-

pocreales, Hysteriales, Discomycetes, Laboulbeniaceae.
Teil 4, Abteilung 1. iv + 614 pp. 90 pis. 1921. Fungi Imperfecta
Band 4, Teil 1. viii + 527 pp. 82 pis. 1929. Flechten.
Teil 2. iv + 868 pp. 143 pis. 1931. Flechten (conclusion).
Jaczewski, a. a.: Identification of Fungi, vol. 1, 1913; vol. 2, 1917. (In Russian.)

, AND P. A. Jaczewski: Identification of Fungi. Perfect Forms (Diploid

Stages), Tom I. Phycomycetes, ed. 3, 294 pp. 329 figs. Leningrad and Mos-
cow, 1931. (In Russian.)
Cooke, M. C: Handbook of British Fungi with Full Descriptions of all the
Species and Illustrations of the Genera, 2 vols, ii + 981 pp. 7 pis. 408 figs.
London, Macmillan and Co., 1871; ed. 2, 1883.

: Handbook of Austrahan Fungi, xxxii + 457 pp. 36 pis. London, Williams

and Norgate, 1892.
Massee, George: British Fungus Flora, a Classified Text-book of Mycology,
London, George Bell and Sons.
Vol. 1. xii + 432 pp. Illustrated. 1892. Gastromycetes ; Tremellineae; Cla-
varieae; Thelephoreae ; Hydneae; Polyporeae; Agaricineae: Melanosporeae
and Porphyrosporeae.
Vol. 2. vii + 460 pp. Illustrated. 1893. Agaricineae: Ochrosporeae, Rhodo-

sporeae, Leucosporeae.
Vol. 3. viii + 512. Illustrated. 1893. Agaricineae: Leucosporeae; Hyphomycetes.
Vol. 4. viii + 522 pp. Illustrated. 1895. Ascomycetes.

: British Fungi with a Chapter on Lichens, 551 pp. Colored pis. 1-40. Pis.

A-B. 19 unnumbered figs. London, George Routledge and Sons. Undated
(about 1911).

666 GUIDE TO THE LITERATUEE FOR THE IDENTIFICATION OF FUNGI

Massee, George, and Ivy Massee: Mildew, Rusts and Smuts: A Synopsis of
the Families Peronosporaceae, Erysiphaceae, Uredinaceae and Ustilaginaceae,
229 pp. Pis. 1-5. London, Dulau and Co., 1913.
Stevens, F. L.: The Fungi which Cause Plant Disease, ix + 754 pp. 449 figs.
New York, Macmillan, 1913.

: Hawaiian Fungi, Bernice P. Bishop Museum Bulletin 19 :i-ii, 1-189.

Pis. 1-10. Figs. 1-35. Honolulu, 1925.
North American Flora, published by the New York Botanical Garden, New York.
Of this work various parts describing fungi have been issued, as follows:
Vol. 1, Pt. 1. 1949. Mycetozoa.

Vol. 2, Pt. 1. 1937. Blastocladiales, Monoblepharidales and Saprolegniales.
Vol. 3, Pt. 1. 1910. Hypocreales and Fimetariales.
Vol. 6, Pt. 1. 1922. Phyllostictales (part).
Vol. 7, Pts. 1-13. 1906-1931. Ustilaginales and Uredinales.
Vol. 9, 1-542. 1907-1916. Polyporaceae (part), Boletaceae, Agaricaceae (part).
Vol. 10, Pts. 1-5. 1917-1932. Agaricaceae (continued).
Larsen, p.: Fungi of Iceland, vol. 2, pt. 3, in L. Kolderup Rosenvinge and E.
Warming: The Botany of Iceland. Copenhagen and London, Oxford Univ.
Press, 1932.
Heim, Roger, (Ed.) : Flore mycologique de Madagascar et D^pendances, Paris,
Laboratoire de Cryptogamie du Museum National d'Histoire Naturelle,

I. Heim, Roger: Les Lactario-Russul^s. 196 pp. 8 pis. QO figs. 1938.

II. Romagnesi, H. : Les Rhodophylles. 146 pp. 4:Q figs. 1941.

III. Method, Georges: Les Mycenes. 144 pp. 88 ^grs. 1949.

IV. LeGal, Marcelle: Les Discomycetes Opercul^s. (To appear 1950.)

V. Bucket, Samuel: Les Myxomycetes. (To appear 1951.)

VI. Heim, Roger, et Raymond Decary: Les Phalloid^es. (In preparation.)

VII. Bouriquet, Gilbert: Les Rouilles. (In preparation.)

Flora Italica Cryptogama. Firenze, published under the auspices of the Societa

Botanica Italiana. Pars I. Fungi.

Traverso, G. B.: Elenco bibliografico della mycologia Italiana, Fasc.
1:1-118, Supplemento 1:119-135. 1905. Supplemento II, Fasc. 9:1-151.
1912.

: Pyrenomyceteae : Xylariaceae, Valsaceae, Ceratostomataceae, Fasc.

2:1-352. Figs. 1-68. 1906; Sphaeriaceae allantosporae, hyalosporae, phaeo-
sporae. Fasc. 3:353-492. Figs. 69-97. 1907; Sphaeriaceae hyalodidymae,
Fasc. 11:493-700. Figs. 98-116. 1913.

Trotter, Alex: Uredinales, Fasc. 4:1-519. Figs. 1-110. 1908.

Petri, L.: Gasterales, Fasc. 5:1-139. Figs. 1-83. 1909.

Ferraris, T. : Hyphales: Tuberculariaceae, Stilbaceae, Fasc. 6:1-198. Figs.
1-53. 1910; Dematiaceae, Fasc. 8:199-534. 1912; Mucedinaceae, Fasc.
10:535-846. 1913; Indice generale, Fasc. 13:847-979. 1914.

Saccardo, p. a., adiuvante Hier. Dalla Corda: Hymeniales: Leuco-
sporae et Rhodosporae, Fasc. 14:1-576. Pis. 1-6. Figs. 1-7. 1915; Hy-
meniales : ceterae Agaricaceae, Polyporaceae, Hydnaceae, Thelephoraceae,
Tremellaceae, Fasc. 15:577-1386. P/s. 7-11. 1916.

CoLLA, S.: Laboulbenialcs, Fasc. 16:1-157. Figs. 1-108. 1934.

Ciferri, Raphael: Ustilaginales, Fasc. 17:1-443. Figs. 1-23. 1938.

(Other fascicles have probably appeared but their citations are not available.)

Pars. III. Lichenes.

Jatta, a.: Lichenes, Fasc. 1: i-xxii, 1-958. Figs. 1-80. 1909.

LIST 1. GENERAL WORKS 667

Barghoorn, E. S., and D. H. Linder: Marine fungi: their taxonomy and biology,

Farlowia, l(3):395-467. Pis. 1-7. Figs. 1-3. 1944.
Van Overeem, C, und J. Weese: Icones fungorum Malayensium Abbildungen

und Beschreibungen der Malayischen Pilze, Hefte 1-16. 16 colored pis. 1 pi.

in black and white. Weesp, Holland, Mycol, Museum, 1923-1926.
CoRDA, A. C. I.: Icones fungorum hucusque cognitorum, Prag, J. G. Calve (vols.

1-4), Fr. Ehrlich (vols. 5-6).

1:1-32. Pis. 1-7. 1837. Mostly Fungi Imperfecti; a few Uredinales, My-
cetozoa and miscellaneous fungi.

2:1-43. Pis. 8-15. 1838. Fungi Imperfecti, Mucorales, Mycetozoa, a few
Uredinales, Ustilaginales, Pezizales and miscellaneous fungi.

3:1-55. Pis. 1-9. 1839. Uredinales, Fungi Imperfecti, Agaricales, Miscel-
laneous.

4:1-53. Pis. 1-10. 1840. Erineum galls, Fungi Imperfecti, Uredinales, My-
cetozoa, Agaricales and various Ascomyceteae.

5:1-92. Pis. 1-10. 1842. Erineum galls. Fungi Imperfecti, Mucorales, My-
cetozoa, Gasteromycetes, Tuberales, Agaricales and Miscellaneous.

6:i-xix, 1-91. Pis. 1-20. 1854. (Plates by A. C. I. Corda, text by J. B.
Zobel). Uredinales, Fungi Imperfecti, Mucorales, Mycetozoa, Gastero-
mycetes, Tuberales, and Miscellaneous.
Juillard-Hartmann, G.: Iconographie des champignons sup^rieurs, 5 vols. 250

colored pis. Epinal, Juillard et Fils. vol. 1, 1919, others not dated. Illustrations

in color of approximately 2400 species of fungi. Vols. 1-3 and part of vol. 4

represent Agaricaceae, the remainder of vol. 4 illustrates Polyporaceae,

Boletaceae, Fistulinaceae; vol. 5 illustrates Hydnaceae, Clavariaceae, Thele-

phoraceae, Exobasidiaceae, Gastromyceteae, Dacryomycetales, Tremellales,

Auriculariales, Helvellaceae, and a few subterranean Ascomyceteae. No

descriptions accompany the plates.
KoNRAD, P., ET A. Maublanc: Icones selectae fungorum. 5 vols, of plates totaling

500; 1 vol. text, over 500 pp. Paris, Paul Lechevalier, 1924-1937.
Bresadola, J. : Fungi Tridentini novi vel nondum delineati, descripti et iconibus

illustrati, Trieste. Published by author.

1:1-114. P/s. 1-105. 1881.

2:118 pp. Pis. 106-217. 1892.
: Iconographia mycologica, edited by J. Traverso, L. Fenaroli,^ G,

Catoni, and J. B. Traverso. 24 vols. 1200 pis. Milan, Societa Bot. Italica,

Seg. Lombard. 1927-1932. Vols. 1-18, Agaricaceae; vols. 19-21, Polyporaceae

and part of Hydnaceae; vol. 22, remainder of Hydnaceae, Thelephoraceae,

part of Clavariaceae; vol. 23, Clavaria, Auriculariaceae, Tremellaceae,

Dacryomycetaceae, Gastromyceteae; vol. 24, Helvellaceae, Leotiaceae,

Pezizaceae.
CoupiN, Henri: Album general des Cryptogames. Fungi (Champignons).

Les Champignons du Globe, 5 vols. 473 pis. Paris, E. Orlhac, about 1920-

1925. (Gives illustrations of nearly all the recognized genera of fungi except

the lichens.)
ScHWARZE, Carl A.: The parasitic fungi of New Jersey, New Jersey Agr. Expt.

Sta. Bull. 313:1-226. Figs. 1-1056. 1917. (Contains beautiful illustrations of

very many genera and species of parasitic fungi.)

G68 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

List 2. Host Indexes, Local Fungus Lists with Host Indexes,

Lists of Fungi on Special Hosts or Substrata,

Bibliographies, Fungi of Man and Other Animals

Saccardo, p. a. Sylloge fungorum omniuin hucusque cognitorum, 13:1-1340.

1898. Also there is a generic host index (Repertorium) at the close of vols.

14, 16, 17, 18, 21, 25, covering the fungi included in the respective volumes.
OuDEMANS, C. A. J. A.: Enumeratio systematica fungorum. The Hague, Martin

Nijhoff.

1. cxxvi + 1230 pp. 1919. Host Index of Algae, Fungi, Bryophyta, Pterido-
phyta, Gymnosperms and Monocotyledons.

2. xix + 1069 pp. 1920. Dicotyledons: Salicaceae — Basellaceae.

3. xvi + 1313 pp. 1921. Dicotyledons: Caryophyllaceae — Vitaceae.

4. xiii + 1231 pp. 1923. Dicotyledons: Elaeocarpaceae — Compositae. Sup-
plement.

5. vii + 999 pp. 1924. Index to species of hosts and fungi in vols. 1-4. (This
is a host index of all parasitic fungi reported in any part of the world on
plants native to Europe or introduced into Europe.)

Seymour, Arthur Bliss: Host Index of the Fungi of North America, xiii + 732

pp. Cambridge, Harvard Univ. Press, 1929.
Anderson, Paul J.; Royal J. Haskell; Walter C. Muenscher; Clara J.

Weld; Jessie I. Wood; and G. Hamilton Martin: Check list of diseases of

economic plants in the United States, U.S. Dept. Agr. Dept. Bull. 1366:1-

111. Figs. 1-4. 1926.
Weiss, Freeman: Check list revision. Plant Disease Reptr., 24-33, various num-
bers, 1940-1949. (A revision of the foregoing.)
Alstatt, G. E.: Diseases of plants reported in Texas since 1933, Pla7it Disease

Reptr., supplement 135:37-50. 1946.
Waterston, J. M.: The fungi of Bermuda, Dept. Agr. Bermuda, Bulletin 23:

i-iii, 1-305. Figs. 1-38. 1947.
BiSBY, G. R., with the collaboration of A. H. R. Buller, John Dearness, W. P.

Eraser, R. C. Russell, and with a preface by H. T. Gtissow: The Fungi of

Manitoba and Saskatchewan, 189 pp. 1 map. 49 figs. Ottawa, National

Research Council of Canada, 1938. (Lists over 2700 species of fungi, with

host index.)
Butler, E. J., and G. R. Bisby: Fungi of India, Imperial Council of Agricultural

Research of India. Science Monograph l:i-xviii, 1-237. 1931.
MuNDKUR, B. B., and M. J. Thirumalachar: Revisions of and additions to

Indian Fungi, I, Mycological Papers. Commonwealth Mycological Inst.,

16:1-27. 19 figs. 1946.
, and Sultan Ahmad: Revisions of and additions to Indian Fungi, II,

ibid., 18:1-11. 8 figs. 1946.
Brown, Charles C: Contributions toward a host index to plant diseases in

Oklahoma, Oklahoma Agr. Expt. Sfa. Circ. 33. 1939. Revised edition 1941.

— : Supplement No. 1. ibid., Mimeographed Circular M. 104:1-32. 1943.

Preston, D. A.: Host Index of Oklahoma plant diseases, supplement, 1948,

Plant Disease Reptr., 32(9):398-401. Sept. 15, 1948.
Melchers, L. E.: a check list of plant diseases and fungi occurring in Egypt,

Trans. Kansas Acad. Sci., 34:41-106. 1931.
CooKE, Wm. Bridge: Preliminary host index to fungi of Mt. Shasta, CaUfornia,

Pla?it Disease Reptr., supplement 123:125-133. 1940.

LIST 2. HOST INDEXES, ETC. 669

Coons, G. H. : A preliminary host index of the fungi of Michigan, exclusive of the

Basidiomycetes, and of the plant diseases of bacterial and physiological

origin, Mich. Acad. Sci. Rept., 14:232-276. 1912.
Davis, J. J. : A provisional list of the parasitic fungi of Wisconsin, Trans. Wis-
consin Acad. Sci., 17(2) :846-984. 1914.
: Parasitic Fungi of Wisconsin, 157 pp., Madison, Wis., published for the

author posthumously. 1942.
Parris, G. K.: a check list of fungi, bacteria, nematodes and viruses occurring

in Hawaii, and their hosts, P/an^i Disease ^ep/r., supplement 121:1-91. 1940.
Noble, R. J.; H. J. Hynes; F. C. McCleery; and W. A. Birmingham: Plant

diseases recorded in New South Wales, Dept. Agr. of New South Wales Sci.

Bull. 46:1-47, 1934; Supplement to the foregoing, ibid., supplement 1:1-7,

1937.
McAlpine, D.: Systematic arrangement of Australian fungi together with host-
index and list of works on the subject. Department of Agriculture, Victoria,

vii + 236 pp. Melbourne, Government Printer, 1895.
ScHADE, Arthur L.: A preliminary list of the parasitic fungi of Idaho, Plant

Disease Reptr., supplement 95:77-113. 1936.
Stevenson, John A. : A check list of Porto Rican fungi and a host index, /. Dept.

Agr. Porto Rico, 2:125-264. 1918.
Kawamura, S.: The Japanese Fungi, ed. 3, Tokyo, 1930. (In Japanese.)
Maneval, Willis E.: A list of Missouri fungi with special reference to plant

pathogens and wood destroying species, Univ. Missouri Studies, 12(3):1-150.

1937. (A list of the reported species of fungi from Missouri and a host

index.)
GiLMAN, Joseph C: First supplementary list of parasitic fungi from Iowa, Iowa

State Coll. J. Sci., 6:357-365. 1931.
• — : Second supplementary list of parasitic fungi from Iowa, ibid., 23(3) :261-

272. 1949.
, AND W. Andrew Archer: The fungi of Iowa parasitic on plants, ibid.,

3(4) :299-507. 2^^s. 1929.
Sprague, Roderick: A revised check list of the parasitic fungi on cereals and

other grasses in Oregon, Plant Disease Reptr., supplement 134:1-36. 1942.
Boyce, J. S. : Host relationships and distribution of Conifer rusts in the United

States and Canada, Trans. Connecticut Acad. Arts Sci., 35 :329-482. 1943.
Thirumalachar, M. J.: Some fungal diseases of Bryophytes in Mysore, Brit.

Mycol. Soc. Trans., 31(1-2) :7-12. Figs. 1-8. 1947.
Chardon, Carlos E., and Rafael A. Toro: Mycological exploration of Vene-
zuela, Monographs of the Univ. Puerto Rico, Physical and Biological Sciences,

B, 2:1-353. Pis. 1-33 (1 colored). 1 map. 1934. (Keys to various families.

Host index.)
Moller, F. H.: Fungi of the Faeroes: I. Basidiomycetes, 295 pp. Map. 3 pis.

{colored). \Mfigs. Copenhagen, Ejnar Munksgaard Forlag, 1945.
Darker, G. D.: A brief host index of some plant pathogens and virus diseases in

Eastern Asia, Plant Disease Reptr., supplement 122:93-123. 1940.
Garcia Rada, German, y J. A. Stevenson: La flora fungosa Peruana. Lista

preliminar de hongos que atacan a las plantas en el Peru, 112 pp. Lima,

Estacion Experimental Agricola de la Molina, 1942. (List of Peruvian[]fungi

and host index.)
OsTERO, Jos^ I., AND Melville T. Cook: a bibliography of mycology and

phytopathology of Central and South America, Mexico and the West Indies,

/. Agr. Univ. Puerto Rico, 2(3):249-486. 1937.

670 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Reinking, Otto A.: Higher Basidiomycetes from the Philippines and their hosts,

I, Philippine J. Sci., 15(5) :479-490. 1919; II, ibid., 16(2):167-179. 1920; III,

ibid., 16(5) :527-537. 1920.
: Host index of diseases of economic plants in the Philippines, Philippine

4^r., 8(1-2) :38-54. 1919.
Teodoro, Nicanor G.: An enumeration of Philippine fungi. Commonwealth of

the Philippines, Department of Agriculture and Commerce, Technical Bull. 4 :

1-585. 1937. (Contains a host index.)
Raabe, Achilles: Parasitische Pilze der Umgebung von Tubingen. Ein Beitrag

zur Kryptogamenflora Siidwestdeutschlands, Hedwigia, 78(1-2) :1-106. PL

1. Figs. 1-7. 1938.
Savulescu, Trian, et C. Sandu-Ville: Contribution a la connaissance des

Micromycetes de Roumanie, Bull, trimestr. soc. mycol. France, 46(3-4) :177-

192. 1930. (See the three further contributions following below.)
■ — , UND : Beitrage zur Kenntnis der Micromyceten Rumaniens, Hed-
wigia, 73:71-132. 1933.
, UND : Beitrage zur Kenntnis der Micromyceten Rumaniens, ibid.,

75:159-233. 1935.
, et : Quatrieme contribution h la connaissance des Micromycetes

de Roumanie, Acad. Romano, Mem. Sect. Stiintifice {Bucharest), 15:397-502.

15 pis. 1939-40 (1941).
, UND : Die Erysiphaceen Rumaniens, Annates Scientifiques de

V Academic de Hautes Etudes Agronomiques de Bucarest, 1 :47-123. Pis. 1-24.

1929. (Keys and host index.)
: Contributions a la connaissance des Ustilagin^es de Roumanie, Ann.

inst. recherches agron. Roumanie, 7:1-86. Pis. 1-35. 1935. (Distribution list

and host index.)

ET T. Rayss: Contribution a I'^tude de la Mycofiore de Palestine, Ann.

cryptogam, exotique, 8(1-2) :49-87. 1935.

SiEMASZKO, Wincenty: Badania mykologiczne w gorach Kaukazu (Recherches
mycologiques dans les montagnes du Caucase), Archiwum Nauk Biologi-
cznych Towarzystwa Naukowego Warszawskiego (Disciplinarum Biologicarum
Archivum Societatis Scientiarum Varsaviensis), l(14):l-57. Fig. 1. 1923.

Bremer, Hans; H. Ismen; G. Karel; H. Ozkan; und M. Ozkan: Beitrage zur
Kenntnis der parasitischen Pilze der Tlirkei, III, Istanbul Universitesi Fen
Fakultesi Mecmuasi {Rev. Fac. Sci. Univ. Istanbul), Ser. B., Sci. Nat.,
13(l):l-53. 1948.

Cummins, George B.: Annotated check list and host index of the rusts of Guate-
mala, Plant Disease Reptr., supplement 142:79-131. 1943.

Faull, Joseph Horace: Tropical fern hosts of rust fungi, J. Arnold Arboretum
Harvard Univ., 28(3) :309-319. 1947.

Lieneman, Catherine : Host index of the North American species of the genus
Cercospora, Ann. Missouri Botan. Garden, 16(l):l-52. 1929.

Makju, Nazeer Ahmed: Contribution to our knowledge of Indian coprophilous
fungi, J. Indian Botan. Soc, 12(2):153-164. Pis. 1-2. 1933.

GiNAi, Mohammed Asgher: Further contributions to our knowledge of Indian
coprophilous fungi, ibid., 15(5):269-284. Pis. 20-22. 1936.

Niethammer, a.: Die mikroskopischen Boden-Pilze, 1-193. Pis. 1-6. Figs. 1-57.
The Hague, W. Junk, 1937.

Gilman, J. C: a manual of soil fungi, 1-392. Figs. 1-135. Ames, Iowa State
College Press, 1945.

LIST 2. HOST INDEXES, ETC. 671

Jensen, C. N.: Fungous flora of the soil, Cornell Univ. Agr. Expt. Sta. Bull. 315 :

415-501. Illustrated. 1912.
Sabet, Younis S. : On some fungi isolated from soil in Egypt, Fouad I Univ. Bull.

Fac. Sci., 19:61-112. Figs. 1-45. 1939.
BisBY, G. R.: An Introduction to the Taxonomy and Nomenclature of Fungi, 117

pp. Kew, England, Imperial Mycological Institute, 1945.
Lindau, G., et p. Sydow: Thesaurus litteraturae mycologicae et lichenologicae,

Leipzig, Gebriider Borntraeger.

1:1-903. 1908. Authors A to L, up to 1906, incl.

2:1-808. 1909. Authors M to Z, up to 1906, incl.

3:1-766. 1913. Corrections and additions up to 1910, incl.

4:1-609. 1915. Subject lists. Applied mycology, geographical distribution,
pathology.

5:1-526. 1917. Subject lists (continued). Systematically arranged.

(A nearly complete bibliography of all mycological literature up to the close

of 1910, arranged alphabetically by the authors. Vols. 4 and 5 are arranged

by subjects, the plants diseased under their hosts.)
GuBA, E. F., AND P. A. Young: Check list of important references dealing with

the taxonomy of fungi, Trans. Am. Microscop. Soc, 43:17-67. 1924.
Chables, Vera K. : A preliminary check list of the entomogenous fungi of North

America, Insect Pest Surveij Bull. 21:707-785. 1941. (Supplement to No. 9.)
AiNswoRTH, G. C, and G. R. Bisby: A Dictionary of the Fungi, ed. 2, viii + 431

pp. 138 figs. Kew, Surrey, The Imperial Mycological Institute, 1945.
Thaxter, Roland: On certain peculiar fungus-parasites of living insects, Botan.

Gaz., 58(3) :235-253. Pis. 16-19. 1914. (See also Lists 11, 12, and 14, under

Entomophthorales, Zoopagales, Eccrinales, and Laboulbeniales.)
Fetch, T. : A list of the entomogenous fungi of Great Britain, Brit. Mycol. Soc.

Trans. 17(3):170-178. 1932. (A list with hosts and localities of all ento-
mogenous fungi known to occur in Great Britain, with the exception of the

Laboulbeniales.)

: A rei-ised list of British entomogenous fungi, ibid., 31(3-4) :286-304. 1948.

: Studies in entomogenous fungi, ibid., 7-12, about 1922-1927. (Scattered

papers.)

Notes on entomogenous fungi, ibid., 16-27, 1931-1944. (Scattered

papers.)

Watson, W. : List of British fungi parasitic on lichens or which have been included
as lichens (or vice versa), with some notes on their characters and distinc-
tions, Brit. Mycol. Soc. Trans., 31(3-4) :305-339. 1948.

Dodge, Carroll W.: Medical Mycology. Fungous Diseases of Men and Other
Mammals, 900 pp. 142 ^i^s. St. Louis, C. V. Mosby Co., 1935.

Ota, Masao, et Maurice Langeron: Nouvelle classification des Dermato-
phytes, Ann. parasitol., l(4):305-336. Figs. 1-8. 1923.

Sartory, a.: Champignons parasites de I'homme et des animaux, 895 + 47 pp.
50 pis. 91 figs. Paris, Lefrangois, 1920.

: Champignons parasites de I'homme et des animaux, ler supplement, pp.

1-78. Pis. 1-2. Figs. 1-11. Paris, Lefrangois, 1923.

, et J. Bailly: Champignons parasites de I'homme et des animaux, 2me

supplement, pp. 1-95. Paris, Editions Clinique et Laboratoire, 1927.

; A. GoDEAu; R. Sartory; L. Bailly; et J. Meyer: Champignons para-
sites de I'homme et des animaux, 3me supplement, pp. 1-159. Paris, Editions
Clinique et Laboratoire, 1933.

G72 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

VuiLLEMiN, Paul: Les champignons parasites et les mycoses de I'homme, En-
cyclop^die Mycologique, vol. 2, pp. 1-290. Figs. 1-140. Paris, Paul Lechevalier

et Fils, 1931.
CoNANT, N. F.; D. S. Martin; D. T. Smith; R. D. Baker; and J. L. Callaway:
Manual of Clinical Mycology, 348 pp. Illustrated. Philadelphia, W. B.
Saunders Co., 1945.

List 3. Mycetozoa, Including Myxogastrales,
Plasmodiophorales, Acrasiales, Labyrinthulales

Lister, Arthur: A monograph of the Mycetozoa, ed. 3, revised by Gulielma
Lister, xxxii + 296 pp. 222 -ph. 60 figs. London, Trustees of the British
Museum, 1925.
MacBride, Thomas H., and G. W. Martin: The Myxomycetes, a descriptive
list of the known species with special reference to those occurring in North
America, 339 pp. 20 pis. New York, Macmillan Co., 1934.
Hagelstein, Robert: The Mycetozoa of North America, based upon the speci-
mens in the New York Botanical Garden, pp. 1-306. Pis. 1-16. Mineola,
N. Y., published by the author, 1944.
Martin, G. W.: Fungi, Myxomycetes. Ceratiomyxales, Liceales, Trichiales,
Stemonitales, Physarales, North American Flora, 1(1):1-151, with Bibli-
ography, pp. 153-178, by Harold "W. Rickett and Index, pp. 179-190, by
Gussie M. Miller. 1949.
Torrend, C: Flore des Myxomycetes, 271 pp. 9 p/s. 1908. (Reprinted from

Broteria.)

Jaczewski, a. a.: Mycological Flora of European and Asiatic Russia: IL

Myxomycetes, Materialien zur Kenntnis der Fauna und Flora des Rus-

sichen Reiches, Botanischer Teil. Heft 6:1-140. 84 ^^s. 1907. (In Russian.)

Brooks, Travis E.: Myxomycetes of Kansas, I, Trans. Kansas Acad. Sci.,

44:130-157. 1941.
Greene, H. C: Wisconsin Myxomycetes, Trans. Wisconsin Acad. Sci., 27:141-

181. 6 pis. 1932.
Emoto, Yoshikadzu: Die Myxomyceten Japans, Botanical Magazine (Tokyo),
48^ various numbers, 97 figs., 1934; 49, various numbers, 168 figs., 1935;
50, various numbers, mimerous figs., 1936. (Text in Japanese.)
Hattori, H.: Myxomycetes of Nasu District (Japan), pp. 1-280. 23 colored pis.

320 figs. 1935. (Text in Japanese.)
Dennison, Mary Louise: The genus Lamproderma and its relationships, I,
Mycologia, 37(1):80-10S. Figs. 1-22. 1945; II, ihid., 37(2):197-204. Fig. 1.
1945. (Part I contains a key and descriptions of all recognized species of
Lamproderma. Part II includes a discussion of the Family Stemonitaceae and
key to the 13 genera recognized by the author.)
Karling, John S.: The Plasmodiophorales, ix -j- 144 pp. 17 pis. 17 figs. New

York, published by the author, 1942.
Cook, W. R. Ivimey: A monograph of the Plasmodiophorales, Arch. Protistenk.,
8b(2):179-254. Pis. 5-11. Figs. 1-14. 1933.

LIST 5. CHYTRIDIALES 673

Olive, Edgar W.: A preliminary enumeration of the Sorophorae, Proc. Am.

Acad. Arts Sci., 37:333-344. 1901,
: Monograph of the Acrasiae, Proc. Boston Soc. Natural Historij, 30:451-

513. Pis. 5-8. 1902.
Young III, Edward Lorraine : Studies on Labyrinthula. The etiologic agent of the

wasting disease of eel-grass, Am. J. Botany, 30(8) :586-593. Figs. 1-2. 1943.

(Includes a brief monograph of the genus.)

List 4. General Works on Phycomyceteae

Sparrow Jr., Frederick K.: Aquatic Phycomycetes Exclusive of the Saproleg-

niaceae and Pythium, xx + 785 pp. 6Mfigs. Ann Arbor, Univ. Mich. Press,

1943.
: A contribution to our knowledge of the aquatic Phycomycetes of Great

Britain, /. Linnean Soc. London, 50(334) :417-478. Pis. 14-20. Figs. 1-7. 1936.
FiTZPATRiCK, Harry M.: The lower fungi. Phycomycetes, xi + 331 pp. Figs.

1-112. New York, McGraw-Hill Book Co., 1930.
Ito, S.: Mycological flora of Japan: I. Phycomycetes, 340 pp. 125 figs. Tokyo,

1936.
Jaczewski, a. a.: Opredelitel gribov: I. Fikomitsety. (Determination of Fungi:

I. Phycomycetes), 294 pp. 329 figs. Moscow and Leningrad, 1931. (In

Russian.)
VON MiNDEN, M.: Chytridiineae, Ancylistineae, Monoblepharidineae, Sapro-

legniineae, in Krytogamenflora der Mark Brandenburg, vol. 5, pt. 2, pp.

193-352, 1911; pt. 3, pp. 353-496, 1911; pt. 4, pp. 497-608, 1912; pt. 5, pp.

609-630, 1915. (Illustrated.)
Petersen, H. E. : An account of Danish freshwater Phycomycetes, with biological

and systematical remarks, Ann. Mycol., 8(5) :494-560. Figs. 1-27. 1910.
Tiesenkausen, Manfred Baron: Beitrage zur Kenntnis der Wasserpilze der

Schweiz, Arch. HydroUol. Planktonkunde, 7:261-308. Figs. 1-24. 1912.
Vi^GAS, A. P., E A. R. Teixeira: Alguns fungos do Brasil (Phycomycetos),

Bragantia, 3(8) :223-245. 22 pis. 4: figs. 1943.

List 5. Chytridiales

(Note : In some of these references are included fungi now segregated in the
Hyphochytriales and in the nonfilamentous Lagenidiales.)

Petersen, H. E.: Contributions a la connaissance des Phycomycetes marins
(Chytridineae Fischer), Oversigt over Kongelige Danske Videnskabernes
Selskab Forhandlinger, 1905:439-488. Illustrated. 1905.

674 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

ScHERFFEL, A.: Beitragc zur Kenntnis der Chytridineen : II. Einiges iiber neue

oder ungeniigend bekannte Chytridineen, Arch. Protistenk., 54(2): 167-260.

3 pis. 1926.
: Beitrage zur Kenntnis der Chytridineen, III, ibid., 54(3) :510-528. 1926.

(Includes descriptions of all known species of Harpochytrium.)
Atkinson, George F. : The genus Harpochytrium in the United States, Ann.

MycoL, 1(6) :479-502. PL 10. Fig. 1. 1903.
: Notes on the genus Harpochytrium, /. Mycology, 10(1) :3-8. PI. 72. Text

figs. 24-33. 1904.
Jane, Frank W. : A revision of the genus Harpochytrium, /. Linnean Soc. London,

53(348) :28-40. Figs. 1-28. 1946.
Karling, John S.: Brazilian Chytrids: I. Species of Nowakowskiella, Bull. Torrey

Botan. Club, 71(4):374-389. Figs. 1-69. 1944; II. New species of Rhizidium,

Am. J. Botany, 31(5):254-261. Figs. 1-72. 1944; III. Nephrochytrium

amazonensis, Mycologia, 36(4) :351-357. Figs. 1-28. 1944; IV. Species of

Rozella, ibid., 36(6) :638-647. Figs. 1-28. 1944; V. Nowakowskiella macro-

spora n. sp. and other polycentric species. Am. J. Botany, 32(1) :29-35. Figs.

1-51. 1945; VI. Rhopalophlyctis and Chytriomyces, two new chitinophilic

operculate genera, ibid., 32(7):362-369. Figs. 1-61. 1945; VII. Observations

relative to sexuality in two new species of Siphonaria, ibid., 32(9):580-587.

Figs. 1-53. 1945; VIII. Additional parasites of rotifers and nematodes,

Lloydia, 9(1):1-12. Pis. 1-2. 1946; IX. Species of Rhizophydium, Am. J.

Botany, 33(5) :328-334. Figs. 1-37. 1946; X. New species with sunken oper-

cula, Mycologia, 39(l):56-70. Figs. 1-56. 1947.

■ : A synopsis of Rozella and Rozellopsis, ibid., 34(2): 193-208. 1942.

Canter, Hilda M.: Studies on British Chytrids: II. Some new monocentric

Chytrids, Brit. MycoL Soc. Trans., 31(1-2) :94-105. Pis. 1-10. Figs. 1-8. 1947;

III. Zygorhizidium willei Lowenthal and Rhizophidium columnaris n. sp.,

ibid., 31(1-2) :128-135. PL 11. Figs. 1-4. 1947.
ToKUNAGA, Yosio: Studies on the aquatic Chytrids of Japan: II. Olpidiaceae,

Trans. Sapporo Natural History Soc, 13(2):78-84. PL 5. 1933; III. Rhizi-

diaceae, ibid., 13(4) :388-393. PL 11. 1934.
Farlow, W. G. : The Synchytria of the United States, Botan. Gaz., 10(3) :235-240.

PL 4. 1885.
Tobler-Wolfp, Gertrud: Die Synchytrien. Studien zu einer Monographic der

Gattung, Arch. Protistenk., 28:143-238. Pis. 10-13. 1913.
Cook, Melville T. : Species of Synchytrium in Louisiana: I. Descriptions of

species found in the vicinity of Baton Rouge, Mycologia, 37(3) :284-294.

Figs. 1-4. 1945; II. Species of Louisiana Synchytrium, ibid., 37(5) :571-575.

1 fig. 1945; III. The development and structure of the galls, ibid., 37(6) :715-

740. Figs. 1-12. 1945; IV. Two new species of Synchytrium, ibid., 39(3) :351-

357. Figs. 1-4. 1947; V. A new species on Sambucus canadensis, ibid.,

41(l):24-27. Figs. 1-9. 1949.
■ : Synchytrium decipiens and Synchytrium chrysosplenii, ibid., 38(3) :300-

305. Figs. "l-3. 1946.
Quintanilha, a.: Contribui9So ao estudo dos Synchytrium, Boletim da Sociedade

Broteriana II, serie 3, 110 pp. 1 fig. 1926.
Mhatre, J. R., AND B. B. Mundkur: The Synchytria of India, Lloydia, 8(2) :131-

138. 1945.
Cejp, Karel: Some remarks to the knowledge of the parasitic Phycomycetes of

Conjugates in Bohemia, Academic Tcheque des Sciences Mathematiques,

LIST 7. BLASTOCLADIALES AND MONOBLEPHARIDALES 675

Naturelles et de la Medecine, 33:17-23. Pis. 1-2. 1932. (Includes descriptions

of the three known species of Micromycopsis.)
Couch, John N.: Rhizophidium, Phlyctochytrium and Phlyctidium in the United

States, /. Elisha Mitchell Sci. Soc, 47(2) . -245-260. Pis. 14-17. 1932.
: Notes on the genus Micromyces, Mycologia, 29(5) :592-596. Figs. 1-14.

1937.
Canter, Hilda M.: Studies on British Chytrids: VI. Aquatic Synchytriaceae,

Brit. Mycol. Soc. Trans., 32(l):69-94. Pis. 7-11. Figs. 1-13. 1948.

List 6. Hyphochytriales

Karling, John S.: The life history of Anisolpidium ectocarpii gen. nov. et sp
nov., and a synopsis and classification of other fungi with anteriorly uni-
flagellate zoospores, Am. J. Botany, 30(8) : 63 7-648. Figs. 1-21. 1944.

List 7. Blastocladiales and Monoblepharidales

(See also Sparrow, 1943, in List 4.)

CoKER, W. C, AND Velma D. Matthews: Blastocladiales, Monoblepharidales

and Saprolegniales, North Aynerican Flora, 2(l):l-76. 1937.
Sparrow Jr., Frederick K. : The Monoblepharidales, Ann. Botany, 47^87) -517-

542. PI. 20. Figs. 1-2. 1933.
Kanouse, Bessie B.: A monographic study of special groups of water molds: I.

Blastocladiaceae, Am. J. Botany, 14(6) :287-306. Pis. 32-34. 1927.
Wolf, Frederick Taylor: The aquatic Oomycetes of Wisconsin, Pt. I, 64 pp.

Pis. 1-6. Madison, Univ. Wisconsin Press, 1944.
Emerson, Ralph: An experimental study of the life cycle and taxonomy of

AUomyces, Lloydia, 4(2):77-144. Figs. 1-16. 1941.
Couch, John N., and Alma J. Whiffen: Observations on the genus Blasto-

cladiella, Am. J. Botany, 29(7) :582-591. Figs. 1-66. 1942.
: 0})servations on the genus Catenaria, Mycologia, 37(2):163-192. Figs

1-78. 1945.
Cejp, Karel: Sur les affinites des Blastocladiaceae. Revision du genre Gonapodya,

sa position systematique, Bull, trimestr. soc. mycol. France, 62(3-4) :246-=257

1946 (1947).
Couch, John N. : Revision of the genus Coelomomyces, parasitic in insect larvae,

/. Elisha Mitchell Sci. Soc, 61(1-2) :124r-136. Pis. 1-2. 1945.
, and H. R. Dodge : Further observations on Coelomomyces, parasitic on

mosquito larvae, ibid., 63(l):69-79. Pis. 15-20. 1947.
Springer, Martha E.: Two new species of Monoblepharella, Mycologia,

37(2) :208-2lQ. Figs. 1-51. 1945.

676 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Indoh, Hiroharo: Studies on Japanese aquatic fungi: II. The Blastocladiaceae,
Science Repts. Tokyo Bunrika Daigaku, B, 4:237-384. 34: figs. 1940.

List 8. Lagenidiales

(See also Sparrow, 1943, and others in List 4.)

Karling, John S.: The simple holocarpic biflagellate Phycomycetes. Including a
complete host index and bibliography, x + 123 pp. Pis. 1-25. 3 diagrams.
New York, published by the author, 1942.

ToKUNAGA, Yosio: Studies on the aquatic Chytrids of Japan: I. Woroninaceae,
Trans. Sapporo Natural History Soc, 13(l):20-28. PI. 2. 1933.

: Notes on the Lagenidiaceae of Japan, ibid., 13(3) :227-232. 3 figs. 1934.

List 9. Saprolegniales (Including Leptomitales)

(See also Sparrow, 1943, and others in List 4.)

Humphrey, J. E.: The Saprolegniaceae of the United States with notes on other

species. Trans. Am. Phil. Soc, 17:64-148. Pis. 14-20. 1892.
CoKER, W. C: The Saprolegniaceae with notes on other water molds, 201 ]^p.

Pis. 1-63. Chapel Hill, Univ. North Carolina Press, 1923.
, AND Velma D. Matthews: Saprolegniales, North American Flora,

2(i):15-67. 1937.
Nagai, Masaji: Studies on the Japanese Saprolegniaceae, /. Faculty Agr. Hok-
kaido Imp. Univ., 32(l):l-43. Pis. 1-7. 1931.
: Additional notes on the Japanese Saprolegniaceae, Botanical Magazine

(Tokyo), 47(554) :136-137. Fig. 1. 1933.
Maurizio, a.: Zur Entwicklungsgeschichte und Systematik der Saprolegniaceen,

Flora', 79:109-158. Pis. 3-5. 1894.
Wolf, Frederick Taylor: The aquatic Oomycetes of Wisconsin, Pt. I, pp. 1-64.

Pis. 1-6. Madison, Univ. Wisconsin Press, 1944.
Apinis, Arv. : Untersuchungen uber die in Lettland gefundenen Saprolegniaceen

nebst Bemerkungen liber einige andere Wasserpilze, Acta Horti Botan. Univ.

Latviensis, 4:201-246. Figs. 1-4. 1929.
Chaudhuri, H., and p. L. Kochhar: Indian water molds, I, Proc. Indian Acad.

Sci., B, 2 :137-154. Pis. 5-12. 1935.

— , and S. S. Lotus: Indian water molds, II, ibid., 3:328-333. 1936.

Crook's, Kathleen M.: Studies on Australian aquatic Phycomycetes, Proc. Roy.

Soc. Ftctona, 49(2) :206-232.P/. 10. Figs. 1-11. 1937.
Kanouse, Bessie B.: A monographic study of special groups of water molds: II.
Leptomitaceae and Pythiomorphaceae, Am. J. Botany, 14(7) :335-357. PL 48.
1927.

LIST 10. PERONOSPORALES, ALSO PROTOMYCETALES G77

Indoh, Hiroharo: Studies on the Japanese aquatic fungi: I. On Apodachlyella

completa sp. nov., with revision of the Leptomitaceae, Science Repts. Tokyo

Bunrika Daigaku, B, 4:43-50. PL 7. Figs. 1-11. 1939.
CoKER, W. C, AND John N. Couch: Revision of the genus Thraustotheca with a

description of a new species, /. Elisha Mitchell Sci. Soc, 40(3-4): 197-202.

Pis. 38-40. 1924.
Cutter Jr., Victor M.: Observations on certain species of Aphanomyces, Mycol-

ogia, 33(2) :220-240. Figs. 1-15. 1941.

List 10. Peronosporales, Also Protomycetales

Matthews, Velma Dare: Studies on the genus Pythium, 136 pp. 29 pis. Chapel

Hill, Univ. North Carolina Press, 1931.
Sideris, C. p.: Taxonomic studies in the Family Pythiaceae: I. Nematosporan-

gium, Mycologia, 23(4):252-295. Figs. 1-12. 1931; II. Pythium, ibid.,

24(l):14-61.Fi>s. 1-21. 1932.
Drechsler, Charles: Some new species of Pythium, J. Wash. Acad. Sci.,

20(16) :398-418. 1930.
Sparrow Jr., Frederick K. : Two new species of Pythium parasitic on algae, Ann.

Botany, 45(178) :255-277. 1 pi. 2 figs. 1931.
Butler, E. J.: An account of the genus Pythium and some Chytridiaceae, Mem.

Dept. Agri. India, Botan. ser., 1(5):1-161. Pis. I-IO. Feb. 1907.
Fitzpatrick, H. M.: Generic concepts in the Pythiaceae and Blastocladiaceae,

Mycologia, 15(4):166-173. 1923.
MiDDLETON, John T. : The taxonomy, host range and geographic distribution of

the genus Pythium, Mem. Torrey Botan. Club, 20(1):1-171. Figs. 1-17. 1943.
Tucker, C. M.: Taxonomy of the genus Phytophthora de Bary, Univ. Missouri

Agr. Expt. Sta. Research Bull. 153:1-208. Figs. 1-30. 1931.
Rosenbaum, J. : Studies of the genus Phytophthora, /. Agr. Research, 8(7) :233-

276. 7 pis. 1917.
Baker, R. E. D.: Notes on Trinidad fungi: I. Phytophthora, Trop. Agr. Trinidad,

13(12) :330-332. Figs. 1-3. 1936. (The species of Phytophthora on cacao in

Trinidad.)
FoiSTER, C. E. : The white tip disease of leeks and its causal fungus, Phytophthora

Porri n. sp., Trans. Proc. Botan. Soc. Edinburgh, 30(4):257-281. PL 18. Figs.

1-3. 1931. (Describes a new species and gives for comparison the description

of 10 other rather similar species.)
Wilson, G. W.: Studies in North American Peronosporales: I. The genus Albugo,

Bull. Torrey Botan. Club, 34(2):61-84. 1907; II. Phytophthoreae and Rhy-

sotheceae, ibid., 34(8) :387-416. 1907; III. New or noteworthy species

(Species of Albugo and Peronospora), ibid., 35(7):361-365. 1908; IV. Host

Index, ibid., 35(11) :543-554. 1908; V. A Review of the genus Phytophthora,

Mycologia, 6(2):54-83. PL 119. 1914; VI. Notes on Miscellaneous Species,

ibid., 6(4):192-210. Pis. 135-136. 1914; VII. New and Noteworthy Species,

ibid., 10(3):168-169. 1918.
Farlow, W. G.: Enumeration of the Peronosporaceae of the United States

Botan. Gaz., 8(10):305-315, (ll):327-337, 1883.

678 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Farlow, W. G.: Additions to the Peronosporaceae of the United States, ibid.,

9(l):37-40. 1884. (These two papers contain descriptions of every species of

this order known from the United States up to the close of 1883, with full

notes as to known distribution, hosts, etc.)
Swingle, W. T. : Some Peronosporaceae in the Herbarium of the Division of

Vegetable Pathology, /. Mycology, 7(2) :109-130. 1892.
Berlese, a. N.: Saggio di una monografia delle Peronosporaceae, Riv. patol.

vegetale, 6(1):78-101. (2):237-268. Pis. 7-10. 1897; 7:19-37. 1898; 9:1-126.

Figs. 1-21. 1900; 10:185-298. Figs. 22-69, 1902.
: Icones fungorum ad usum Sylloges Saccardianae accomodatae. Phyco-

mycetes: Fasc. I. Peronosporaceae, pp. 1-40. Pis. 1-67. Padua, 1898, pub-
lished by the author. (Beautifully executed illustrations of most of the species

of this family.)
Wakefield, E. M. : The genus Cystopus in South Africa, Bothalia, 2(Ib) :242-246.

1927.
Jaczewski, a. a.: Mycological Flora of European and Asiatic Russia: I.

Peronosporaceae, Materialien zur Kenntnis Fauna und Flora des Russischen

Reiches, Botanischer Teil, Heft 4. 1907. (In Russian.)
Thind, Kartar Singh: The genus Peronospora in the Punjab, /. Indian Botan.

Soc, 21(3-4): 197-2 15. 1942.
Gaumann, Ernst: tJber die Specialisation der Peronospora auf einigen Scrophu-

lariaceen, Ann. MycoL, 16(1-2) :189-199. Figs. 1-6. 1918.
: Ueber die Formen der Peronospora parasitica (Pers.) Fries. Ein Beitrag

zur Speziesfrage bei den parasitischen Pilzen, Beihefte Botan. Centr., Erste

Abteilung, 35:1-143, 305-533. Figs. 1-47. 1918.
— ■ : Zur Kenntnis der Chenopodiaceen bewohnenden Peronospora- Arten,

Mitteilungen der Naturforschenden Gesellschaft in Bern aus dem Jahre 1918 :

45-66. Figs. 1-15. 1919.

Beitriige zu einer Monographie der Gattung Peronospora Corda,

Beitrdge zur Kryptogamenflora der Schweiz, 5(4):l-360. Figs. 1-166. 1923.
Wartenweiler, Alfred: Beitrage zur Systematik und Biologie einiger Plasmo-

para-Arten, Ann. MycoL, 16(3-6) :249-299. Pis. 1-3. Figs. 1-12. 1918.
Savulescu, Trian: Les especes de Peronospora Corda de Roumanie, Sydowia,

Ann. MycoL, 2(1-6) :255-307. 1948.
von BtJREN, GtJNTHER; Die schweizerischen Protomycetaceen mit besonderer

Beriicksichtigung ihrer Entwicklungsgeschichte und Biologie, Beitrdge zur

Kryptogamenflora der Schweiz, 5(l):l-95. Pis. 1-7. Figs. 1-28. 1915.
: Weitere Untersuchungen liber die Entwicklungsgeschichte und Biologie

der Protomycetaceen, ibid., 5(3):l-94. P/s. 1-2. Figs. 1-27. 1922.

List 11. Mucorales, Entomophthorales

Zycha, H.: Mucorineae, Kryptogamenflora der Mark Brandenburg, vol. 6a, pp.
1-264. 114 yigrs. Leipzig, Gebriider Borntraeger,1935.

Naumov, N. A.: Opredelitel Mukorovych (Determination of Mucorales), ed. 2,
pp. 1-136. Figs. 1-49. 1 Diagram. Moscow and Leningrad, Botanical Insti-
tute of the Academy of Sciences of U.S.S.R., 1935. (In Russian.)

LIST 11. MUCORALES, ENTOMOPHTHORALES 679

■: Cl^s des Mucorin^es (Mucorales). Translated by S. Buchet and I. Mou-

raviev, in Encyclop^die Mycologique, ed. 2, vol. 9, pp. 1-137, with Appendix

i-xxxvi. 83 figs. Paris, Paul Lechevalier, 1939.
Lendner, Alf. : Les Mucorin^es de la Suisse, Beitrdge zur Kryptogamenflora der

Schweiz, 3(1):1-180. Pis. 1-3. Figs. 1-59. 1908.
Hagem, 0.: Untersuchungen liber norwegische Mucorineen, Videnskapselskapets-

Skrifter. Mat. naturv. Klasse, 1907(7) :l-50. Figs. 1-22. 1908; 1910(4) :1-1 52.

1910.
• : Neue Untersuchungen liber norwegische Mucorineen, Ann. Mycol.,

8(3) :265-286. Figs. 1-11. 1910.
LiNNEMANN, G. : Beitrag zu einer Flora der Mucorineae Marburgs, Flora, 130(N.S.

30):176-217. Illustrated. 1936.
SuMSTiNE, D. R.: The North American Mucorales, I, Mijcologia, 2(3):125-154.

1910.
Christenberry, George A.: A taxonomic study of the Mucorales in the South-
eastern United States, /, Elisha Mitchell Sci. Soc., 56(2) :333-366. Pis. 13-19.

1946.
Ou, Shih-Kuang: Phycomycetes of China, I, Sinensia, 11(1-2) :33-57. 18 figs.

1940. (Descriptions and keys to the Mucorales found in Szechuan.)
Campbell, Marie E. : An investigation of the Mucorales in the soil, Trans. Roy.

Soc. Edinburgh, 59(2) :41 1-436. 3 pis. 15 figs. 1938.
PovAH, A. H. W.: A critical study of certain species of Mucor, Bull. Torrey

Botan. Club, 44(5) :241-259. (6):287-313. Pis. 17-20. 1917.
Zach, Franz: Zur Kenntnis der Formenkreis von Mucor plumbeus Bonorden,

Oesterreichische Botanische Zeitschrift, 84(2):117-122. Fig. 1. 1935.
: Beitrag zum Formenkreis von Mucor plumbeus Bonorden, ibid.,

85(2):151-153. 1936.
Grove, W. B.: A systematic account and arrangement of the Pilobolidae, in

A. H. Reginald Buller: Researches on Fungi, vol. 6, chap. 4, pp. 190-224.

Figs. 97-111. 1934.
Palla, E.: Zur Kenntnis der Pilobolus-Arten, Oesterreichische Botanische Zeit-
schrift, 50(10) :349-370, (11):397-401, PI. 10. 10 text figs. 1900.
MoRiNi, Fausto: Materiali per una monografia delle Pilobolee, Mem. reale accad.

sci. inst. Bologna, ser. 6, 3:111-129. 1906.
Hanzawa, J.: Studien iiber einige Rhizopus-Arten, Mycolog. Centr., 5(5) :230-246.

(6):257-281. Figs. 1-12. 1915. (A morphological, systematic, and physio-
logical study of 12 species.)
Yamamoto, Yoshihiko: Ein Beitrag zur Kenntnis der Gattung Rhizopus: I.

Morphologisches, J. Faculty Agr. Hokkaido Imp. Univ., 28(1) :1-101. P/s. 1-4.

1930. (Divides Rhizopus into 2 sections. Gives diagnoses for 15 species.)
Yamazaki, MoMOji: On the classification of Rhizopus species, Bidl. Utsunomiya

Agr. Coll., 5:1-16. 1934. (Japanese, with English summary.)
Dauphin, J.: Contribution a I'^tude des Mortierell^es, Ann. sci. not. Botan.,

9me s6r., 8:1-112. Figs. 1-45. 1908.
Vuillemin, Paul: Sur les Mortierella des groupes polycephala et nigrescens.

Bull. soc. mycol. France, 34:41-46. Figs. 1-3. 1918.
Alcorn, Gordon D., and Charles C. Yeager: A monograph of the genus

Cunninghamella with additional descriptions of several common species,

Mijcologia, 30(6) :653-658. Figs. 1-2. 1938.
Cutter Jr., Victor M.: The genus Cunninghamella (Mucorales), Farlowia,

2(3) :321-343. Pis. 1-2. 1946.

680 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

LiNDER, D. H.: The genera Kickxella, Martensella, and Coemansia, Farlowia,
l(l):49-77.P;s. 1-4. 1943.

BucHOLTZ, Fedor: Beitrage zur Kenntnis der Gattung Endogone Link, Beihefte
Botan. Centr., Zweite AbL, 29:147-225. Pis. 3-10. 1912. (Besides morpho-
logical and cytological data this paper also contains descriptions of most of
the known species.)

BoEDiJN, K. B.: The genera Endogone and Sclerocystis in the Netherlands
Indies, Bull. Jardin Botan. de Buitenzorg, III, 13(3) :503-508. Illustrated. 1935.

Thaxter, Roland: A revision of the Endogoneae, Proc. Am. Acad. Arts Sci.,
57(12) :289-350.P^s. 1-4. 1922.

: The Entomophthorae of the United States, Mem. Boston Soc. Natural

History, 4:133-201. Pis. 14-21. 1888.

Berdan, Helen: Revision of the genus Ancylistes, Mycologia, 30(4):396-415.
Figs. 1-22. 1938.

List 12. Zoopagales, Eccrinales

(Note: For both these orders no publication brings together in one place a
discussion of the described species. Therefore the many papers bearing on these
fungi must be referred to.)

Drechsler, Charles: Some conidial Phycomycetes destructive to terricolous

Amoebae, Mycologia, 27(1) :6-40. Pis. 1-7. 1935.
: Some non-catenulate conidial Phycomycetes preying on terricolous

Amoebae, ibid., 27(2):176-205. Figs. 1-5. 1935.
: A new species of conidial Phycomycete preying on nematodes, ibid.,

27{2):20Q-215. Fig. 1. 1935.
: A new species of Stylopage preying on nematodes, ibid., 28(3) :241-246.

Fig. 1. 1936.
: New conidial Phycomycetes destructive to terricolous Amoebae, ibid.,

28(4) :363-389. Figs. 1-7. 1936.
: New Zoopagaceae capturing and consuming soil Amoebae, ibid.,

30(2):137-157.Figrs. 1-4. 1938.
: A few new Zoopagaceae destructive to large soil rhizopods, ibid.,

31(2):128-153. Figs. 1-7. 1939.
: Five new Zoopagaceae destructive to rhizopods and nematodes, ibid.,

31(4):388-415.F2>s. 1-5. 1939.
: Four Phycomycetes destructive to nematodes and rhizopods, ibid.,

33(3):248-269. Ft^s. 1-5. 1941.
: New species of Acaulopage and Cochlonema destructive to soil Amoebae,

ibid., 34(3) :274-297. Figs. 1-6. 1942.
: Several additional Phycomycetes subsisting on nematodes and Amoebae,

ibid., 37(1):1-31. Figs. 1-3. 1945.
: Three new Zoopagaceae subsisting on soil Amoebae, ibid., 38(2) :120-143.

Figs. 1-6. 1946.

Three zoopagaceous fungi that capture and consume soil-inhabiting

rhizopods, ibid., 39(3):253-281. Figs. 1-7. 1947.

LIST 12, ZOOPAGALES, ECCRINALES 681

- — — : Three new species of Zoopage predaceous on terricolous rhizopods, ibid.,

39(4) :379-408. Figs. 1-7. 1947.
: Three Zoopagaceae that subsist by capturing soil Amoebae, ibid.,

40(1):85-105. A>. 1-4. 1948.

(The following five papers concern Family Harpellaceae, including Genistel-
laceae). ,

Leger, Louis, et 0. Duboscq: Harpella melusinae n. g. et n. sp. Entophyte

eccriniforme parasite des larves de Simulie, Compt. rend. 188(14) :951-954.

Figs. 1-6. 1929.
, ET Marcelle Gauthier : Endomycetes nouveaux des larves aquatiques

d'insectes, ibid., 194(26) :2262-2265. Figs. 1-3. 1932.
, ET : La spore des Harpellac^es (Leger & Duboscq), champignons

parasites des insectes, ibid., 200(17) :1458-1460. 1935.

ET : Graminella bulbosa nouveau genre d'entophyte parasite des

larves d'Ephemerides du genre Baetis, ibid., 204(1) :27-29. Figs. 1-5. 1937.
Gauthier, Marcelle: Sur un nouvel entophyte du groupe des Harpellac^es
Leg. & Dub., parasite des larves d'Ephemerides, ibid., 202(12) :1096-1098.
Figs. 1-4. 1936.

(The following papers treat of the Eccrinales.)

Leger, Louis, et 0. Duboscq: Eccrinoides Henneguyi n. g. et n. sp. et la sys-
tematique des Eccrinides, Archives d' Anatomic Microscopique, 25:309-324.
Figs. 1-6. 1929. (The order is classified into families with lists of the con-
tained genera, but no generic distinctions are given. Therefore the following
papers will have to be consulted.)

Duboscq, O., L. Leger, et 0. Tuzet: Contribution a la connaissance des
Eccrinides: les Trichomycetes, Arch. Zool. exptl. et gen., 86(2) :29-144. Pis
1-5. Figs. 1-42. 1948.

Leidy, Joseph: Enterobrus, a new genus of Confervaceae, Proc. Acad. Nat. Sci.
Phila., 4:225-227. 1849.

: A flora and fauna within living animals, Smithsonian Inst. Pubs. Contribs.

to Knowledge, 5(2):l-67. Pis. 1-10. 1851 (1853).

Leger, Louis, et 0. Duboscq: Les Eccrinides, nouveau groupe de Protophytes
parasites, Compt. rend., 141(9) :425-427. 1905.

, et : Sur les Eccrinides des Hydrophilides, Arch. zool. exptl. et gen.,

56(2):21-31. A>s. 1-4. 1916.

ET -: L'^volution du Paramoebidium, nouveau genre d'Eccrinides,

parasite des larves aquatiques d'insectes, Compt. rend., 189(2) :75-77. Figs

1-15. 1929.
LicHTENSTEiN, Jean L. : Sur un Amoebidium a commensalisme interne du rectum

des larves d'Anax imperator Leach: Amoebidium fasciculatum n. sp., Arch.

zool. exptl. et gen., 56(3):49-62. Figs. 1-7. 1917.
: Sur un mode nouveau de multiplication chez les Amoebidiac^es, i6id.,

56(4) :95-99. Fi^. 1. 1917.
PcissoN, Raymond: Recherches sur quelques Eccrinides parasites de Crustac^s

amphipodes et isopodes. Arch. zool. exptl. et gen., 69(3):179-216. Figs. 1-23

1929.
: Recherches sur les Eccrinides. Deuxieme contribution, ibid., 74(4):53-68.

Figs. 1-7. 1931. (Volume jubilaire.)

682 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

PoissoN, Raymond: Asellaria CauUeryi N. G., N. Sp., type nouveau d'entophyte
parasite intestinal des Aselles (Crustaces isopodes). Description des stades
connus et d'une partie de son cycle 6volutif, Bull. biol. France Belg.,
66:232-254. PL 3. Figs. 1-9. 1932.

List 13. Ascomyceteae, Miscellaneous

TuLASNE, L. R., ET C: Selecta fungorum carpologia. Paris, Typographie Im-
periale. Vol. 1. Erysiphei, pp. xxviii + 1-242, Pis. 1-5, 1861; vol. 2. Xylariei,
Valsei, Sphaeriei, pp. xx + 1-319, Pis. 1-34, 1863; vol. 3. Nectriei, Phacidiei,
Pezizei, pp. xvi + 1-221, Pis. 1-22, 1865.

GiLMAN, J. C.: A partial list of the parasitic Ascomycetes of Iowa, Proc. Iowa
Acad. Sci., 32:235-264. 1925.

Fink, Bruce, and Sylvia C. Fuson: An arrangement of the Ascomycetes of
Indiana, Proc. Indiana Acad. Sci., 1919:113-133. 1921.

Miller, Julian H.: The Ascomycetes of Georgia, Plant Disease Reptr., supple-
ment 131:31-93. 1941.

ViEGAS, A. P.: Alguns fungos do Brasil: II. Ascomicetos, Bragantia, 4(1-6) :5-392,
220 pis. 34 figs. 1944.

List 14. Laboulbeniales

Thaxter Roland: Contribution toward a monograph of the Laboulbeniaceae,
I u'em. Am. Acad. Arts Sci., 12:195-429. Pis. 1-26. 1895; II, ibid., 13:219-
469 Pis. 28-71. 1908; III, ibid., 14:309-426. Pis. 1-12. 1924; IV, ibid.,
15(4):427-580. Pis. 1-24. 1926; V, ibid., 16(l):l-435. Pis. 1-60. 1 ^^. 1931.

: Laboulbeniales parasitic on Chrysomelidae, Proc. Am. Acad. Arts Sci.,

50(2):17-50. 1914.
-: New Laboulbeniales from Chile and New Zealand, ibid., 54:207-232.

1918.

New Dimorphomyceteae, ibid., 55:211-282. 1920.

Paoli, G.: Nuovi Laboulbeniomiceti parassiti di Acari, Redia, 7:283-295. 1 pi.

1911.
Spegazzini, Carlos: Contribucion al estudio de las Laboulbeniomicetas Argen-

tinas, Anales del Museo Nacional de Historia Natural de Buenos Aires, 23 :167-

244. 71 figs. 1912.
. : Laboulbcniali ritrovate nelle coUezioni de alcuni nmsei italiani, ibid.,

26:451-511. Illustrated. 1915.
-: Revision de las Laboulbeniales Argentinas, ibid., 29 :445-688. Figs. 1-213.

1917.

Siemaszko, Janina I Wincenty: Owadorosty polskie i palearktyczne (Laboul-
beniales polonici et palaearctici), I-III, Polskie Pismo Entomologiczne {Bui-

LIST 15. LECANORALES AND PYRENULALES 683

letin Entomologique de laPologne), 6(3-4): 188-211. PL 7. 1928; 10(3-4): 149-
188. Pis. 7-10. Figs. 1-2. 1931; 12(1-4) :1 15-138. Pis. 9-10. 1933.

CoLLA, S.: Laboulbeniales : Peyritschiellaceae, Dimorphomycetaceae, Laboul-
beniaceae Heterothallicae, Laboulbeniaceae Homothallicae, Ceratomyce-
taceae, Flora Italica Cryptogama, Fasc. 16:1-157. 108 ^grs. 1934.

MiDDELHOEK, A.: Laboulbeniaceae in Nederland, Nederlandsch Kruidkundig
Archief, 53:86-115. 12 pis. 1943; 56:249-260. 10 figs. 1949.

Lepesme, P.: Revision des Rhachomyces pal^arctique (Laboulbeniaceae), Bull,
trimestr. soc. mycol. France, 58:57-80. Pis. 2-6. 1942.

List 15. Lecanorales and Pyrenulales

(See also in List 1, Zahlbruckner, Band 8 in Rabenhorst: Kryptogamen-
Flora, etc., and Migula in Thomie: Flora von Deutschland, etc.)

Fink, Bruce: The Lichen Flora of the United States. Completed for publication

by Joyce Hedrick, x + 426 pp. 47 pis. 4: figs. Ann Arbor, Univ. Mich. Press,

1935.
TucKERMAN, Edward : A synopsis of the North American lichens: Pt. I. Parme-

liacei, Cladoniei and Coenogoniei, xx + 262 pp. Boston, S. E. Cassino, 1882.

Pt. IL Lecideacei and (in part) Graphidacei. 177 pp. New Bedford, Mass.,

E. Anthony and Sons, 1888.
Schneider, Albert: A Text-book of General Lichenology, xvii + 230 pp. 76 pis.

Binghamton, N. Y., Williard N. Clute and Co., 1897. (Contains a systematic

account with keys to families and genera but not to species.)
Smith, Annie Lorrain : Lichens, xxviii + 464 pp. 135 figs. Cambridge, Cambridge

Univ. Press, 1921. (Pp. 313-342 give keys to families and genera.)
: A monograph of the British lichens, ed. 2, London, published by the

Trustees of the British Museum. Pt. I. xxiii + 519 pp., Pis. 1-71, 1918;

Pt. IL ix + 447 pp.. Pis. 1-63, 1926.
Leighton, W. A.: The lichen-flora of Great Britain, Ireland and the Channel

Islands, ed. 2, pp. i-xlvii, 1-502. Shrewsbury, published by the author, 1872.
Maas Geesteranus, R. A.: Revision of the lichens of the Netherlands. I.

Parmeliaceae, Blumea 6(1): 1-199. 18 figs. 1947-1948.
Zahlbruckner, Alexander: Catalogus lichenum universalis, 9 vols. Leipzig,

Gebriider Borntraeger, 1922-1933. (Vols. 1-8: A systematically arranged

catalogue of all of the families, genera and species of lichens with complete

citation of places of publication, illustrations, etc., but without keys or

descriptions. Vol. 9: Index.)
Fries, Theodor Magnus: Lichenographia Scandinavica sive dispositio lichenum

in Dania, Suecia, Norvegia, Fennia, Lapponia, Rossica hactenus collectorum,

Iv + 639 pp. Illustrated. Uppsala, E. Berling, 1873-1874.
Gall0e, Olaf: Natural History of the Danish lichens. Copenhagen, H. Asche-

houg, Dansk Forlag. Pt. 1. 93 pp., 160 pis., 1927; Pt. 2. 84 pp., 129 pis., 1929;

Pt. 3. 114 pp., 127 pis., 1930; Pt. 4. 81 pp., 133 pis., 1932; Pt. 5. 117 pp., 140

pis., 1936; Pt. 6. 103 pp. 88 pis., 1939; Pt. 7. 72 pp., 101 pis., 1948.
MoREAU, F.: Les lichens: Morphologic, biologic, systematique, in Encyclopedic

Biologique, vol. 3, pp. 1-148. 2 pis. 65 figs. Paris, Paul Lechevalier, 1929.

684 GUIDE TO THE LITERATUEE FOR THE IDENTIFICATION OF FUNGI

Vainio, E. a.: Lichenographia fennica: I. Pyrenolichenes iisque proximi Pyreno-
mycetes et Lichenes Imperfect!, Acta Societatis pro Fauna et Flora Fennica,
49(2):l-274, 1921; II. Baeomyceae et Lecideales, ibid., 53(l):l-340, Map,
1922; III. Coniocarpeae, ibid., 57(1):1-138, 1927; IV. Lecideales, pt. 2
(edited by B. Lynge), ibid., 57(2):1-531, Pis. 1--4, 1934.

: Etude sur la classification naturelle et la morphologie des lichens du

Bresil, ibid., 7(l):l-247, (2):l-256, 1890.

Dodge, Carroll W.: The foliose and fruticose lichens of Costa Rica, I, Ann.
Missouri Botan. Garden, 20(3) :373-467. 1933.

Magnusson, a. H., and A. Zahlbruckner: Hawaiian lichens: I. The families
Verrucariaceae to Peltigeraceae, Arkiv for Botanik, 31A(l):l-96. 1944; II.
The families Lecideaceae to Parmeliaceae, ibid., 31A(6) :1-109. 1944; III. The
families Usneaceae to Physciaceae, ibid., 32A(6):l-89. Pis. 1-10. 1945.

van der Byl, p. a.: Korsmosse van die Unie van Suid-Afrika: I. Familie Rocel-
laceae, II. Familie Teloschistaceae, Ann. Univ. Stellenbosch, 11A(6):1-18.
10 ^^s. 1933; III. Familie Cladoniaceae, ibid., 11A(4):1-13. 5 figs. 1933;
IV. Die geslag Ramalina, ibid., 13A(1):1-13. 12 figs. 1935; V. Familie Col-
lemaceae, ibid., 13A(4):1-11. 4: figs. 1935.

Hasse, Hermann E.: The lichen flora of Southern California, Contribs. U.S.
National Herbarium, 17:1-132. 1913.

Herre, Albert W. C. T.: The lichen flora of the Santa Cruz Peninsula, Cali-
fornia, Proc. Wash. Acad. Sci., 12:27-269. 1910.

Nearing, G. G.: The Lichen Book, 624 pp. 690 drawings. Ridgewood, N.J.,
published by the author, 1947.

Dodge, Carroll W., and Gladys E. Baker: The second Byrd Antarctic expedi-
tion— Botany: II. Lichens and lichen parasites, Ann. Missouri Botan. Garden,
25(2):515-718. PZs. 38-65. 1938.

Magnusson, A. H.: Lichens from Central Asia, Report of the Scientific Expedi-
tion in the North-Western Provinces of China under Sven Hedin, The Sino-
Swedish Expedition Publication No. 13, pp. 1-168, 12 pis., 1940; No. 22, pp.
1-71, 8 pis., 1944.

Fink, Bruce: The lichens of Minnesota, Contribs. United States National Her-
barium, 14:1-269. Pis. 1-51. Figs. 1-18. 1910.

Rasanen, Veli: Das System der Flechten. tjbersicht mit Bestimmungstabellen
der natiirlichen Flechtenfamilien, ihrer Gattungen, Untergattungen, Sek-
tionen und Untersektionen, Acta Botanica Fennica, 33:1-82. 1943.

Zahlbruckner, A.: Additamenta ad Lichenographiam Japoniae, Botanical
Magazine (Tokyo), 41:313-364. Pis. 11-12. 1927.

Vainio, E. A.: Monographia Cladoniarum universalis. Acta Societatis pro Fauna
et Flora Fennica, 4:1-509, 1887; 10:1-499, 1894; 14:1-268, 1897.

Sandstede, H.: Erganzungen zu Wainio's Monographia Cladoniarum uni-
versalis unter besonderer Beriicksichtigung des Verhaltens der Cladonien zu
Asahina's Diaminprobe, Repertorium Specierum Novarum Regni Vegetabilis,
Beiheft 103, 103 pp. 16 pis. 1938.

Evans, Alexander W.: The Cladoniae of Connecticut, Trans. Connecticut Acad.
Arts Sci., 30:357-510. 1930.

: Notes on the Cladoniae of Connecticut, Rhodora, 34(403) :121-142,

(404):153-164, 1932; 40(469) :4-26, 1938.

The Cladoniae of New Jersey, Torreya, 35(4):81-109. 1935.

The Cladoniae of New Jersey. Supplement, ibid., 38(6):137-149. 1938.

A study of certain North American Cladoniae, Bryologisf, 50(1):14-31.

Pis. 1-5. Figs. 1-7. 1947.

LIST 15. LECANOKALES AND PYRENULALES 685

: Supplementary report on the Cladoniae of Connecticut, Trans. Con-
necticut Acad. Arts Sci., 35:519-626. 1944.
Des Abbayes, H.: Revision monographique des Cladonia du sous-genre Cladina

(Lichens), Bull. soc. sci. Bretagne, 16(2):51-156. 1939.
Savicz, V. P.: Die Cladonien Kamtschatkas, Repertorium Specierum Novarum

Regni Vegetabilis, 19 :337-372. 1924.
Hansen, H. Molholm, og Mogens Lund: De Danske arter af slaegten Cladonia

med angivelse af deres udbredelse og forekomst, Botanisk Tidsskrift, 41(1) :1-

80. Apis. S7 figs. 1929.
ToRREY, Raymond H.: Cladoniae in the range of the Torrey Botanical Club,

Torreya, 33(5) :109-129. P/s. 1-4. 1933.

: Cladoniae of the North Woods, ibid., 34(3):57-74. Pis. 1-3. 1934.

Howe Jr., R. Heber: Preliminary notes on the genus Usnea, as represented in

New England, Bull. Torrey Botan. Club, 36(6):309-327. Pis. 21-23. Figs.

A-C. 1909.
• : A manual of the genus Usnea as represented in North and Middle

America, north of the 15th parallel, ibid., 37(1):1-18. Pis. 1-7. 1910.
: Classification de la famille des Usneaceae dans I'Am^'ique du Nord,

Theses presentees a la Faculte des Sciences de TUniversit^ de Paris, s^rie 60,

ordre 71, pp. 1-32. 1912.
: A monograph of the North American Usneaceae, Rept. Missouri Botan.

Garden, 23:133-146. PL 7. 1912. (Part 1, general, apparently all that was

issued.)
: The genus Evernia as represented in North and Middle America, Botan.

Gaz., 51(6) :431-442. Pis. 24-25. 1911.
■ : Oropogon loxensis and its North American distribution, Mycologia,

4(3):152-156. Fififs. 1-2. 1912.
: North American species of the genus Ramalina, The Bryologist, 16 :65-74,

Pis. 5-7. 81-89, 2 pis. 1913; 17:1-7, 2 pis. 17-27, 33-40, 2 pis. Zfigs. 49-52,

65-69, 81-87, 2 pis. 1914.
: The genus Teloschistes in North America, Bull. Torrey Botan. Club,

42(10) :579-583. Figs. 1-2. 1915.

American species of Alectoria occurring north of the fifteenth parallel,

Mycologia, 3(3):106-150. Pis. 41-47. 1911.
DU RiETZ, G. Einar: Vorarbeiten zu einer "Synopsis Lichenum": L Die Gat-

tungen Alectoria, Oropogon und Cornicularia, Arkiv for Botanik, 20A(4):1-

43. 2 pis. 2 figs. 1926.
: Morfologi och systematik hos slaket Ramalina, sarskillt dess skandi-

navisks arter, Svensk Botan. Tid., 20:295-298. 1926. (Keys to sections and to

the Scandinavian species of Ramalina.)
•: Bestamningskema ofver Skandinaviens Stereocaulonarter, ibid., 20(1) :95-

96. 1926.
Dodge, Carroll W.: A synopsis of Stereocaulon with notes on some exotic

species, Ann. cryptogam, exotique, 2(2):93-153. 1929.
Lamb, I. Mackenzie: A monograph of the Uchen genus Placopsis Nyl., Lilloa,

13:151-288. Pis. 1-16. Figs. 1-7. 1947.
Lowe, Josiah L.: The genus Lecidea in the Adirondack Mountains of New York,

Lloydia, 2(4):225-304. Pis. 1-4. Fig. 1. 1939.
Frey, Eduard: Beitrage zur Biologic, Morphologie und Systematik der Um-

bilicariaceen, Hedwigia, 69(5) :2\9-252. Figs. 1-9. 1929.
: Weitere Beitrage zur Kenntnis der Umbilicariaceen, ibid., 71(1-2) :94-

119. Figs. 1-8. 1931.

G8G GUIDE TO THE LITEKATUEE FOR THE IDENTIFICATION OF FUNGI

ScHOLANDER, P. F.: On the apothecia in the lichen family Umbilicariaceae, Nytt

Magasinfor Naturvidenskapene, 75:1-31. Pis. 1-5. Figs. 1-12. 1936.
Berry, Edward Cain: A monograph of the genus Parmelia in North America,

north of Mexico, Ann. Missouri Botan. Garden, 28(1):31-146. 1941.
Tavares, C. das Neves: Contribuicao para o estudo das Parmeliaceas portu-

guesas, Portugaliae Ada Biologica, ser. B, 1(1-2) :1-120. 1945.
Herre, Albert W.: The Parmelias of California, Contribs. Dudley Herbarium of

Stanford Univ., 3(10):313-350. 1946.
• : The Gyrophoraceae of California, Contribs. U.S. National Herbarium,

13:313-321. Pis. 68-73. 1911.
DU RiETZ, G. Einar: Die europaischen Arten der Gyrophora "anthracina"

Gruppe, Arkivfor Botanik, 19(12) :1-14. 1925.
Gyelnik, v.: tJber einige Arten der Gattung Parmeliopsis (Stizenb.) Nyl., Ann.

Mycol., 30(5-6) :456-459. 1932. (Descriptions of some species and key to all

known species of this genus of lichens.)
: Lichenologische Substratstudien (Squamaria radiosa-Gruppe), Hedwigia,

71(1-2) :120-132. 1931.
■: Nephroma-Studien, ibid., 72(l):l-30. Figs. 1-2. 1932.

Darbishire, Otto V.: Monographia Rocellarum: Ein Beitrag zur Flechten-
systematik, Bibliotheca Botanica, 45:1-102. Pis. 1-30. 1898.

HiLLMANN, Johannes: Studien iiber die Flechtengattung Teloschistes Norm,
Hedwigia, 69:303-343. Figs. 1-2. 1930.

: tjbersicht liber die Arten der Flechtengattung Xanthoria (Th. Fr.) Arn.,

ibid., 63(3-4) :198-208. 1922.
-: Bemerkungen iiber einige Arten der Flechtengattung Parmelia, I, ibid.,

78(5-6) :249-267. 1939.
Asahina, Yasuhiko: Leptogium (Section MaUotium) aus Japan. J. Japanese

Botany, 11 :544-566. 20 figs. 1935. (In Japanese.)
Schede, Alwin: Die Sachsischen Arten der Flechtengattung Rhizocarpon (Ram.)

Th. Fr., Beihefte Botan. Centr., Abt. B, 54(1-2) :75-107. Figs. 1-18. 1935.
Lynge, B. a.: a monograph of Norwegian Physciaceae, Videnskapselskapets-

Skrifter, I, Mat.-naturv. Klasse, No. 8, 110 pp. 3 p^s. 1916.
Rasanen, Veli: Bestimmungsschliissel flir die gelben Rhizocarpon-Arten,

-Varietaten und Formen, Repertorium Spec. Nov. Reg. Veg., 52 :127-136. 1943.
Grespi, L.: Notas liquenalogicas. El genero Rhizocarpon en Espana, Boletin

Sociedad Espanola de Historia Natural, 30(5):261-269. 1930.
Bachmann, E.: Die Moriolaceen, Nyt Magazin for Naticrvidenskaberne, 64:170-

328. 3p^s. ISfigs. 1926.
: A revision of the genus Rhizocarpon (Ram.) Th. Fr. in Greenland,

Norges Svalbard og Ishavs-undersokelser, Skriften om Svalbard og Ishavet,

47:1-30. 1932.
Thompson Jr., John W.: The Wisconsin species of Peltigera, Trans. Wisconsin

Acad. Sci., 38:249-271. Pis. 1-6. 1946 (1947).
Zahlbruckner, A.: Die Guttung Lecanoru. Report of the scientific results of the

Norwegian expedition to Novaya Zemlya 1921, Botany, 44:1-32. 4 pis. 1928.
Lettau, Georg: Monographische Bearbeitung einiger Flechtenfamilien, Reper-
torium Specierum Novarinn Regni Vegetabilis, Beiheft 69 (1, Lief. l):l-96. 3

pis. 1932. (Contains key to species of Lecanactidaceae.)
Magnusson, a. H.: a monograph of the genus Acarospora, Kgl. Svenska Vet-

enskapsakad. Handl., tredje ser., 7(4):l-400. 18 maps. Stockholm. 1929.
: The yellow species of Acarospora in North America, Mycologia, 21(5) :249-

260. 1929.

LIST 15. LECANORALES AND PYRENULALES 687

— Supplement to the monograph of the genus Acarospora, Ann. cryptogam.

exotique, 6(1) :13-48. 1933.
— : The lichen genus Acarospora in New Mexico, Meddelanden frdnGotcborgs

Botaniska Tradghrd, 5 :55-72. 1930.

— : New or otherwise interesting Lecanora species, ibid., 6:1-20. 1930.
— : Beitrage zur Systematik der Flechtengruppe Lecanora fusca, ibid.,

7:65-87. 1931.
-: Studien liber einige Arten der Lecidea armeniaca und elata Gruppen,

ibid., 6:93-144. 1930.
: On saxicolous species of the genus Lecidea proper to North America, ibid.,

10:1-53. 1935.
-: Contribution to the taxonomy of the Lecidea goniophila group, ibid.,

16:125-134. 1944.

: A monograph of the lichen genus lonaspis, ibid., 8:1-46. 1933.

: Die Flechtengattung Maronea Mass., ibid., 9:41-66. 1934.

: On the species of Biatorella and Sarcogyne in America, Ann. cryptogam.

exotique, 7(3-4) :115-146. 1934.

On North American, nonsaxicolous species of the genus Rinodina, Botan.

Notiser, 1947(1) :32-54. 1947.
Ericksen, C. F. E.:Lichenologische Beitrage, Hedwigia, 70:21 6-233. Ftgr. 1. 1930.

(Contains a key to the bark-inhabiting crustose lichens of the lowlands of

North Germany, mainly sterile or nearly sterile specimens.)
Olivier, H.: Les lichens pyr^nocarp^s de la flore d'Europe, Bull, de Geographic

Botaniquc, 28:146-152, 168-183. 1918; 29:6-16, 35-48, 97-110. 1919.
ZscHACKE, H.: Die mitteleuropaischen Verrucariaceen, Hedwigia, 60:1-9, 1918;

62:90-154, 1921; 65:46-64, 1924; 67:45-85, 1927.
Malme, Gust. 0. A.-N.: De sydsvenska formerna af Rinodina sophodes (Ach.)

Th. Fr. och Rinodina exigua (Ach.) Th. Fr., Bihang till Kgl. Svenska Vet-

enskapsakad. Handl., Band 21, Afd. Ill, No. 11:1-40. Pis. 1-2. 1895.

(The following series of papers throws light upon the lichen flora of Paraguay
and Southern Brazil, being based almost entirely upon the collections of Regnell.)

Malme, Gust. 0. A.-N.: Die Pannariazeen des Regnellschen Herbars, Arkiv for

Botanik, 20A(3):l-23. 1926.
: Lichenes blasteniospori herbarii Regnelliani, ibid., 20A(9):1-51. 1926.

(Theloschistaceae.)
: Die im Regnellschen Herbar aufbewahrten Arten der Flechtengattung

Lecanactis (Eschw.) Vainio, ibid., 20B(2):l-6. 1926.

: Buelliae itineris Regnelliani primi, ibid., 21A(4):l-42. 1 fig. 1928.

: Lichenes pyrenocarpi aliquot in herbario Regnelliano asservati, ibid.,

22A(6):1-11. 1928.
— : Pyrenulae et Anthracothesia herbarii Regnelliani, ibid., 22A(ll):l-40.

3 figs. 1929.
: Porinae et Phylloporinae in itinere Regnelliani coUectae, ibid., 23A(1,

paper l):l-37. 1930.
■: Die Ramalinen der ersten Regnell'schen Expedition, ibid., 26A(12):l-9.

2 ph. 1934.
: Die Gyalectazeen der ersten Regnell'schen Expedition, ibid., 26A(13):1-

10. 1934.
: Die Stictazeen der ersten Regnell'schen Expedition, ibid., 26A(14):1-18.

3 pis. 2 figs. 1934.

688 GUIDE TO THE LITERATUKE FOR THE IDENTIFICATION OF FUNGI

Malme, Gust. 0. A.-N.: Bacidiae itineris Regnelliani primi, ibid., 27A(5):l-40.
1935.

: Lecideae expeditionis Regnellianae primae, ibid., 28A(7):l-53. 1936.

: Thelotremaceae Brasilienseae imprimis ex herbario Regnelliano cognitae

praeteraque in herbariis Krempelhaberi, Muller Arg., Nylanderi, Wainionis
et Zahlbruckneri asservatae, ibid., 28A(8):1-122. Figs. 1-75. 1936.

Pertusariae expeditionis Regnellianae primae, ibid., 28A(9):l-27. 1936.

Redinger, Karl M.: Die Graphidineen der ersten Regnell'schen Expedition nach
Brasilien 1892-1894: I. Glyphis, Medusulina, und Sarcographa, Arkiv. for
Botanik., 25A(13):l-20. 1 pi. 7 figs. 1933; II. Graphina und Phaeographina,
ibid., 26A(1):1-103. 7 pis. 1934; III. Graphis und Phaeographis, nebst einem
Nachtrage zu Graphina, ibid., 27A(3):1-103. 7 pis. 1935.

List 16. Pezizales Operculati and Inoperculati, Including

"Phacidiales"

(See BiGEARD ET GuiLLEMiN, J. B. Cleland, J. Ramsbottom, v. 0. Graham
in List 34; Massee, vol. 4, in List 1.)

Boudier, Emile : Histoire et classification des Discomycetes d'Europe, vii + 223
pp. Paris, Librairie des Sciences Naturelles, Paul Klincksieck, 1907.

: Icones mycologicae ou iconographie des champignons de France, prin-

cipalement Discomycetes, 4 vols. 600 colored plates. Paris, Paul Klincksieck,
1905-1910.

Seaver, Fred Jay: The North American Cup-Fungi (Operculates), 284 pp.
Frontis. (col.) 45 pis. 15 figs. New York, published by the author, 1928.

: Supplement to North American Cup-Fungi, pp. i-viii + 285-377. Pis.

46-74. Figs. 16-23. New York, published by the author, 1942.

: Photographs and descriptions of Cup-Fungi: I. Peziza, Mycologia, 7(2) :90-

93. Pis. 155-156. 1915; II. Sepultaria, ibid., 7(4):197-199. PI. 161. 1915;
III. Peziza domiciliana and Peziza repanda, ibid., 8(4) :195-198. Pis. 188-189.
1916; IV. Peziza clypeata, ibid., 8(5):235-238. PI. 191. 1916; V. Peziza
proteana and Peziza violacea, ibid., 9(l):l-3. PL 1. 1917; VI. Discina venosa,
ibid., 9(2):53-54. PI. 5. 1917; VII. The genus Underwoodia, ibid., 10(1) :l-3.
Fig. 1. 1918; VIII. Elvela infula and Gyromitra esculenta, ibid., 12(1) :l-5.
PL 1. 1920; IX. North American species of Discina, ibid., 13(2) :67-71. PL 4.
1921; X. Ascotremella, ibid., 22(2):51-54. Pis. 11-12. 1930; XI. Solenopezia,
ibid., 22(3):122-124. PL 16. 1930; XII. Elvelaceae, ibid., 22(4):163-164. Pis.
17-19. 1930; XIII. Subhypogeous forms, ibid., 22(5):215-218. Pis. 22-23.
1930; XIV. A new genus, ibid., 23(4):247-251. Pis. 23-24. 1931 (the genus
Chloroscypha with key and description of the 4 known species) ; XV. The
giant Elvela, ibid., 23(0):409-410. PL 29. 1931; XVI. Stamnaria, ibid.,
24(1) :l-3. PL 1. 1932; XVII. A new species of Godronia, ibid., 24(4) :353-354.
PL 9. 1932; XVIII. Rare species of Godronia, ibid., 25(1) :55-57. PL 15. 1933;
XIX. The cabbage-head fungus, ibid., 25(3) :157-159. Pis. 24-25. 1933; XX. A
new Lamprospora, ibid., 26(1):102-103. PL 14. 1934; XXI. The genus
Calycina, ibid., 26(4) :344-347. PL 40. 1934; XXII. Dasyscypha. ibid.,
28(1) :l-6. Fig. 1. 1936; XXIII. Stamnaria, ibid., 28(2):186-187. Fig. 1. 1936;

LIST 16, PEZIZALES OPERCULATI AND INOPEECULATI 689

XXIV. Chlorociboria, iUd., 28(4):390-394. Figs. 1-2. 1936; XXV. Urnula
geaster, ibid., 29(l):60-65. Figs. 1-3. 1937; XXVI. The genus Diplocarpa,
iUd., 29(2):174-177. Fig. 1. 1937; XXVII. Pezicula on Cornus, ibid.,
29(3) :334-337. Figs. 1-2. 1937; XXVIII. A proposed genus, ibid., 29(6) :678-
680. Fig. 1. 1937 {Wolfina, nov. gen.); XXVIII (bis). A new Helotium, ibid.,
30(1):79-81. Fig. 1. 1938; XXIX. Chloroscypha, ibid., 30(5):594-596. Fig. 1.
1938; XXX. Arachnopeziza, ibid., 30(6):659-663. Fig. 1. 1938; XXXI. Mol-
lisiella, ibid., 31(l):93-95. Fig. 1. 1939; XXXII. Podophacidium, ibid.,
31(3):350-353. Fig. 1. 1939; XXXIII. A new Boudiera, ibid., 31(4) -.499-501.
Fig. 1. 1939; XXXIV. A new Humarina, ibid., 31(5):533-536. Fig. 1. 1939;
XXXV. A new species of Patella, ibid., 32(4) :567-569. Fig. 1. 1940; XXXVI.
A new species and genus, ibid., 34(3):298-301. Fig. 1. 1942 (Pestalopezia
nov. gen.); XXXVII. Pezicula purpurascens, ibid., 34(4):412-415. Fig. 1.
1942; XXXVIII. The genus Kriegeria, ibid., 35(4) :492-493. 1943; XXXIX. A
new Helotium, ibid., 37(2) :267-269. Fig. 1. 1945; XXXIX (bis). The genus
Godronia and its allies, ibid., 37(3):333-359. S figs. 1945; XLI. Catinella
nigro-olivacea, ibid., 38(4) :473-476. Fig. 1. 1946; XLII. Gorgoniceps, ibid.,
38(5):549-5o3. 2 figs. 1946; XLIII. Seaverinia, ibid., 39(1):113-119. Fig. 1.
1947.

Nannfeldt, J. A.: Studien iiber die Morphologie und Systematik der nicht
lichenisierten inoperculaten Discomyceten, Nova Acta Regiae Soc. Sci.
Upsaliensis, Ser. IV, 8(2):l-368. Pis. 1-19. Figs. 1-47. 1932.

Phillips, William: A manual of British Discomycetes, in International Science
Series, vol. 61, 462 pp. 12 pis. London, Kegan Paul, Trench, Triibner and
Co., 1887; ed. 2, 1893.

Grelet, L. J. : Les Discomycetes de France d'apres la classification de Boudier,
Fascicules 1-8, Bull. soc. botanique du Centre-Ouest, 1932-1940; Fasc. 9-19,
Rev. mijcoL, N. S., vols. 7-14, 1942-1949 (to be continued.)

LeGal, Marcelle: Florule mycologique des Bois de la Grange et de I'Etoile
(Seine-et-Oise) : Discomycetes (Opercules), Rev. mycol., N.S., 2:150-162,
197-222. Figs. 1-27. 1937; Discomycetes (Inopercules), ibid., 3:129-147.
Figs. 1-9. 1938; 4:25-63. Figs. 10-29. 1939.

: Quelques Galactinia de la flore Frangaise, ibid., 4:169-186. Figs. 1-8.

1939; 5:102-112. Figs. 1-3. 1940; 10:90-95. Figs. 1-3. 1945.
-: Les Aleuria et les Galactinia, ibid., 6(supplement 3):56-82. Figs. 1-4.

1941.
Lagarde, J.: Contribution a I'etude des Discomycetes charnus, Ann. Mycol.,

4(2):125-201, (3):203-256. Pis. 3-4. Figs. 1-58. 1906.
: Discomycetes de France: I. Les morilles, pp. 1-35, Pis. 1-5, 1923; II. Les

Helvelles, pp. 39-82, Pis. 6-12, 1924. Paris, La Pens^e Frangaise.
Ramsbottom, J. : A list of the British species of Discomycetes arranged according

to Boudier's system with a key to the genera, Brit. Mycol. Soc. Trans.,

4(2):343-381. 1914.
Kanouse, Bessie B.: A survey of the Discomycete flora of the Olympic National

Park and adjacent areas, Mycologia, 39(6) :635-689. Figs. 1-35. 1947.
: The genus Plectania and its segregates in North America, ibid., 40(4) :482-

497. Figs. 1-12. 1948.

Studies in the genus Otidea, Mycologia, 41(6) :660-677. Figs. 1-21. 1949.

Velenovsky, J.: Monographia Discomycetum Bohemiae, vol. 1:1-436; vol. 2:

pis. 1-31. Prague, 1934.
KiLLERMANN, S.: Bayerische Becherpilze: I. Eupezizaceen. Mit kritischen

Bemerkungen, Kryptogamische Forschungen herausgegeben von der Krypto-

690 GUIDE TO THE LITERATUBE FOR THE IDENTIFICATION OF FUNGI

gamenkommission der Bayerischen Botanischen Gesellschaft zur Erforschung
derHeimischen Flora, 2(1) :27-47. P/s. 1-3. 1929; II. Pezizeae, ibid., 2(3):255-
296. Pis. 4-5. 1935.
Rick, J.: Monographia Pezizinearum Riograndensium, Broteria, Ser. Boian.,
25(2):77-122. 1931; continued in Broteria, Serie Trimestral, Ciencias Na-
turais, l(l):35-46, (2):89-96, (3):97-105. 1932.
McLennan, Ethel, and Isabel Cookson: Addition to the Australian Asco-

mycetes, I, Proc. Roy. Soc. Victoria, N.S., 35(2):153-158. Ph. 8-10. Fig. 1.

1923; II, ibid., 38:69-76. Pis. 4-6. 1926.
, and Francis Halsey: Addition to the Australian Ascomycetes, III, ibid.,

49:51-62. PL 2. 1936.
Rod WAY, L.: Tasmanian Discomycetes, Papers and Proc. Roy. Soc. Tasmania,

1924:90-122. 1925.
KiLLERMANN, S. : Morcheln und andere Helvellaceen aus Bayern, Kryptogamische

Forschungen der Bayerischen Botanischen Gesellschaft, 3 :148-154. 1 fig. 1918.
Imai, Sanshi : Contributions to the knowledge of the classification of the Helvel-

laceae, Botanical Magazine {Tokyo), 46(544) : 172-175. 1932.
Heim, Roger: Tableaux pratiques de determination des principales morilles, in

"Le culture des Morilles," Rev. Mycol., l(supplement to 2) :22-25. 2 pis. 1936.
MosER, Meinhard: Uber das Massenauftreten von Formen der Gattung Mor-

chella auf Waldbrandflachen, Sydowia, Ann. Mycol. , 3(1-6) :176-195. Figs.

1-4. 1949. Includes critical descriptions of the several species occurring in

the burned area.
Overholts, L. 0.: The morels of Pennsylvania, Proc. Penna. Acad. Sci., 8:108-

114. Figs. 1-8. 1934.
Rick, J.: Monographia Helvellinearum Riograndensium, Broteria, Ser. Botan.,

25(2) :72-76. 1931.
Klika, Jaromir: Prispevek k poznani hub chrapd,covitych v Ceskoslovensku

(Czechoslovakian Helvellaceae), Vestnik Krdlovski Ceske Spolesnosti Nauk

Tr. II, 1924:1-54. Hfigs. 1925.
BouDiER, £.: Revision analytique des morilles de France, Bull. soc. mycol. France, I

13:129-153.1897.
Hone, Daisy S.: Minnesota Helvellineae, Minnesota Botanical Studies, 3:309-

321. Pis. 48-52. 1904.
Anderson, P. J., and Marguerite G. Ickis: Massachusetts species of Helvella,

Mycologia, 13(4-5) :201-229. Pis. 11-12. 1921.
Massee, George: A monograph of the Geoglossaceae, Ann. Botany, 11(42) :225-

306. Pis. 12-13. 1897. ^

DuRAND, E. J. : The Geoglossaceae of North America, Ann. Mycoi., 6(5) :387-477. j

Pis. 5-22. 1908. (A very valuable work with full descriptions of all species I

and illustrations of many of them.)
: New or noteworthy Geoglossaceae, Mycologia, 13(3):184-187. 1921.

(Supplementary to the preceding.)
SiNDEN, J. W., AND H. M. Fitzpatrick: a new Trichoglossum, Mycologia,

22(2) :55-61. PL 13. 1930. (A new species and corrections to Durand's paper.)
Lloyd, C. G.: The Geoglossaceae, Mycological Writings 5, pp. 1-24 (separate

pagination). Figs. 782-807. 1916. (Based largely on Durand's paper but with

some modification of generic names and limits.)
VAN LuiJK, A.: Fungi van Nederland: I, Geoglossaceae van Nederland, Neder-

landsch Kruidkundig Archie}, 1918:111-144. 12 figs. 1919.
Imai, Sanshi: Studies on the Geoglossaceae of Japan: I. Trayis. Sapporo Natural
History Soc, 13(3):179-184. PI. 7. 1934; II. The genus Leotia, Botanical

LIST 16. PEZIZALES OPERCULATI AND INOPERCULATI 691

Magazine {Tokyo), 50(589) :9-16. 1936; III. The genus Cudonia, ihid.,

50(600) :671-676. 1936; IV. The genus Microglossum, ibid., 52(620) :41 7-424.

1938.
: Geoglossaceae Japoniae, J. Faculty Agr. Hokkaido Lmp. Univ. 45(4) :155-

264. Pis. 6-10. Figs. 1-6. 1941.
: Contributiones ad studia monographica Geoglossacearum Botanical

Magazine (Tokyo), 56(671): 523-527. Figs. 1-3. 1942.
: The Geoglossaceae of Norway, Ann. Mijcol., 38(2-4) :268-278. Figs. 1-4.

1940.
Nannfeldt, J. a.: The Geoglossaceae of Sweden (with regard also to the sur-

roundmg countries), Arkiv for Botanik, 30A(4):l-67. 5 pis. 6 figs. 1942.
Tai, F. L.: Studies in the Geoglossaceae of Yunnan, Lloydia, 7(2):146-162. 35

figs. 1944.
Whetzel, H. H. : a synopsis of the genera and species of the Sclerotiniaceae, a

family of stromatic inoperculate Discomycetes, Mycologia, 37(6) :648-714.

Figs. 1-36. 1945.
: North American species of Sclerotinia: I. ibid., 18(5) :224-235. Pis. 27-29.

1 fig. 1926; II. Two species on Carex, S. Duriaeana (Tul.) Rehm. and S.

longisclerotialis n. sp., ibid., 21(l):5-32. Pis. 1-5. Fig. 1. 1929.
: Septotinia, a new genus of the Ciborioideae, ibid., 29(1) :128-146. Figs.

1-18. 1937.
: A new genus and new species of brown-spored inoperculate Discomycetes

from Panama, ibid., 34(5):584-591. Figs. 1-5. 1942. (Martinia.)

A monograph of Lambertella, a genus of brown-spored inoperculate

Discomycetes, Lloydia, 6(l):18-52. Pis. 1-6. Figs. 1-7. 1943.
— : A new genus of the Sclerotiniaceae, Farlowia, l(3):483-488. Figs. 1-6.

1944. (Coprotinia.)
— : The cypericolous and juncicolous species of Sclerotinia, ibid., 2(3) :385-

437. Pis. 1-10. Figs. A-D. 1946.
— , AND N. Fabritius Buchwald: North American species of Sclerotinia and

related genera: III. Ciboria acerina, Mycologia, 28(6) :514-527. Figs. 1-19.

1936.
-, AND W. G. Solheim: Sclerotinia Caricis-ampuUaceae, a remarkable sub-

arctic species, ibid., 35(4) :385-398. Figs. 1-6. 1943.
Honey, Edwin E.: The monilioid species of Sclerotinia, ibid., 20(3):127-157.

Pis. 17-19. Figs. 1-4. 1928.
: North American species of Monilinia: I. Occurrence, grouping and life

histories, Am. J. Botany, 23(2):100-106. Figs. 1-4. 1936.
Weiss, Freeman: Ovulinia, a new generic segregate from Sclerotinia, Phyto-
pathology, 30(3) :236-244. Figs. 1-3. 1940.
Buchwald, N. Fabritius: Sclerotiniaceae Daniae. En floristik-systematisk over-

sigt over de i Danmark fundne knoldbaegersvampe, Friesia, 3(4) :235-330.

33 ^^s. 1947.
White, W. Lawrence: A monograph of the genus Rutstroemia (Discomycetes),

Lloijdia, 4(3):153-240. Figs. 1-75. 1941.
, AND H. H. Whetzel: Pleomorphic life cycles in a new genus of the

Helotiaceae, Mycologia, 30{2) -.187-203. Figs. 1-21. 1938. (Pycnopeziza.)
Whetzel, H. H., and W. Lawrence White: MoUisia tetrica, Peziza sejournei,

and the genera Phaeociboria and Pycnopeziza, ibid., 32(5) :609-620. 1940.
Klika, Jaromir: Poznamky k vyskytu druhu r. Humaria v Ceskoslovensku

(Species of Humaria in Czechoslovakia), Vestnik Krdlovske Ceske Spolecnosti

Nauk, 1926(12): 29 pp. 1 fig. 1927.

692 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Klika, Jaromir: 0 druzich r. Barlaea v Ceskoslovensku (Species of the genus
Barlaea in Czechoslovakia), Preslia, 1926(4) :14-19. 1 fig. 1926.

KuPFER, Elsie N.: Studies on Urnula and Geopyxis, Bull. Torrey Botan. Club,
29(3):137-144. 1902.

RouppERT, C: Revision du genre Sphaerosoma, Bull, de VAcademie des Sciences,
Cracovie, Classe Sci., Math., Nat., 1909:75-95. 1909.

KiLLERMANN, S. : Die Bulgaria Fr.-Gruppe, Hedwigia, 69(1-2) :84-93. 1 pi. 1929.
(A revision of the genera Coryne, Bulgaria, Burkardia, Bulgaropsis, and
Neobulgaria.)

BoEDijN, K. B.: The genus Sarcosoma in Netherlands India, Bull. Jardin Bot-
aniq^ie de Buitenzorg, ser. Ill, 12(2):273-279. Illustrated. 1932.

: The genera Phillipsia and Cookeina in Netherlands India, ibid., 13(1) :57-

76. Ilhistrated. 1933.

KoBAYASHi, Yosio: On the gelatinous cup fungi, Bulgaria group, J. Japanese
Botany, 13:510-520. \Q figs. 1937.

LoHMAN, M. L., AND Edith K. Cashi Atropellis species from pine cankers in the
United States, J. Wash. Acad. Sci., 30(6):255-262. Figs. 1-2. 1940.

White, W. Lawrence: Studies in the genus Helotium: I. A review of the species
described by Peck, Mycologia, 34(2): 154^1 79. Figs. 1-16. 1942; II. Lachnum
pygmaeum and the status of the genus Helolachnum, Am. Midland Natural-
ist, 28:512-523. 1942; III. History and diagnosis of certain European and
North American foliicolous species, Farlowia, 1(1):135-170. Figs. 1-17. 1943;
IV. Some miscellaneous species, ibid., 1(4):599-617. Figs. 1-40. 1944.

Groves, J. Walton: Three Pezicula species occurring on Alnus, Mycologia,
32(1) :1 12-123. Figs. 1-12. 1940.

: The genus Dermea in North America, ibid., 38(4):351-431. Figs. 1-57.

1946.

AND A. Mavis Leach: The species of Tympanis occurring on Pinus,

Mycologia, 41(l):59-76. Figs. 1-7. 1949.
Seaver, Fred J., and J. M. Waterston: Contributions to the mycological flora

of Bermuda, II, Mycologia, 33(3):310-317. Figs. 1-2. 1941. (Key to the eight

species of Stictis known from Bermuda.)
Butler, Ellys Theodora: Studies in the Patellariaceae, Mycologia, 32(6) :741-

823. Figs. 1-10. 1940. (A monograph of the North American species of

Lecanidion ( = Patellaria Fr.) and Karschia Koerb.)
Darker, Grant Dooks: The Hypodermataceae of Conifers, Contribs. Arnold

Arboretum Harvard Univ., 1:1-131. Pis. 1-27. 1932.
Hahn, Glenn Gardner, and Theodore T. Ayers: Dasyscyphae on conifers in

North America: I. The large-spored white excipled species, Mycologia,

26(1):73-101. Pis. 8-13. 1934; II. D. Ellisiana, ibid., 26(6):167-1S0. Pis.

21-23. 1934; III. Dasyscypha Pini, ibid., 26(6):479-501. Pis. 52-53. 1934;

IV. Two new species on Douglas fir from the Pacific Coast (senior author,

only), ibid., 32(2):137-147. Figs. 1-2. 1940.
Dennis, R. W. G. : A revision of the British Hyaloscyphaceae, with notes on

related European spec-ies, Commonwealth Mycological Institute Mycological

Papers, 32:1-97. Figs. 1-104. 1949.
Tehon, Leo Roy: A mono^^iaphic rearrangement of Lophodermium, Illinois

Biological Memoirs, 13(4) :1 -151. PZs. 1-5. Fig. 1. 1935.
: New si)ecies and taxouomic changes in the Hypodermataceae, Mycologia,

31(6) :674-692. Fi^s. 1-6. 1939.
Terrier, Charles A.: Essai sur la syst^matique des Phacidiaceae (Fr.) sensu

LIST 17. TUBERALES 693

Nannfeldt (1932), Beitr. zur Kryptogamenflora der Schweiz, 9(2):l-99. Pis.

1-12. Figs. 1-10. 1942.
VON HoHNEL, Fr.: System der Phacidiales v. H., Ber. deut. botan. Ges., 35:416-

422. 1917. (Includes keys to families and genera believed by the author to be

intermediate between Pezizales and Dothideales.)
Petrak, F.: tJber die Leptopeltineen, Sydowia, Ann. Mycol., 1(4-6) :232-247.

1947. (Contains a key distinguishing the genera Leptopeltella, Leptopeltis,

Leptopeltopsis, Leptopeltina, Moeszopeltis, all accredited to the "Order"

Phacidiales.)
DuRAND, E. J.: The genus Catinella, Bull. Torrey Botan. Club, 49(1):15-21. 1922.
Palm, B. T. : On Cyttaria Berk, and Cyttariella n. gen., Ann. Mycol., 30(5-6) :405-

420. Figs. 1-3. 1932. (A morphological and taxonomic study of the known

species of Cyttaria and of Cyttariella, an imperfect stage of that genus.)

List 17. Tuberales

Fischer, Eduard: Abteilung Eumycetes (Fungi): Klasse Ascomycetes: Reihe

Euascales; Unterreihe 8, Tuberineae, in A. Engler und K. Prantl: Die

Natiirlichen Pflanzenfamilien, Zweite Auflage, vol. 5b, pp. 1-42. figs. 1-22.

Leipzig, Wilhelm Engelmann, 1938.
Malencon, Georges: Les truffes europeennes. Historique, morphogenie, organo-

graphie, classification, culture. Rev. Mycol., N.S., 3, Memoire hors-serie No.

1, pp. 1-92. Pis. 1-2. Figs. 1-10. 1938.
Bataille, F. : Flore analytique et descriptive des Tuberoidees de I'Europe et de

I'Afrique du Nord, Bull. soc. mycol. France, 37:155-207. 1921.
DE Ferry de la Bellone, C: La truffe. Etude sur les truffes et les truffieres,

viii + 312 pp. 1 pi. 21 figs. Paris, Librairie J.-B. Bailliere et Fils, 1888.
Hesse, R.: Die Hypogaen Deutschlands. Natur, und Entwicklungsgeschichte

sowie Anatomie und Morphologie der in Deutschland vorkommenden Triif-

feln und der diesen verwandten Organismen nebst praktischen Anleitungen

bezliglich deren Gewinnung und Verwendung, vol. 2, pp. 1-140. Halle a.S.,

Ludw. Hofstetter, 1894. (Tuberales.)
Bucholtz, F. Beitrage zur Morphologie und Systematik der Hypogaen nebst

Beschreibung aller bis jetzt in Russland angetroffenen Arten, 196 pp. 5 pis.

Text figs. Riga, 1902. (Russian, with German summary.)
Jaczewski, Arthur Louis: Monographie des Tuberacees de la Suisse, Bidl.

VHerbier Boissier, 4:591-602. 1909.
Fries, T. M. : Skandinaviens tryfflar och tryffelliknande svampar, Svensk Botan.

Tid., 3:224-300. 1909.
Massee, George: The structure and affinities of the British Tuberaceae, Anyi.

Botany, 23(90) :243-263. PI. 17. 1909.
Harkness, H. W. : California hypogaeous fungi, Proc. Calif. Acad, of Sci., ser. 3,

1 :241-292. Pis. 42-45. 1899.
GiLKEY, Helen M.: A revision of the Tuberales of California, Univ. Calif. Pubs.

Botany, 6:275-356. Pis. 26-30. 1916.

694 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

GiLKET, Helen M.: Tuberales of North America, Oregon State Monographs.

Studies in Botany, No. 1:1-63. Pis. 1-5. 1939.
Mattirolo, Oreste: Catalogo ragionato dei funghi ipogei raccolti nel Canton

Ticino e nelle provincie Italiane confinanti, Beitrdge zur Kryptogamen flora

der Schweiz, 8 (2) :l-53. Pis. 1-2. 1935.
SoEHNER, Ert: Bayerische Pachyphloeus-Arten, Hedwigia, 75(4) :243-254. Figs.

1-20. 1935.
Imai, Sanshi: Second note on the Tuberales of Japan, Proc. Imp. Acad. Tokyo,

16:153-154. 3 A<7s. 1940.

List 18. Taphrinales

(See also F. Neger: Exoascaceae, in Kryptogamenflora der Mark Branden-
burg.)

Mix a J • The genus Taphrina: I. An annotated bibliography; II. A list of valid
species, Vniv. Kansas Sci. Bull, 24(9):113-149, (10):151-176. 1936. (In-
cludes brief descriptions of the 104 recognized species of Taphrina.)

: Species of Taphrina on North American ferns, Mycologia, 30(5):563-579.

Figs. 1-3. 1938.

: New species of Taphrina and new records from western North America,

Am. J. Botany, 26(1) :44-48. Figs. 1-2. 1939.

A monograph of the genus Taphrina, Univ. Kansas Sci. Bull., 33, Pt.

1(1) ■.1-1Q7. Figs. 1-39. 1949.
Ray, W. Winfield: Contributions to the knowledge of the genus Taphrina in

North America, Mycologia, 31(l):56-75. Figs. 1-23. 1939. (Discusses espe-
cially the species found on Alnus and on Prunus.)
Jenkins Anna E., and W. Winfield Ray: A new host for Taphrina dearnessii

and 'geographic distribution of Taphrina on North American Maples, Mtj-

cologia, 32(3):404-414. Figs. 1-4. 1940.
Sadebeck, R. : Die parasitische Exoasceen, eine Monographic, Jahrh. der Ham-

hurgischen Wissenschaftlichen Anstalten, 10(2):1-110. Pis. 1-3. 1893.
Giesenhagen, Karl: Die Entwicklungsreihen der parasitischen Exoasceen,

Flora Oder Allgemeine Botanische Zeitschrift, 81:267-361. 1895.
: Taphrina, Exoascus und Magnusiella, Botan. Zfg., 59(Erste Abt.):115-

142. PL 5. 1901.
NiSHiDA, T. : A contribution to the monograph of the parasitic Exoascaceae of

Japan, in Miyabe Festschrift, pp. 157-212. Pis. 15-19. Tokyo, Rokumeikwan,

1911. (Japanese, with English summary.)
RosTRUP, E.: Taphrinaceae Daniae. Danmarks Taphrinaceer, Dansk Natur-

historisk Forening i Kjohenhavn, Videnskabelige Meddelelser, 1890:246-264.

Illustrated. 1890.
Johanson, C. J.: Studier ofver Svampslaget Taphrina, Kgl. Svenska Vetenskap-

sakad. Handl. Bihang, 13, Abt. 3, No. 4, pp. 3-28. Plate. 1887.
Patterson, Flora W.: A study of the North American paiasitic Exoascaceae,

Bull. Laboratory of Natural History of the Univ. of Iowa, 3 :89-135. Pis. 1-4.

1895.

LIST 20. SPHAERIALES G95

JuEL, 0.: Om Taphrina-Arter pa Betula, Svensk Botan. Tid., 3:183-191. 1909.
Bataille, F.: Monographie des Exoascees d'Europe, Ann. soc. linneenne Lyon,

79:121-130. 1935.
Jaczewski, a. a.: Exoasci of Caucasus, Izvyestia S. Petersburgskovo Botanitsche-

skovo Sada, 1:5-18. 1901. (In Russian.)
■ : Pocket key for the determination of fungi: I. Exoascales, Leningrad,

A. A. Jaczewski Mycological Laboratory, State Institute of Experimental

Agriculture, 1926. (In Russian.)
Palm, B.: Svenska Taphrinaarter, Arkiv for Boianik, 15(4):1-41. Figs. 1-9. 1918.

List 19. Hysteriales

(See also Nannfeldt, in List 16, for some genera sometimes included in this
order.)

BiSBY, G. R.: The literature on the classification of the Hysteriales, Brit. Mycol.

Soc. Trans., 8:176-189. 1923.
: British species of Hysterium, Gloniopsis, Dichaena and Mytilidion, ibid.,

25(2):]27-140. 1 pi. I fig. 1941.
VON Hohnel, Franz: Mycologische Fragmente: CCLXXII. Uber die Hysteri-

aceae, Ann. Mycol., 16(1-2) :145-154. 1918.
LoHMAN, M. L. : Studies in the genus Glonium as represented in the Southeast,

Bull. Torrey Botan. Club, 64(2):57-72. Pis. 1-2. 1 rnap. 1937.

List 20. Sphaeriales

(Many of the fungi included in this list probably should be distributed among
the Pseudosphaeriales, Hemisphaeriales, Myriangiaceae, and possibly other
groups. In all but the newest works these are sometimes lumped with the Sphae-
riales from which only developmental studies can with certainty distinguish
them.)

Ellis, J. B., and B. M. Everhaet: The North American Pyrenomycetes, iii +
793 pp. Pis. 1-41. Newfield, N.J., Ellis and Everhart, 1892. (This work
contains not only Sphaeriales but also Erysiphales, Dothideales, Hypocreales,
and genera now removed to other orders as well.)

Berlese, a. N. : Icones fungorum omnium hucusque cognitorum ad usum Syl-
loges Saccardianae accommodatae. Padua, published by the author, 1894-
1905. Vol. 1. Lophiostomaceae and Sphaeriaceae : Phaeo- and Hyalophrag-
miae, xiv + 243 pp.. Pis. 1-162, Generic plates 1-22, 1894; vol. 2. Sphae-
riaceae: Phaeodictyae, Hyalodictyae, Scolecosporeae, 216 pp.. Pis. 1-178,
Generic plates 1-10, 1900; vol. 3. Sphaeriaceae: Allantosporae, 120 pp., Pis.
1-172, 1905.

696 GUIDE TO THE LITERATUEE FOR THE IDENTIFICATION OF FUNGI

Chenantais, J. S.: Etudes sur les Pyrenomvcetes, Bull. soc. mijcol. France,
34:47-73. 123-236. Figs. 1-7. 1918; 35:46-98. 113-139. Pis. 1-6. Figs. 8-25.
1919. (Takes up many of the fundamental bases for our present classification
of this group and shows the errors. Discusses in particular the genera Nitsch-
kea, Lophiotrema, Rosellinia, Otthia, Massarinula, Lasiosordaria, Podo-
spora, and various species of other genera.)

Cotton, A. D.: Notes on marine Pyrenomycetes, Brit. Mtjcol. Soc. Trans.,
3:92-99. 1907.

Sutherland, George K.: New marine Pyrenomycetes, Brit. Mycol. Soc. Trans.,
5(1):147-155. PL 3. 1915.

: Additional notes on marine Pyrenomycetes, ibid., 5(2) :257-263. PL 5.

1917.

: New marine fungi on Pelvetia, New Phytologist, 14:33-43. ^ figs. 1915.

: Additional notes on marine Pyrenomycetes, ibid., 14:183-193. 5 figs.

1915.
Barghoorn, E. S., and D. H. Linder: Marine fungi: their taxonomy and biology,

Farlowia, l(3):395-467. Pis. 1-7. Figs. 1-2. 1944.
Seaver, Fred J.: Fimetariales, North American Flora, 3:57-88. 1910. (Includes

Families Chaetomiaceae, by H. L. Palliser and Fimetariaceae (Sordariaceae)

by F. J. Seaver.)
Griffiths, David: The North American Sordariaceae, Mern. Torreij Botan. Club,

11:1-134. Pis. 1-19. Figs. 1-6. 1901.
Stratton, Robert: The Ascomycetes of Ohio: III. The Fimetariales of Ohio,

Ohio Biological Survey, 3:75-144. Pis. 1-18. 1921.
Cain, Roy F.: Studies of coprophilous Sphaeriales in Ontario, Univ. Toronto

Studies, Biol. Ser., 38:1-126. Figs. 1-96. 1934.
, and J. W Groves: Notes on seed-borne fungi: VI. Sordaria, Ca7i. J.

Research, C,2Q{5) A8Q^ 495. Figs. 1-27.19^8.
Bayer, August: Monogratickd studia stredoevropskych druhii celedi. Sordari-
aceae. (A monograph of the central European species of the family Sorda-
riaceae), Acta Societatis Scientiarum Naturalium Moravicae Brno, 1(4):1-185.

Qfigs. 1894. (With French resum^.)
Bainier, G.: Monographie des Chaetomidium et des Chaetomium, Bull. soc.

mycol. France, 25:191-237. Pis. 10-26. 1910.
Chivers, a. H. : a monograph of the genera Chaetomium and Ascotricha, Meyn.

Torrey Botan. Club, 14:155-240. Pis. 6-17. 1915.
Greathouse, Glenn A., and L. M. Ames: Fabric deterioration by thirteen

described and three new species of Chaetomium, Mycologia, 37(1):138-155.

Figs. 1-7. 1945.
Ames, L. M. : New cellulose destroying fungi isolated from military material and

equipment, Mycologia, 41(6):637-648. /'\>s. 1-42. 1949. (Descriptions of nine

new species of Chaetomium.)
Skolko, a. J., AND J. W. Groves: Notes on seed-borne fungi: V. Chaetomium

species with dichotomously branched hairs. Can. J. Research, C, 26(3) :209-

280. Pis. 1-7. 1948.
Benjamin, R. K. : Two species representing a new genus of the Chaetomiaceae,

Mycologia, 41(3) :346-354. Figs. 1-33. 1949. (The new gonus Lophotrichus.)
Jaczewski, Arthur Louis: Les Chaetomi^es de la Suisse, Bidl. VHerbier Boissier,

3:494-496. 1895.

: Monographie des Cucurbitariees de la Suisse, Bull. soc. Vaudoise des

Sciences Naturelles, Lausanne, 31:67-128. 2A figs. 1895.

LIST 20. SPHAERIALES 697

— : Monographie des Calosphaeriees de la Suisse, Bull. I'Herbier Boissier,
4:78j86. 1896.

Etude monographique de la Famille des Sphaeriacees (Fuckel Jacz.) de la

Suisse, Bull. soc. mycol. France, 12:86-119. PI. 8. 1896.
Seaver, Fred J.: The genus Lasiosphaeria, Mycologia, 4(3) :1 15-124, Ph. 66-67.

1912.
Petrak, F.: tJber Gibbera und verwandte Gattungen, Sydoioia, Ann. Mycol.,

1(4-6): 169-201. 1947. (Contains a key distinguishing the seven closely re-
lated genera Trichodothis, Spilosticta, Coleroa, Parodiella, Pseudoparodia,

Gibbera, and Neogibbera.
Welch, Donald S.: A monographic study of the genus Cucurbitaria in North

America, Mycologia, 18(2):51-86. Pis. 7-S. Figs. 1-5. 1926.
Ramsey, Glen B.: The genus Rosellinia in Indiana, Proc. Indiana Acad. Sci.,

1914:3-16. PZs. 1-3. 1914.
Rick, J.: Monografia das Roselinias Riograndenses, Broteria, Serie Trimestral,

Ciencias Naturais, 1(4) :183-192. 1932.

: Monografia das Valsineas do Rio Grande do Sul, ibid., 2(2):83-99. 1933.

: Monographia Sphaerialium astromaticorum Riograndensium, ibid.,

2(3):133-145, (4):169-201. 1933.
Orton, C. R. : Graminicolous species of Phyllachora in North America, Mycologia,

36(l):18-53. 1944.
Holm, L. : Taxononiical notes on Ascomycetes: I. The Swedish species of the

genus Ophiobolus Reiss sensu Sacc, Svensk Botan. Tidskr. 42(4):337-347.

1 pi. I fig. 1948.
Duces, Paul.: Sur quelques Pleospora d'Auvergne, Bull, trimestr. soc. mycol.

France, 53(2):168-174. 1937.
Wolf, Frederick A., and Ross W. Davidson: Life cycle of Piggotia fraxini

causing leaf disease of ash, Mycologia, 33(5):526-539. Figs. 1-2. 1941, (Con-
tains comparisons of eight species of Mycosphaerella occurring on Fraxinus.)
VON Arx, J. Adolph: Beitrage zur Kenntnis der Gattung Mycosphaerella.

Sydowia, Ann. Mycol., 3(1-6) :38-100. Figs. 1-24. 1949.
FiTZPATRicK, Harry M.: Monograph of the Coryneliaceae, Mycologia,

12(4):206-237, (5):239-267. Pis. 12-18. 1920.
: Revisionary studies in the Coryneliaceae: I. Mycologia, 34(4) :464-488.

Figs. 1-43. 1942; II. The genus Caliciopsis, ibid., 34(5) :489-514. Figs. 1-35.

1942.
: Monograph of the Nitschkieae, ibid., 15(l):23-44, (2):45-67. Pis. 1-7.

1923.

The genus Fracchiaea, ibid., 16(3):101-114. PI. 10. 1924.

Chenantais, J. E.: Notice taxonomique sur le groupe Melanomma, Bull. soc.

mycol. France, 38:88-92. 1922.
: fitudes sur les Pyrenomycetes: VII. Les Lasiosordariees, ibid., 35:68-86,

PL I. Figs. 12-15. 1919.
Shear, C. L.; N. E. Stevens; and R. J. Tiller: Endothia parasitica and related

species, U.S. Dept. Agr. Bull. 380:1-82. Pis. 1-23. Figs. 1-5. 1917.
Verplancke, G. : Herziening van de Belgische soorten van het geslacht Diaporthe

Nitschke, Natuurw. Tijdschr., 24(6-7) :125-156. 1942.
VON HoHNEL, Franz: System der Diaportheen, Ber. deut. botan. Ges., 35 :631-638.

1917. (Including a key to the genera placed in the group by the author.)
: Mycologische Fragmente: CCIL. tJber die Diaporthe-Arten auf Aesculus;

CCL. tJber die Diaporthe-Arten auf Caprifoliaceen; CCLI. tJber die Dia-

698 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

porthe-Arten auf Cornus; CCLII. tjber die Diaporthe-Arten auf Corylus;
CCLIII. tJber die Diaporthe-Arten auf Eichen; CCLIV. tJber die Dia-
porthe-Arten auf Weiden; Ann. MycoL, 16(1-2) :116-122, 1918; CCLXII.
tJber die allantoidsporigen Sphaeriaceen, ibid., 16(1-2) :127-132. 1918.

Wehmeyer, Lewis E.: The genus Diaporthe Nitschke and its segregates, Univ.
Michigan Studies, Scientific Series 9, pp. i-x, 1-349. Pis. 1-18. Ann Arbor,
Univ. Mich. Press, 1933.

: The British species of the genus Diaporthe Nits, and its segregates,

Brit. Mycol. Soc. Trans., 17(4) :237-295. 1933.

: The genus Thyridaria (Pyrenomycetes), Lloydia, 4(4):241-261. 4 pis.

1941.
-: A revision of Melanconis, Pseudovalsa, Prosthecium and Titania, Univ.

Mich. Studies, Scientific Series 14, pp. i-viii, 1-161. Pis. 1-11. Ann Arbor,

Univ. Mich. Press, 1941.
Savulescu, Trian: Une nouvelle espece du genre Paranthostomella et con-
siderations syst^matiques sur les Sphaeriales Pseudostromatae, Arch, rou-

maines -path, exptl. microbioL, 7(l):7-32. Fig. 8. 2 diagrams. 1934.
Jaczewski, Arthur Louis: Monographie des Massari^es de la Suisse, Bull.

I'Herbier Boissier, 2:661-698. 1894.
: Les Xylari^es de la Suisse, Bull. soc. mycol. France, 11:108-137. PL 12.

1895.
Ellis, J. B., and B. M. Everhart: Synopsis of the North American species of

Xylaria and Poronia, /. Mycology, 3(9):97-102, (10):109-113. 1887.
: Synopsis of the Nortli American species of Hypoxylon and Nummularia,

ibid., 4(4-5) :38-44, (7):66-70, (9):85-93, (ll):109-il3. 18SS; 5(l):19-33.

1889.
Lloyd, C. G.: Synopsis of some genera of the larger Pyrenomycetes: Camillea,

Thamnomyces, Engleromyces, Mycological Writings, 5:1-16. Figs. 826-857.

-1919. (This section is paged separately from the remainder of this volume.)
: The larger Pyrenomycetes, second paper, ibid., 5:1-16. Figs. 1444-1460.

1919. (This is also paged separately from the remainder of the volume.)

: Xylaria notes, I, ibid., 5 :1-16. Figs. 1200-1236. 1918. (Paged separately.)

-: Xylaria notes, II, ibid.,5:l-\6. Figs. 1324-1357. 1918. (Paged separately.)

Hawkins, Stacy: Some Xylarias of Indiana, Proc. Indiana Acad. Sci., 35:225-

229. 1925.
VAN der Byl, p. a.: Die Swamfamilie Xylariaceae in die Unie van Suid-Afrika,

Ann. Univ. Stellenbosch, 10A(3):1-10'. 1932.
: South African Xylarias occurring around Durban, Natal, Trans. Roy.

Soc. S. Africa, 9(2):181-183. Pis. 7-8. 1921.
Child, Marion: The genus Daldinia, Ann. Missouri Botan. Garden, 19(4) :429-

496. Pis. 26-33. 1932.
Theissen, Ferdinand: Xylariaceae Austro-Brasilienses: I. Xylaria, Denkschrift

der K. Akad. Wiss., Wicn, Math.-naturw. Klasse, 83 :47-S6. 1 1 pis. 7 figs. 1927.
Rick, J.: Monographia das Hypoxyleas Riograndenses, Broteria, Ser. Botan.,

25(l):21-50. 1931.
: Monographia Bolinearum Riograndensium, ibid., 25(2):65-71. 1931.

(Bolineaceae, near Xylariaceae. Also a key to the genera of Xylariaceae,

including the above family.)
Miller, Julian: British Xvlariaceae, Brit. Mycol. Soc. Trans., 15:134-154.

Pis. 6-7. 1930; 17:125-146. Pis. 4-6. Fig. 1. 1932.
: Notes on Hypoxylon species, I, Ann. cryptogam, exotique, 4(2):72-73.

PL 1. 1931.

LIST 21. HYPOCBEALES 699

— : South African Xylariaceae, Bothalia, 4(2):251-272. 1942.
— : Georgia Pyrenomycetes, II, Mycologia, 33(1):74-81. 1941. (Gives notes
on the nomenclature of Hypoxylon and Nummularia.)

List 21. Hypocreales

(See also Ellis and Everhart, in List 20.)

Seaver, Fred J.: Hypocreales, North American Flora, 3:1-56. 1910.

: The Hypocreales of North America, I, Mycologia, l(2):41-76. Pis. 4-5.

1909; II, Ibid., l(5):177-207. PI. 13. 1909; III, Ibid., 2(2):48-92. Pis. 20-21.

1910; IV, Ibid., 3(5):207-230. Pis. 53-54. 1911.
Ellis, J. B., and B. M. Everhart: Synopsis of the North American Hypocre-

aceae, with descriptions of the species, /. Mycology, 2(3):28-31, (5):49-51,

(6):61-69, (7):73-80, (9):97-99, (10):109-111, (11):121-125, (12):133-137.

1886; 3(1) :l-6. 1887.
Weese, J.: Beitrage zur Kenntnis der Hypocreaceen : I. Mitteilung, Sitz. ber. K.

Akad. Wiss. Wien, Math.-naturw. Klasse, 125 :465-575. PZs. 1-3. Figs. 1-15.

1916.
Fitzgerald, Ina S.: Hypocreales of Iowa, State Univ. Iowa Studies Nat. Hist.,

19(2) -.1-32. Figs. 1-18. 1949.
LoHMAN, Marion L., and Alice J. Watson: Identity and host relation of Nectria

species associated with diseases of hardwoods in the eastern states, Lloydia,

6(2):77-108. Figs. 1-2. 1943. (Keys for cultural distinctions of the species

discussed.)
Seeler Jr., Edgar V.: A monographic study of the genus Thyronectria, /.

Arnold Arboretum Harvard Univ., 21(4) :429-460. Pis. 1-5. 1940.
Fetch, T.: Studies in entomogenous fungi: II. The genera Hypocrella and

Aschersonia, Ann. Royal Botanical Garden of Peradeniya, 7:167-278. Pis. 2-7.

1921.

British Hypocreales, Brit. Mycol. Soc. Trans., 21:243-305. 1932.
Further notes on British Hypocreales, ibid., 25(2):166-178. 1941.
Additional notes on British Hypocreales, ibid., 27(3-4) :148-154. 1944.

Boedijn, K. B. : The genus Podostroma in the Netherlands Indies, Bull. Jardin
Botanique de Buitenzorg, ser. Ill, 13(2) :269-275. Fig. 1. 1934.

: A new species of Podostroma from Africa, Ann. Mycol., 36(4):314-317.

1 fig. 1938. (Includes a key to the nine species accepted by the author.)

Imai, Sanshi: Studies on the Hypocreaceae of Japan: I. Podostroma, Trans.
Sapporo Natural History Soc, 12:114-118. 2 figs. 1932; II. ibid., 14(2):101-
106. 1 fig. 1935. Part II contains a key to and description of species of
Cordyceps parasitic on Elaphomyces.

PoDziMEK, Jan: K monografi ceskych namehi (Claviceps), Casopus Narodniho
Musea. Cast Prirodovedna, 106:16-35. 1932.

Langdon, R. F. : Ergot of native grasses of Queensland, Proc. Roy. Soc. Queens-
land, 54:23-32. 1942 (1943).

Atanasopf, Dimitr: Ergot of grains and grasses, 127 pp. Mimeographed and
distributed by the Office of Cereal Investigations, Bureau of Plant Industry,
U.S. Department of Agriculture. 1920

700 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Massee, George: A revision of the genus Cordyceps, Ann. Botany, 9(33):l-44.

Pis. 1-2. 1S95.
Lloyd, C. G.: Synopsis of the Cordyceps of Australasia, Mycological Writings,

4:1-12. Figs. 611-626. 1915. (Separate pagination.)
Cunningham, G. H. : The genus Cordyceps in New Zealand, Trans, and Proc.

New Zealand Inst., 53:372-382. 4 pis. 8 figs. 1921.
Mains, E. B.: The genera Cordyceps and Ophiocordyceps in Michigan, Proc.

Am. Phil. Soc, 74(4):263-271. 4 pis. 1934.
: Cordyceps from the mountains of North Carolina and Tennessee, J.

Elisha Mitchell Sci. Soc, 55(1):117-129. Pis. 18-21. 1939.
: Cordyceps species from British Honduras, Mycologia, 32(l):16-22.

Figs. 1-2. 1940.

': Species of Cordyceps, ibid., 32(3) :310-320. Figs. 1-2. 1940.

: New and interesting species of Cordyceps, ibid., 39(5) :535-545. Figs. 1-3.

1947.

: Entomogenous fungi, ibid: 40(4) :402-416. Figs. 1-12. 1948. (Mainly

Cordyceps, also Stilbellaceae.)
-: Cordyceps bicephala Berk, and C. australis (Speg.) Sacc. Bull. Torrey

Botanical Club, 76(l):24-30. Figs. 1-4. 1949.
MouREAU, J.: Cordyceps du Congo beige, Mem. Inst. Roy. Colon. Beige, 7(5) :55

pp. 5 pis. (3 colored). 1949.
Imai, Sanshi: On the fungus inhabiting Cordyceps and Elaphomyces in Japan,

Trans. Sapporo Natural History Soc, ll(l):31-37. 1929.
; On a new species of Cordyceps parasitic on Elaphomyces in Japan, Proc.

Imp. Acad. Japan, 10:677-679. 1934.
OvERHOLTS, L. 0.: The genus Cordyceps in central Pennsylvania, Proc. Penna.

Acad. Sci., 12:68-74. Figs. 1-9. 1938.
KoBAYASi, Y.: The genus Cordyceps and its allies, Tokyo Bunrika Daikagu, B,

5(84) :53-260. S7 figs. 1941.
Teng, S. C: Notes on Hypocreales from China, Sinensia, 4:269-298. 1934.

List 22. Dothideales

(For graminicolous species of Phyllachora see Orton in List 20. Also see
Ellis and Everhart in List 20 and some of the references in Lists 23 and 24.)

Theissen, F., und H. Sydow: Die Dothideales, Ann. Mycol., 13(3-6): 149-746.

Pis. 1-6. 1915. (A very fine monograph, including some families subsequently

removed to some of the groups listed below.)
Stevens, F. L., and Nora Dalby: Some Phyllachoras from Porto Rico, Botan.

Gaz., 68(l):54r-59. Pis. 6-8. 1919.
Jaczewski, Arthur Louis: Les Dothideales de la Suisse, Bull. soc. mycol. France,

11:155-195. PZ. 14. 1895.
DoiDGE, Ethel M.: Revised descriptions of South African species of Phyllachora

and related genera, Bothalia, 4(2):421-463. 1942.
VON Hohnel, Franz: Fragmente zur Mykologie: XL Mitteilung, Sitz. ber. K.

Akad. Wiss. (Wien), Math.-naturw. Klasse, 119:617-679. 1910. (Key to

LIST 23. PSEUDOSPHAERIALES 701

genera of the Capnodiaceae and to those Dothideaceae with superficial ascus
stroma.)

List 23. Pseudosphaeriales

Thiessen, F., und H. Sydow: Vorentwvirfe zu den Pseudosphaeriales, Ann.
MycoL, 16(1-2) :l-34. Figs. 1-5. 1918.

VON HoHNEL, Franz: Fragmente zur Mykologie. Attention may be called to the
following papers under this title: IV. Mitteilung (A discussion of Family
Pseudosphaeriaceae), Sitz. her. K. Akad. Wiss. (Wien), Math.-natxirw. Klosse,
116:615-647, 1907; VI. Mitteilung (Discussion of the relationship of
Dothideaceae and Pseudosphaeriaceae to Myriangiaceae, Saccardia, and
Cookellaceae), ibid., 118:275-458. PL 1. Figs. 1-35. 1909.

Theissen, F. : Mykologische Abhandlungen, I-III, Verhandhingen der Zoologisch-
Botanischen Gesellschaft, Wien, 66:296-400. PL 1. Figs. 1-14. 1916. (A discus-
sion of the families Pseudosphaeriaceae and Englerulaceae and of the genus
Physalospora.)

: vStudie iiber Botryosphaeria, Ann. MycoL, 14(5) :297-340. 1 fig. 1916.

Miller, Julian H., and Gwendolyn Burton: Study of Bagnisiopsis species on
the Melastomaceae, Mycologia, 35(3):312-324. Figs. 1-23. 1943. (These
authors consider this genus to belong in the Pseudosphaeriales and not in
the Dothideales or Sphaeriales where it is placed by some authors.)

Petrak, F. : tJber Bagnisiopsis und verwandte Gattungen, Hedwigia, 68:251-290.
1928.

(In addition to the foregoing the following series of papers by G. Arnaud
includes keys to families and genera and in some cases to species which do not
correspond to the classifications followed in this textbook but which might be
distributed in several orders.)

Arnaud, G.: Les Ast^rin^es, Ann. ecole nat. agr. Montpellier, N.S., 16:1-288. Pis.

1-53. Figs. 1-22. 3 maps. 1918.
: Les Asterin^es: II. Etude sur les champignons parasites (Parodiellinacees,

inclus Erysiphees), Awn. epipMjL, 7:1-115. Pis. 1-10. Figs. 1-25. 1921.
: Les Asterin^es: III. Etude sur les champignons parasites (Parodiellinacees,

suite), ibid., 9:1-40. Pis. 1-10. 1923.
: Les Ast^rin^es: IV. Etudes sur la syst^matique des champignons pyr^no-

mycetes, Ann. sci. not. Botan., Xme s6r., 7:643-723. Pis. 1-16. Figs. 1-25.

1925.
: Les Ast^rinees: \. Etudes sur les champignons parasites (Caliciacees,

Hemisph^riac^es, etc.), Ann. epiphyt., 16(5) :235-302. Pis. 1-14. Figs. 1-15.

1930.
: Les Astdrinees: VI. Champignons ast^rinoides de I'Herbier du Museum.

Recueil de Travaux Cryptogamiques d^di^s a Louis Mangin, 8 pp. Pis. 4-5.

3 ^^s., Paris, 1931.
: Les Ast^rin^es, VII, Ann. cryptogam, exotique, 4:74-97. Pis. 2-7. 1931.

702 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

List 24. Hemisphaeriales

(Many of the forms probably more properly placed here are assigned by some
authors to the Erysiphales with which some species appear to possess relationship.
Accordingly List 26 should also be consulted.)

Theissen, F., und H. Sydow: Synoptische Tafeln, Ann. Mycol., 15(6):389-491.
Figs. 1-38. 1917. (This paper contains keys to the families and genera of the
Orders Hemisphaeriales and Myriangiales, and also of the Order Perispo-
riales. It is the culmination of the work reported in part in the papers im-
mediately below.)

: Hemisphaeriales, Ann. Mycol., ll(5):468-469. 1913. (Contains keys to

Family Hemisphaeriaceae.)

: Lembosia-Studien, ihid., ll(5):425-467. PL 20. 1913.

: Ueber Membranstrukturen bei den Microthyriaceen als Grundlage fur

den Ausbau der Hemisphaeriales, Mycolog. Cent., 3(6) :275-286. PI. 1. 1913.
(Contains keys to the families of the order and to the genera of the families
Microthyriaceae and Hemisphaeriaceae.)

: Trichopeltaceae n. fam. Hemisphaerialium, Centr. Bakt. Parasitenk.,

Zweite Abt., 39(23-25) :625-640. 1 pi. 7 figs. 1914.

: Zur Revision der Gattung Dimerosporium, Beihefte Botan. Centr., Zweite

Abt., 29:45-73. 1912. (Since the name Dimerosporium is unavailable, the
species for which this name has been used have been distributed among six
or more genera. This paper gives a key to these genera and brief descriptions
of the more available species.)

: Die Gattung Asterina, Abhandlungen der K.K. Zoologisch-Botanischen

Gesellschaft in Wien, 7(3):1-130. Ph. 1-8. 1913. (Gives full descriptions of
119 species in the three sections Euasterina, Dimerosporium, and Clypeol-
aster. Gives a generic key distinguishing Asterina and the 10 other genera of
Microthyriaceae with two-celled ascospores.)

Zur Revision der Gattungen Microthyrium und Seynesia, Oesterreichische

Botan. Z., 62:216-221, 275-280, 327-329, 395-396, 420-435, 1912; 63:121-

131, 1913.
Doidge, Ethel M.: South African Microthyriaceae, Trans. Roy. Soc. S. Africa,

8:235-282. Pis. 13-19. 1920.
: A revision of the South African Microthyriaceae, Bothalia, 4(2) :273-420.

Pis. 1-76. 1942.
Stevens, F. L., and W. H. Manter: The Hemisphaeriaceae of British Guiana

and Trinidad, Botan. Gaz., 79(3) :265-296. Pis. 18-21. 1925.
, and Sister Mary Hilaire Ryan: The Microthyriaceae, Illinois

Biological Monographs, 17(2):1-138, Urbana, Univ. Illinois Press, 1939.
Ryan, Ruth W.: The Microthyriaceae of Porto Rico, Mycologia, 16(4):177-196.

1924. (Mostly consists of new species, some old species being listed but not

described.)
LuTTRELL, E. S.: The genus Stomiopeltis (Hemisphaeriaceae), Mycologia,

38(5) :565-586. Figs. 1-21. 1946.
Petrak, F.: liber die Leptopeltineen, Sydowia, Ann. Mycol., 1(4-6) :232-247.

1947.
Fraser, Lilian: Notes on the occurrence of the Trichopeltaceae and Atichiaceae
in New South Wales, and on their mode of nutrition, with a description of a
new species of Atichia, Proc. Linnean Soc. New South Wales, 61 :277-284. Pis.
13-14. Figs. 1-10. 1936.

LIST 25. ERYSIPHACEAE 703

Tehon, L. R., and G. L. Stout: Notes on the parasitic fungi of Illinois, IV,

Mycologia, 21(4): 180-1 96. PL 13. 1929. (Contains a key distinguishing five

genera of the Stigmateaceae, Order Hemisphaeriales.)
DiPPENAAR, B. J. : 'n Bydrae tot ons kennis van die Suid-Afrikaanse geslagte en

soorte van die Famielie Polystomellaceae Theiss. en Syd., Ann. Univ. Stel-

lenbosch, 8Ai2) -.1-38. Figs. 1-3. 1930.
Theissen, F., und H. Sydow: Die Gattung Parodiella, Ann. Mijcol., 15(1-2) :125-

142. 1917.
Mendoza, Jose Miguel: The Philippine species of Parasterina, Philippine J.

Sci., 49(3) :443-459. Pis. 1-15. 1932.

List 25. Erysiphaceae

(See also in List 23, Arnaud: Les Ast^rin^es, II, and in List 20, Ellis and

EVERHART.)

Salmon, Ernest S.: A monograph of the Erysiphaceae, Mem. Torrey Botan.

Club, 9:1-292. Pis. 1-9. 1900.
: Supplementary notes on the Erysiphaceae, Bull. Torrey Botan. Club,

29(l):l-22, (2):83-108, (4):181-210, (5):302-316, (ll):647-649. Pis. 9-11.

1902.
BuRRiLL, T. J., AND F. S. Earle: Parasitic Fungi of Illinois: II. Erysipheae, Bull.

Illinois State Laboratory of Natural History, 2:387-432. Figs. 1-8. 1887.
Kelsey, F. D.: The genus Uncinula, Oberlin College Laboratory Bull., 7:1-15. 10

^^s. 1897.
O'Kane, W. C.: The Ohio Powdery Mildews, The Ohio Naturalist, 10(7):166-176.

Pis. 2-10. 1910.
Fink, Bruce : Notes on the powdery mildews of Ohio, Ohio J. Sci., 21(6) :211-216.

Figs. 1-2. 1921.
OvERHOLTS, L. 0., AND W. A. Campbell: The powdery mildews of Central

Pennsylvania, Proc. Penna. Acad. Sci., 8:114-124. 1934.
Salmon, E. S.: The Erysiphaceae of Japan, Bull. Torrey Botan. Club, 27(8) :437-

450. PZ. 1. 1900.
J0RSTAD, Ivar: The Erysiphaceae of Norway, Norske Videnskaps-Akad. Math.

naturv. Klasse. Skrifter, 1925(10) :1-116. 2 figs. 1926. (Discusses distribution,

hosts, etc., of the 25 species of the family in Norway.)
Tai, F. L., and C. T. Wei: Notes on Chinese fungi: II. Erysiphaceae, Sinensia.

Contribs. Metropolitan Museum of Natural History, Nanking, 3(4):93-130.

Illustrated. 1932.
: Further studies on the Erysiphaceae of China, Bull. Torrey Botan.

Club, 73(2): 108-1 30. Figs. 1-13. 1946. (Contains key to the Chinese species

of Uncinula.)
Klika, Jaromir : Monografie ceskych padll (Monograph of Czech Erysiphaceae) ,

Spisy Masarykova Akademie Prdce, 23:1-80. Illustrated. 1924.
Savulescu, Trian, und C. Sandu-Ville: Die Erysiphaceen Rumaniens, Ann.

Scientifiques de l' Academic des Hautes JStudes Agronomiques de Bucarest,

1:47-123. Pis. 1-24. 1929. (Also published separately by Tipografia "Buco-
vina" in Bucarest.)

704 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

PoLLACCi, GiNo: Monographia delle Erysiphaceae italiane, Atti reale ist. boian.

Univ. Pavin, 2 ser., 9:155-181. 1 pi. 1905.
Jaczewski, Arthur Louis: Monographie des Erysiphees de la Suisse, Bull.

I'Herbier Boissier, 4:721-755. 1896.
Homma, Yasu: Erysiphaceae of Japan, Journ. Faculty Agr. Hokkaido Imp. Univ.

38(3):183-466. S pis. 1937.
Blumer, S.: Die Formen der Erysiphe cichoracearum DC, Cenlr. Bakt. Para-

sitenk., Zweite Abt., 57(1-3) :45-60. 1922.
: Die Erysiphaceen Mitteleuropas mit besonderer Berucksichtigung der

Schweiz, Beitrdge zur Kryptoganienflora der Schweiz, 7(1): 1-483. Figs.

1-167. 1933.
LiNDER, David H.: A new species of Phyllactinia, Mycologia, 35(4) :465-468.

Figs. 1-5. 1943. (Gives a comparative discussion of the five species of the

genus.)
Hashioka, Yoshio: Specialization in Sphaerotheca fuliginea (Schlecht.) Poll.,

Ann. Phytopathological Soc. Japan, 8:113-123. 1 fig. 1938. (Contains a key to

six biological species based upon spore size and host inoculations.)
Maurizio, Anna Maria: Zur Biologie und Systematik der Pomaceen bewohn-

enden Podosphaeren. Mit Beriicksichtigung der Frage der Empfanglichkeit

der Pomaceenpropfbastarde fur parasitische Pilze, Centr. Bakt. Parasitenk.,

Zweite Abt., 72(1-7) :129-14S. Figs. 1-6. 1927.
Reed, George M.: The powdery mildews — Erysiphaceae, Trans. Am. Microscop.

Soc, 32(4) -.219-258. Pis. 13-16. 1913.

List 26. Meliolaceae (Perisporiaceae)

(For some genera formerly included here see Lists 24 and 27, also Ellis and
Everhart in List 20.)

Gaillard, Albert: Contribution a I'etude des champignons inf^rieurs. Famille

des Perisporiac^es. Le genre Meliola: anatomie, morphologie, syst&natique.

These, 164 pp. 24 pis. Paris, P. Klincksieck, 1892.
: Le genre Meliola. Supplement I, Bull. soc. mijcol. France, 8:176-188. Pis.

14-16. 1892.
Stevens, F, L. : The genus Meliola in Porto Rico, Illinois Biological Monographs,

2 :475-554. FZs. 1-5. 1916.
: Spegazzinian Meliola types, Botan. Gaz., 64(5) :421-425. Pis. 24-26. 1917.

(Illustrations of the type specimens of the species of Meliola described,

mainly from Argentina, by Dr. Carlos Spegazzini.)
: The Meliolineae, I, Ann. MycoL, 25(5-6) :405-469. Pis. 1-2. 1927; II,

ibid., 26(3-4): 165-383. Pis. 2-6. 1928.

AND L. R. Tehon: Species of Meliola and Irene from British

Guiana and Trinidad, M ijcologia, 18(l):l-22. Pis. 1-2. 1926.
Beeli, M.: Note sur le genre Meligla, Bull. Jardin Botanique de I'Etat, Bruxelles,

7(1):89-160. 1920. (Contains a key to all known species of Meliola.)
: Notes Mycologiques: I. Contribution a la flore mycologique du Congo,

ibid., 8(1):1-11. PI. 1. 1922. (Descriptions of additional species of Meliola

and of other fungi.)

LIST 27. REMAINDER OF ERYSIPHALES : CAPNODIACEAE, ETC. 705

Spegazzini, Carlos: Revision de las Meliolas Argentinas, Andes do Museo

Nacional de Historia Natural Buenos Aires, 32:339-393. 1924.
Deighton, F. C: West African Meliolineae: I. Meliolineae on Malvaceae and

Tiliaceae, Mycological Papers. Commonwealth Mycological Institute, 9:1-24.

28 figs. 1944.
Hansford, C. G., and F. C. Deighton: West African Meliolineae: II. Melio-
lineae collected by F. C. Deighton, Mycological Papers. Commonwealth

Mycological Institute, 23:1-79. 1948.
Doidge, E. M., and H. Sydow: The South African species of the Meliolineae,

Bothalia, 2(2) :424-472. 1928.
: South African Perisporiaceae, I, Trans. Roy. Soc. S. Africa, 5:713-750.

Pis. 55-66. 1917; II, Revisional Notes, ibid., 7:193-197. 3 figs. 1919; III,

ibid., 8:107-143. 1920.
Martin, George : Synopsis of the North American species of Asterina, Dimero-

sporium and Meliola, /. Mycology, 1(11) :133-139, (12):145-148. 1885.
Miller, Vera Mentzer, and Lee Bonar: A study of the Perisporiaceae, Capno-

diaceae and some other sooty molds from California, Univ. Calif. Pubs.

Botany, 19(12) :405-428. Pis. 67-70. 1941.
Hansford, C. G.: Chinese fungi collected by S. Y. Cheo, Farlowia, 3(3):269-283.

Figs. 1-18. 1948.
■ , AND M. J. Thirumalachar: Fungi of South India, ibid., 3(3):285-314.

Figs. 1-37. 1948. (These two papers treat mainly of the Meliolaceae, and the

first paper also of a few Hemisphaeriales.)
Yamamoto, Wataro: Formosan Meliolineae, Trans. Natural History Soc. For-
mosa. 30(200-201) :148-1 59. 1940; 30(206-207) :414-425. Figs. 1-36. 1940;

31(208): 14-30. Figs. 1-40. 1941; 31(209) :47-60. Figs. 1-43. 1941.

List 27. Remainder of Erysiphales : Capnodiaceae,
Englerulaceae, Trichothyriaceae, Atichiaceae

(See Also Lists 24 and 26 for Some of the Capnodiaceae)

Theissen, F., und H. Sydow: Synoptische Tafeln, Ann. Mycol., 15(6):389-491.

Figs. 1-38. 1917.
Fraser, Lilian: An investigation of the sooty molds of New South Wales, Proc.

Linnean Soc. New South Wales, 58:375-395. 1933; 59:123-142. Figs. 1-59.

1934; 60:97-118. Figs. 1-65; 159-578. Figs. 1-91; 280-290. Figs. 1-39. 1935.

(Aside from the Capnodiaceae, Englerulaceae, and Atchiaceae, consideration

is given to the Meliolaceae.)
: Notes on the occurrence of the Trichopeltaceae and Atichiaceae in New

South Wales, and on their mode of nutrition, with a description of a new

species of Atichia, ibid., 61:277-284. Pis. 13-14. Figs. 1-10. 1936.
Fisher, Eileen E.: A study of Australian "sooty moulds," Ann. Botany, N.S.,

3(10):399-426. PZ. 12. Figs. 1-4. 1939.
Jaczewski, Arthur Louis: Les Capnodi^es de la Suisse, Bidl. VHerbier Boissier,

3:603-606. 1895.

706 GUIDE TO THE LITERATUKE FOR THE IDENTIFICATION OF FUNGI

Arnaud, Gabriel: Contribution a I'^tude des Fumagines, Ann. ecole nat. agr.

Montpellier, N.S., 9:239-277. Pis. 1-3. Figs. A-C. 1909; 10:211-330. Figs.

1-29. 1910; 12:23-54. Figs. 1-13. 1912.
Cotton, A. D.: The genus Atichia, Roy. Botan. Garden, Kew. Bull. Misc. Inform.,

1914:54-63. Figs. 1-5. 1914. (The genus Atichia is known by some French

writers as Seuratia.)
Mangin, L., et N. Patouillard: Les Atichiales, groupe aberrant d'Ascomyc^tes

inf^rieurs,/7omp. Rend., 154(23) :1475-14S1. Figs. 1-2. 1912.
Petrak, F. : tlber Englerula und die Englerulaceen, Ann. Mycol., 26(5-6) :385-

413. 1928.
Theissen, F. : Die Trichothyriazeen, Beihefte Botan. Centr., Zweite Abt., 32(1) :1-

16. PL 1. Figs. 1-3. 1914.
von Hohnel, Franz: Ueber die Trichothyriazeen, Ber. deut. botan. Ges., 35:411-

416. 1917. (Discussion of structure of perithecium and relationship of the

family, and of the composition of the Order Perisporiales.)
Theissen, F. : Mykologische Abhandlungen, I-III, Verhandlungeyi der Zoologisch-

Botanischen Gesellschaft, Wien, 66:296-400. PI. 1. Figs. 1-14. 1916. (Among

other fungi discusses Family Englerulaceae.)

List 28. Myriangiaceae

(See also Theissen und Sydow, 1917, in List 27.)

Petch, T.: Studies in entomxOgenous fungi: V. Myriangium, Brit. Mycol. Soc.

Trans., 9:45-80. Pis. 2-3. Fig. 1. 1924.
Miller, Julian H.: The genus Myriangium in North America, Mycologia,

32(5):587-600. 1940.
VON Hohnel, Franz: Fragmente zur Mykologie: VL Mitteilung, Sitzber. K.

Akad. Wiss. Wien. Math.-naturiv. Klasse, 118:275-452. PI. 1. Figs. 1-35.

1909. (Revision of the Family Myriangiaceae, and of the genus Saccardia and

of the Family Cookellaceae and discussion of their relationships to the

Pseudosphaeriaceae and Dothideaceae.)

(In addition to the foregoing there is a series of papers by Miss Anna E.
Jenkins and A. A. Bitancourt and others on the genus Elsinoe and its conidial
stage, Sphaceloma, to whicli reference should be made pending the appearance in
the future of a monograph of this genus by these two authors.)

Jenkins, Anna E., and A. A. Bitancourt: Revised descriptions of the genera
Elsinoe and Sphaceloma, Mycologia, 33(3):338-340. 1941.

, and : lUustraQoes das doen^as causadas por "Elsinoe" e "Spha-
celoma" conhecidas na America do Sul ate Janeiro de 1936 (Illustrations of
South American "Elsinoe" and "Sphaceloma" diseases known up to
January 1936), Arquiv. inst. bioL, 10(2):31-60. Pis. 1-11. Sao Paulo. 1939.

Bitancourt, A. A., and Anna E. Jenkins: Elsinoe fawcetti, the perfect stage of
the Citrus scab organism, Phytopathology, 26(4) :393-396. Fig. 1. 1936.

, AND : Perfect stage of the sweet orange fruit scab fungus, My-
cologia, 28(5) :4S9 492. Figs. 1-2. 1936.

LIST 30. SACCHAROMYCETALES AND ASPOROGENOUS YEASTS 707

— ; AND •: New discoveries of Myriangiales in the Americas, Proc.

Eighth Am. Sci. Congr., Biol. Sci.: Botany, Washington, 1940, 3:149-172,
1942.

Hansford, C. G.: Contributions toward the fungus flora of Uganda: III. Some
Uganda Ascomycetes, Proc. Linnean Soc. London, 153:4-52. 1940-41. (Has
descriptions of seven new species of Sphaceloma.)

Jenkins, Anna E.; A. A. Bitancourt; and Flora G. Pollack: Spot anthrac-
noses in the Pacific Coast States, /. Wash. Acad. Sci., 36(12) :416-421. Figs.
1-2. 1946.

, AND : Spot anthracnoses in the United States and some island

possessions. Plant Disease Reptr., 31(3):114r-117. 1 map. 1947. (This has no
descriptions but gives the distribution of 23 spot anthracnoses in the United
States, Puerto Rico, Guam, and Hawaii as well as references to their pub-
lished descriptions.)

Thirumalachar, M. J.: Some new Sphaceloma diseases of economic plants in
Mysore, Brit. Mycol. Soc. Trans., 31(1-2) :l-6. Figs. 1-9. 1947.

(In addition to the foregoing consult Mycologia for various articles describing
separate species of Elsinoe (Sphaceloma) in the last eight years.)

List 29. Aspergillales (Plectascales)

(For extensive systematic studies of Aspergillus and Penicillium and closely
related genera see List 49.)

Dodge, Carroll W.: The higher Plectascales, Ann. Mycol., 27(3-4) :145-184.

Pis. 1-2. Figs. 1-2. 1929. (This treats of the Trichocomaceae and Elapho-

mycetaceae.)
Imai, Sanshi: Fourth note on Elaphomyces in Japan, Proc. Imp. Acad.

(Japan), 15:146-147. 1939.
GoiDANicH, Gabriele: II genere di Ascomiceti "Grosmannia" G. Gold., Boll.

staz. patol. vegetate, N.S., 16(l):26-60. PI. 1. Figs. 1-19. 1936.

List 30. Saccharomycetales and Asporogenous Yeasts

Stelling-Dekker, N. M.: Die Hefesammlung des "Centraalbureau voor
Schimmelcultures." Beitrage zu einer Monographic der Hefesorten. Erster
Teil: Die sporogenen Hefen, Verhandelingen der Koninklijke Akademie
van Wetenschappen te Amsterdam. Afdeeling Natuurkunde {Tiveede Sectie),
Deel 28, No. 1, vii + 547 pp. Illustrated. 1931.

LoDDER, J.: Die Hefesammlung des " Centraalbureau voor Schimmelcultures."
Beitrage zu einer Monographie der Hefearten : Teil II. Die anaskosporogenen
Hefe. Erste Halfte, ihid., Deel 32, ix + 256 pp. 114^^s. 1934.

708 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

DiDDENs, H. A., UND J. Lodder: Die Hefesammlung des " Centraalbureau voor

Schimnielcultures." Beitrage zu einer Monographie der Hefearten: Teil II.

Die anaskosporogenen Hefe. Zweite Halfte, xii + 511 pp. 99 figs. Amsterdam,

N.V. Nord-Hollandsche Uitgevers Maatschappij, 1942.
Guilliermond, Alexandre: Recherches eytologiques et taxonomiques sur les

Endomycetees, Rev. gen. hotan., 21:353-391, 401-419. Pis. 12-19. Figs. 1-33.

1909.

■ : Les levures, xii + 565 pp. \%?> figs. Paris, 0. Doin et fils, 1912.

— : The Yeasts. Translation of the foregoing by F. W. Tanner, xix + 424

pp. 163;i^s. New York, Wiley and Sons, 1920.
' : Clef dichotomique pour la determination des levures, 124 pp. Ilhistrated.

Paris, Librairie le Francois, 1928.
-: La classification des levures, An7i. fermentations, 2:474-491, 540-551.

Figs. 1-23. 1936.
Zender, Justin: Sur la classification des Endomycetacees, Bull. soc. hotan.

Geneve, 17:272-302. 1925.
Bedford, C. L. : A taxonomic study of the genus Hansenula, Mycologia,

34(6) :628-649. 1942.
Niokerson, Walter J.: Studies in the genus Zygosaccharomyces : I. Transfer of

pellicle-forming yeasts to Zygopichia, Farloivia, 1(3):469-481. 1944. (With

key to and description of the species of Zygopichia.)
Konokotina, a. G., and N. A. Krasil'nikov: Yeasts of the genus Debaryomyces

Klock. and their distribution in nature, J. Microbiologie de VInstitut Bac-

teriologiqiie Pasteur (Leningrad), 9(1):93-107. 1 pi. 1929. (In Russian.)
Guilliermond, Alexandre: fitude cytologique et taxinomique sur les levures

du genre Sporobolomyces, Bull, trimestr. soc. mycol. France, 43 :245-258. PL

10. Figs. Ij6. 1927.
Derx, H. G.: Etude sur les Sporobolomycetes, Ann. Mycol., 28(1-2) :l-23. PI. 1.

1930.
Verona, 0., e R. Ciferri: Revisione dei lieviti asporogeni del genero Sporo-
bolomyces Kluyver et van Niel, Atti ist. hotan. '^Giovanni Briosi" e lab.

crittogam. Italiano ■imiv. Pavia, ser. IV, 10:241-255. 1938.
Ciferri, Raffael, e Piero Redaelli: Caratteri e posizione sistematica dell'

agente della "malattia di Darling," Histoplasma capsulatum Darling e note

sugli H. farciminosum, H. pyriforme e H. Muris, ibid., ser. IV, 6:247-309.

PL 1. Figs. 1-27. 1935. (Gives keys to the families Nectaromycetaceae,

Histoplasmaceae, and Torulopsidaceae.)
, E : Monografia delle Torulopsidaceae a pigmento rosso, Atti ist.

hotan. rcgia univ. Pavia, ser. Ill, 2:147-303. 8 pis. 1925.
• — , E : Studies on the Torulopsidaceae, Ann. Mycol., 27(3-4) :243-295.

Pis. 4-6. 1929.
, E : Contriljuzione alia sistematica delle Torulopsidaceae, XV-

XXXIII, Arch. MikrohioL, 6:9-72. 1935.
■ : Studi suUe Toruloi)sidaceae-Sui nomi generic! di Torula, Eutorula,

Torulopsis, Cryptococcus e sul nome di gruppo Toruhiceae, Atti ist. hotan.

regia univ. Pavia, ser. Ill, 2:129-142. 1925.
: Morpliological relations of the genera of asporogenous yeasts, Ann.

Mycol., 28(5-6) :372-376. 1930.

E O. Verona: Chiavi analitiche dei lieviti segnalati nell' uve, nei mosti e

nei vini, Mycopathologia, 4(3) :243-24S. 1948.
Harrison, F. C: Cheese Torulae, Trans. Roy. Soc. Can., 21(Sect. 5, part 2):341-

LIST 31. USTILAGINALES (INCLUDING GRAPHIOLACEAE) 709

380. 5 pis. 1927. (Divides the genus Torula into three genera and describes
the species.)
-: A systematic study of some Torulae, ibid., 22:(Sect. 5, part 2):187-225.

5 ph. 1928.
Castellani, Aldo: I miceti della blastomicosi Nord-Americana, Ann. med.

navale e coloniale, 2(516) :239-257. 2 pis. 1929. (All species of Blastomycoides

are described.)
Negroni, P., y C. A. N. Daglio: Sobre el g&iero Nectaromyces, Andes soc. dent.

argentina, 144:484-491. i^tgrs. 1-2. 1947.
Berkhout, Christine Marie : De schimmelgeslachten Monilia, Oidium, Oospora

en Torula, Doctor's Thesis, Univ. Utrecht, pp. 1-77. Pis. 1-14. Scheveningen,

Edauw and Johannissen, 1923.
CoNANT, Norman F.: Studies in the genus Microsporum: I. Cultural studies.

Arch. Dermatol, and SyphiloL, 33:665-683. 1936 (reprint with additions and

change of pagination); II. Biometric studies, ibid., 34:79-89. 1936 (reprint

with additions and change of pagination).
Martin, Donald S.; Claudius P. Jones; K. F. Yao; and L. E. Lee, Jr.: A

practical classification of the Monilias, /. Bact., 34(1):99-128. Pis. 1-3. 1937.

(Deals with the species of Monilia (Candida) parasitic upon Man.)

List 31. Ustilaginales (Including Graphiolaceae)

(See also Plowright in List 32.)

Clinton, George P.: North American Ustilagineae, Proc. Boston Soc. Natural

History, 31:329-529. 1904.
: The Ustilagineae, or smuts, of Connecticut, Connecticut State Geological

and Natural History Survey Bull. 5:1-45. Figs. 1-55. 1905.
Ustilaginales, North American Flora, 7(l):l-82. 1906.

Zundel, George Lorenzo Ingram: Additions and corrections to Ustilaginales,

North American Flora, 7(14):971-1045. 1939.
Cunningham, G. H. : The Ustilagineae, or smuts of New Zealand, Trans. Proc.

N. Zealand Inst., 55:307-433. 7 pis. 1924.
: Third supplement to the New Zealand Uredinales and Ustilaginaceae,

ibid., 56:74-80. 1926.
: Fourth supplement to the Uredinales and Ustilaginales of New Zealand,

ibid., 57:186. 1926.
: Fifth supplement to the Uredinales and Ustilaginales of New Zealand,

ibid., 58:47-50. 1927.
: Sixth supplement to the Uredinales and Ustilaginales of New Zealand,

ibid., 59:491-505. 1928.

Seventh supplement to the Uredinales and Ustilaginales of New Zealand,

ibid., 61:402-418. 1930.
McAlpine, D.: The Smuts of Australia, vii + 288 pp. Pis. 1-56. Figs. 1-15.

Melbourne, Department of Agriculture, Victoria, 1910.
Schellenberg, H. C: Die Brandpilze der Schweiz, Beitrdge zur Kryptogamen-

flora der Schweiz, 3(2): i-xlvi, 1-180. Figs. 1-79. 1911.

710 GUIDE TO THE LITERATUEE FOR THE IDENTIFICATION OF FUNGI

CiFERRi, R.: Prima contribuzione alio studio degli "Ustilaginales," Boll. soc.

botan. Hal, 2-3:46-59. 1924. (Discusses 22 species of Tolyposporium, Enty-

loma, and Melanotaenium.)
: Seconda contribuzione alio studio degli Ustilaginales, Atti ist. botan.

regia univ. Pavia, ser. Ill, l(2):77-97. 1924. (Describes various species of

Tuburcinia and Entyloma.)
: Terza contribuzione alio studio degli Ustilaginales. Alcuni micromiceti

della flora Spagnola e Svizzera, ibid., 2 :7-14. 1925. (Describes several species

of Entyloma and of other genera.)
: Quarta contribuzione alio studio degli Ustilaginales, Ann. Mycol.,

26(1-2) :l-68. PL 1. 1928. (Includes a synopsis of the known species of

Entyloma.)
-: A few interesting North American smuts: I. Revision of the smuts on

Bouteloua spp., Brit. Mycol. Soc. Trans., 18(4) :257-262. 1934.
BuBAK, F.: Die Pilze Bohmens: Teil II. Brandpilze (Hemibasidii), Archiv der

Naturwissenschaftlichen Landesdurchforschung von Bohmen, 15(3):1-81. Figs.

1-24. 1916.
ViEGAS, A. P.: Alguns fungos do Brasil: III. Ustilaginales, Bragantia, 4(12) :739-

762. Pis. 1-10. Figs. 1-4. 1944.
Nagorny, p. I.: Caucasian species of the genus Ustilago Pers"., Zapiski Nautchno-

prikladnikh Otdyelov Tiflisskovo Botanitcheskovo Sada, 5:109-128. Pis. 1-2.

1926. (In Russian.)
: Caucasian species of the genus Urocystis Rabenhorst, ibid., 6:104-108.

1929. (In Russian.)
: Caucasian representatives of the genus Tilletia Tulasne, Vyestnik Tiflis-
skovo Botanitcheskovo Sada, 1926-27(3-4) :89-96. 1927. (In Russian.)
: Caucasian species of the genus Entyloma DeB., Stravopolskaya Stantsia

Zashtchity Rastyenii, 1926:49-52. 1926. (In Russian.)

Caucasian species of the genus Doassansia Cornu, Izvyestia Terskoi

Okruzhnoi Stantsii Zashtchity Rastyenii, 1-2:84-85. 1927. (In Russian.)
GuTNER, L. S.: Golovnevye griby (po materialam A. A. Jachevskovo) (The smut

fungi of the U.S.S.R. after the materials of the late A. A. Jaczewski), 383 pp.

137 figs. Moscow and Leningrad, Lenin Academy of Agricultural Science.

Institute of Plant Protection, 1941. (In Russian.)
KocHMAN, Jozef: Grzby glowniowe polski. Ustilaginales Poloniae, Planta Po-

lonica Materjaly do Flory Polskiej. Wydawane przez Towarzystwo Naukowa

Warszawskie, 4:1-161. 12 pis. 1 fig. 1936. (In Polish.)
Beeli, M.: Notes Mycologiques: II. Relev^ des Ustilagin^es r^colt^es dans le

bassin du Congo, Bull. Jardin Botanique de I'Stat. Bruxelles, 8(1) :12-15. 1922.
: Notes Mycologiques: III. Relev^ des Ustilagin^es d'Afrique et de leurs

botes, i6i(^., 8(1) :16-22. 1922.
Verwoerd, Len: n'Bydrae tot ons kennis van die Suid-Afrikaanse Ustilaginales

of Brandswamme, Ann. Univ. Stellenbosch, 4A(2):l-34. Figs. 1-6. 1926.
MuNDKUR, B. B.: A contribution towards a knowledge of Indian Ustilaginales,

Brit. Mycol. Soc. Tran^., 23(1):86-121. 2 figs. 1939.
: A second contribution towards a knowledge of Indian Ustilaginales, ibid.,

24(3-4) :312-336. 1940.
: Some rare and new smuts from India, Indian J. Agr. Sci., 14(l):49-52.

Figs. 1-2. 1944.
: Fungi of the Northwestern Himalayas. Ustilaginales, Mycologia

36(3):286-292. 1944.

LIST 31. USTILAGINALES (INCLUDING GRAPHIOLACEAE) 711

Yen, Wen-Yu: Premiere note sur quelques Ustilagin^es de Chine, Ann. Crypto.

Exotique, 7:11-18. PZs. 1-2. 1934.
— : Deuxieme note sur quelques Ustilagin^es de Chine, ihid., 7:85-95. Pis.

3-4. Figs. 1-4. 1934.
— : Note sur les Ustilagin^es de Chine, Contrih. Inst. Bot. Nat. Acad. Peiping,

1:165-175. 1934; 3:5-15, 41-58. 1935.

Recherches syst^matiques, biologiques et cytologiques sur les Ustila-

gin^es de Chine, Theses presentees a la Faculty des Sciences de 1' University
de Paris. S^r. A, No. 341. No. d'ordre 365:157-310. Pis. 8-25. Figs. 1-52.
Paris, 1937.

Ling, Lee: Taxonomic notes on Asiatic smuts, I., Sydowia, Ann. Mycol., 3(1-
6):123-134. 1949.

Whetzel, H. H., and F. D. Kern: The smuts of Porto Rico and the Virgin
Islands, Mycologia, 18(3) :1 14-124. PL 16. 1926.

LiRO, J. Ivar: Uber die Gattung Tuburcinia Fries, Ann. univ. Fennicae Aboensis,
Ser. A, 1:1-153. 1922.

: Die Ustilagineen Finnlands, I, Ann. Acad. Sci. Fennicae, Serie A, 17:1-

636. Figs. 1-9. 1924; II, ibid., 42:i-xiii, 1-720. Figs. 1-8. 1 map. 1935-1938.

Zundel, George Lorenzo Ingram: The Ustilaginales of South Africa, Bothalia,
3(3):283-320. 1938.

: Monographic studies on the Ustilaginales attacking Andropogon, My-
cologia, 22(3):125-158. 1930.

Savile, D. B. 0.: a study of the species of Entyloma on North American com-
posites, Can. J. Research, C, 25(3):105-120. 1 pi. 1947.

Garrett, A. 0.: The Ustilaginales or Smuts of Utah, Bull. Univ. Utah, Biological
Series 4(2), 29(9):l-23. Pis. 1-4. 1939. (A list of all smuts known in Utah,
with hosts and distribution, and a host index. A key to the genera and illus-
trations of every genus and of many species.)

Fischer, George W.: The stem smuts of Stipa and Oryzopsis in North America,
Butler Univ. Botan. Studies, 7:25-39. Qjigs. 1945.

— -, and Elisa Hirschhorn: A critical study of some species of Ustilago

causing stem smut on various grasses, Mycologia, 37(2) :236-266. Figs. 1-6.
1945.

and : Observations on certain species of Ustilago on Hilaria,

Stenotaphrum, and Muhlenbergia, ibid., 37(3):318-325. 2 figs. 1945.
Davis, W. H. : Summary of investigations with Ustilago striaeformis parasitizing

some common grasses, Phytopathology, 25(8):810-817. 1935.
Fischer, George W. : Fundamental studies of the stripe smut of grasses (Ustilago

striaeformis) in the Pacific Coast, Phytopathology, 30(2):93-118. Figs. 1-4.

1940.
Hirschhorn, Elisa, y Julio Hirschhorn : Los carbones del maiz en Argentina,

Rev. facultad agron. Univ. nacl. La Plata, 20(2):108-139. Pis. 1-5. 1935.
Jackson, H. S. : The Ustilaginales of Indiana, Proc. Indiana Acad. Sci., 1917 : 119-

132. 1918; 1920:157-164. Fig. 1. 1921. (Distribution lists and host index.)
Savulescu, Trian: Contributions a la connaissance des Ustilagin^es de Rou-

manie, Ann. inst. recherches agron. Roumanie, 7:1-86. Pis. 1-35. 1936.
Thirumalachar, M. J.: Species of the genera Doassansia, Doassansiopsis, and

Burrillia in India, Mycologia, 39(5):602-611. Figs. 1-9. 1947.
Fischer, Edward: Weitere Beitrage zur Kenntnis der Gattung Graphiola, Ann.

Mycol, 20(3-4) :228-237. Figs. 1-4. 1922. (Distinguishes all the recognized

species of the genus.)

712 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

List 32. Uredinales

General Works

Sydow, p. et H.: Monographia Uredinearum seu specierum omnium ad hunc
usque diem descriptio et adumbratio systematica, Leipzig, Gebrlider Born-
traeger. Vol. 1, pp. i-xxxv, 1-972. 45 pZs. 1904 (The genus Puccinia); vol. 2,
pp. i-xix, 1-396. 14 pis. 1910 (The genus Uromyces) ; vol. 3, pp. 1-728. 32 pis.
1915 (The remainder of the Pucciniaceae and families Melampsoraceae,
Zaghouaniaceae and Coleosporiaceae) ; vol. 4, pp. i-iv, 1-670. 1924 (Ured-
ineae Imperfecti: Peridermium, Aecidium, Monosporidium, Roestelia, Cae-
oma, Uredo, Mapea).

Klebahn, H.: Die wirtswechselnden Rostpilze, xxxvii + 447 pp. Pis. 1-6. Berlin,
Gebrlider Borntraeger, 1904.

Hariot, Paul: Les Uredin^es (Rouilles des plantes), xv + 392 pp. 47 figs. Paris,
Octave Doin, 1908. (This is the first volume issued of the Bibliothcque de
Botanique Cryptogamique directed by L. Mangin, one of the series of
Toulouse, Encyclopedie Scientifique.)

Guyot, a. L.: Les Uredin^es (ou rouilles des vegetaux). fitude morphologique et
biologique des champignons de ce groupe, qui vivent en Europe, Asie Oc-
cidentale, Afrique Septentrionale et revision des especes connues dans les
autres parties du monde: Tome I. Genre Uromyces (a) Especes parasites des
plantes appartenant aux families des Graminees, Cyperacees, Juncacees,
Renonculac^es, Polygonac^es, Ombelliferes, Campanulacees, in Encyclopedie
Mycologique, 8:1-439. Figs. 1-83. Paris, Paul Lechevalier, 1938.

, ET AL.: Uredineana: Tome 1, pp. 1-205, Figs. 1-11, Encyclopedie My-
cologique, 8 (suppl.), 1939; Tome 2, pp. 1-228, Figs. 1-5, ibid. vol. 13, 1946.

Arthur, Joseph Charles: Manual of the rusts in the United States and Canada.
Illustrations by George B. Cummins, xv + 438 pp. 4^7 figs. 1 map. Lafayette,
Ind., Purdue Research Foundation, 1934.

Plo WRIGHT, Charles B. : A monograph of the British Uredineae and Ustilagineae,
vii -f 347 pp. Ph. 1-8. Figs. 1-13. London, Kegan Paul, Trench and Co.,

1889.
Grove, W. B.: The British rust fungi (Uredinales): their biology and classifica-
tion, xi -f 412 pp. 290 figs. Cambridge, Cambridge Univ. Press, 1913.
ViEGAS, A. P.: Alguns fungos do Brasil: IV. Uredinales, Bragantia, 5(1):1-144.

Pis. 1-48. Figs. 1-89. 1945.
Fischer, E.: Die Uredineen der Schweiz, Beitrdge zur Kryptog amen flora der

Schweiz, 2(2):i-xciv, 1-591. Figs. 1-342. 1909.
Cunningham, G. H.: The rust fungi of New Zealand together with the biology,

cytology and therapeutics of the Uredinales, xx + 261 pp. 177 figs. Dunedin,

N. Z., Mclndoc, 1931.
• : The Uredinales, or rust fungi, of New Zealand: I. Pucciniaceae, tribe

Puccineae, Trans. Proc. N. Zealand Inst., 54:619-704. 1 pi. 1923.
■ : The Uredinales, or rust fungi, of New Zealand: Supplement to Part I;

and Part II, ibid., 55:1-58. 1924.

: Second supplement to the Uredinales of New Zealand, ibid., 55:392-396.

1924. (For the third to seventh supplement see List 31 under Cunningham:
The Ustilagineae of New Zealand.)
McAlpine, D.: The Rusts of Australia, vii + 349 pp. 55 pis. 28 figs. Melbourne,
Department of Agriculture, Victoria, 1906.

LIST 32. UREDINALES 713

BuBAK, Franz: Die Pilze Bohmens: Erster Teil. Rostpilze (Uredinales), Archiv

der Nat'urwissenschaftlichen Landesdurchforschung von Bohmen, 13(5): 1-234.

Figs. 1-59. 1908.
Savulescu, Trian et Olga: Materiaux pour la flore des Uredinees de Rounianie,

Academic Romdna Sectimied Stiinfificd Memoriile, 17:114-261. 18 figs.

1941-42 (1943).
: Materiaux pour la flore des Uredinees de Rounianie. Supplement, Acad-

emie Roumaine Bull, Sect. Sci., 26(5) :308-332. 2 figs. 1944.
DoiDGE, Ethel M. : A preliminary study of the South African rust fungi, Bothalia,

2(la):l-228. 6 col. pis. 221 figs. 1926.
Fragoso, Romualdo Gonzales: Flora Iberica. Uredales, Museo Nacional de

Ciencias Naturales, Madrid, 1 :i-lxxi, 1-416. Figs. 1-208. 1924; 2 :i-viii, 1-421.

Figs. 1-174. 1925. (The first volume contains the genus Puccinia, alone; the

second volume the other genera of rusts.)
Eraser, W. P. : The rusts of Nova Scotia, Proc. Trans. Nova Scotian Inst. Sci.

Halifax, 12(4) :313-443. 1913.
Burrill, T. J.: Parasitic fungi of Illinois: I. Uredinales, Bull. Illinois State

Laboratory of Natural History, 2:141-255. 1885.
Ramsbottom, J.: Some notes on the history of the classification of the Uredinales,

Brit. Mijcol. Soc. Trans., 4(1):77-105. 1913. (Contains keys to the families

and genera and lists of the species occurring in Great Britain.)
YosHiNAGA, Torama, AND Naohide Hiratsuka: a list of Uredinales collected

in the Province of Tosa, Botanical Magazine (Tokyo), 44(528) :627-667. 1930.
J0RSTAD, Ivar: a study on Kamtchatka Uredinales, Skrifter Norske Videnskaps-

Akad. Oslo. I. Mat.-naturv. Klasse, 1933(9) :1-183. Figs. 1-22. 1934. (Keys to

the genera and species and a host index.)
: Uredinales of Northern Norway, ibid., 1940(6) :1-145. 1940. (A list of the

rusts of the districts Nordland, Troms, and Finmark. Host index.)
Camara, Emmanuele de Sousa da; Antonis Lopes Branquinho de Oliveira;

et Carlos Gomes da Luz: Uredales ahquot Lusitaniae, III, Agronomia

Lusitana, 5(4):317-347. 1943.
Arthur, J. C, and George B. Cummins: Phillippine rusts in the Clemens col-
lection, 1923-1926, I, Philippine J. Sci., 59(3) :437-449. Pis. 1-3. 1936.
Gobi, Christian J., and W. Tranzschel: On the rust fungi of St. Petersburg and

some adjacent portions of Estland, Viborg and Novgorod Governments.

Scripta Botanica Horti U Oliver sitatis Petropolitanae, 3(2):65-123. 1891. (In

Russian.)
Kern, F. D.; H. W. Thurston, Jr.; and H. H. Whetzel: Annotated index of the

rusts of Colombia, Mycologia, 25(6):448-503. 1933.
, R. Ciferri, and H. W. Thurston, Jr. : The rust-flora of the Dominican

Republic, Ann. Mycol, 31(1-2) :l-40. 1933.
Sydow, H.: Fungi Venezuelani, Ann. Mycol., 28(1-2) :29-224. 1930. (Uredineae

in pp. 37-52.)
Kern, F. D.; H. W. Thurston, Jr., and H. H. Whetzel: Uredinales, in Myco-

logical exploration in Venezuela, Monograph of the Univ. Puerto Rico, B,

2 :262-303. 1934.
: Additions to the Uredinales of Venezuela, I, Mycologia, 30(5) :537-552

1938.
-, AND H. W. Thurston, Jr.: Additions to the Uredinales of Venezuela,

II-III, ibid., 35(4):434-445. 1943; 36(l):54-64. 1944.
Hiratsuka, Naohide: Zweiter Beitrag zur Uredineen-flora von Sudsachalin,
Trans. Tottori Soc. Agr. Sci., 2(3) :233-246. 1931.

714 GUIDE TO THE LITEEATURE FOR THE IDENTIFICATION OF FUNGI

Hashioka, Yoshio: Mat^riaux pour la flore des Ur^din^es de I'lle de Saghaline
septentrionale, /. Japanese Botany, 12:882-886. 1936. (A list and host names
of 29 species from the Russian portion of Sakhalin.)

Jackson, H. S.: The Uredinales of Indiana, Proc. Indiana Acad. Sci., 1915:429-
475. 1916.

— : The Uredinales of Delaware, ibid., 1917:311-385. 1918.

: The Uredinales of Oregon, Brooklyn Botanic Garden Mem., 1:198-287

1918.
-: The rusts of South America based on the Holway collection, Mycologia,

18(4) :139-162. PL 19. 1926; 19(2) :51-65. 1927; 23(2) :96-116. PL 11. Fi^s. 1-5.

(5):332-364, (6):463-503. 1931; 24(1):62-186. 1932. (Keys to the genus

Mainsia and to the rusts on various groups of hosts.)
Garrett, A. 0.: The Uredinales or rusts of Utah, Bull. Univ. of Utah,

28(7):1-81.PL 1-8. 1937.
HoTsoN, John William: Key to the rusts of the Pacific Northwest, Univ.

Washington Pub. Biology, 3:193. Illustrated. 1934.
Barclay, A.: Descriptive list of the Uredineae occurring in the neighborhood of

Simla (Western Himalaya), /. Asiatic Soc. Bengal, 56:350-375. 4 pis. 1887;

58:232-251. 3 pis. 1889; 59:75-112. 4 pis. 1890.
: Additional Uredineae from the Neighborhood of Simla, ibid., 60:211-230.

2 pis. 1891.
Cummins, George B.: Uredinales of New Guinea, Mycologia, 32(3):359-373.

Figs. 1-14. 1940; 33(l):64r-68. 1 fig. (2):143-154. Figs. 1-7. (4):380-389.

Figs. 1-14. 1941.
Tax, F. L.: Uredinales of Western China, Farlowia, 3(1):95-139. 27 figs. 1947.

M elampsoraceae

Mains, E. B.: Species of Melampsora occurring upon Euphorbia in North
America, Phytopathology, 7(2) :101-105. 1917.

Thirumalachar, M. J., and Frank D. Kern: Notes on some species of Pha-
kopsora and Angiopsora, Mycologia, 41(3):283-290. Figs. 1-3. 1949. (Con-
tains a key for distinguishing the seven nearly related genera of rusts,
Phakopsora, Angiopsora, Bubakia, Baeodromus, Dasturella, Arthuria, and
CeroteUum.)

HiRATSuKA, Naohide: Studies on the Melampsoraceae of Japan, /. Faculty Agr.
Hokkaido Imp. Univ., 21(l):l-42. 2 figs. 1927.

: Notes on the Melampsoraceae of Japan: II. Chrysomyxa of Japan,

Botanical Magazine (Tokyo), 43(513) :466-478. 1929.

: Notes on the Melampsoraceae of Japan: III. Pucciniastrum of Japan,

ibid., 44(521) :261-284. 1930.

: Beitrage zu einer Monographie der Gattung Pucciniastrum Otth., J.

Faculty Agr. Hokkaido Imp. Univ., 21(3):63-119. 1 pi. 1927.

: A contribution to the knowledge of the Melampsoraceae of Hokkaido,

Japanese J. Botany, 3(4):289-322. 1927.

: Additional notes on the Melampsoraceae of Hokkaido, Botanical Maga-
zine (Tokyo), 42(503) :503-504. 1928.

• : Additional notes on the Melampsoraceae of Hokkaido, II, Trans. Tottori

Soc. Agr. Sci., 4(2) :1 11-1 15. 1932.

: Additional notes on the Melampsoraceae of Saghalien, Trans. Sapporo

Natural History Soc, 10:119-121. 1929.
-, and Y. Uemura: On Japanese species of Hyalopsora, Trans. Tottori

Soc. Agr. Sci., 4(l):ll-27. 2 figs. 1932. (In Japanese.)

LIST 32. UREDINALES 715

— : Thekopsora of Japan, Botanical Magazine {Tokyo), 43(505): 12-22. 1929.
— : Phakopsora of Japan, ibid., 49(587) :781-788, (588):853-860. 1935;
50(589) :2-8. 1936.

A monograph of the Pucciniastreae, ix + 374 pp. Pis. 1-11. Tottori,

Japan, 1936. (Reprinted from Mem. Tottori Agr. Coll., 4.)
Faull, Joseph Horace: Taxonomy and geographical distribution of the genus

Milesia, Contribs. Arnold Arboretum Harvard Univ., 2:1-138. Pis. 1-9. 1932.
: Taxonomy and geographical distribution of the genus Uredinopsis, ibid.,

11:1-120. Pis. 1-6. 1938.

Tropical fern hosts of rust fungi, J. Arnold Arboretum Harvard Univ.,

28(3):309-319. 1947.
HiRATSUKA, Naohide, AND Y. Yoshida: Two species of Milesina on some Japan-
ese species of Polystichum, Trans. Tottori Soc. Agr. Sci., 4(1):7-10. 1932.
: On some new species of Milesina, Botanical Magazine (To^yo), 48(565) :39-

47. Figs. 1-6. 1934.
Kamei, Senji: On new species of heteroecious fern rusts. Trans. Sapporo Natural

Histonj Soc, 12(3):161-174. 1932.
Bell, H. P.: Fern rusts of Abies, Botan. Gaz., 77(l):l-30. Pis. 1-5. 1924.
Hunter, Lilian M.: Comparative studies of spermogonia of rusts of Abies,

Botan. Gaz., 83(l):l-23. Pis. 1-4. Figs. 1-2. 1927.
Weir, James R., and Ernest E. Hubert: Observations on forest tree rusts, Am.

J. Botany, 4(6) :327-335. Figs. 1-2. 1917.
Hedgcock, George G., and N. Rex Hunt: Notes on some species of Coleo-

sporium, I, Mycologia, 14(5) :244-257. Pis. 20-21. 1922. (Distinguishes

several species of Coleosporium inhabiting Composites.)
Weir, James R.: The genus Coleosporium in the northwestern United States,

Mycologia, 17(6) :225-239. Pis. 22-24. Fig. 1. 1925.
Hedgcock, George G. : A key to the known aecial forms of Coleosporium occur-
ring in the United States and a list of the host species, Mycologia, 20(2) :97-

100. 1928.
Arthur, J. C, and F. D. Kern: North American species of Peridermium, Bull.

Torrey Botan. Club, 33(8):403-438. 1906.
: North American species of Peridermium in pine, Mycologia, 6(3) :109-138.

1914.
LuDWiG, C. A.: Notes on some North American rusts with Caeoma-like sori,

Phytopathology, 5(5):273-281. 1915. (Gives keys for the determination of

such rusts.)
Rhoads, a. S.; G. G. Hedgcock; E. Bethel; and C. Hartley: Host relation-
ships of the North American rusts other than Gymnosporangium which

attack Conifers, Phytopathology, 8(7) :309-352. 1918.
Mains, E. B.: Angiopsora, a new genus of rusts on grasses, Mycologia, 26(2) :122-

132. Pis. 17-20. 1934.
Weir, James R.: The genus Chrysomyxa, Mycologia, 15(4):183-187. PL 17.

1923.

Pucciniaceae

Dietel, p. : Monographie der Gattung Ravenelia, Beihefte Botan. Centr. Zweite

Abt., 20:343-413. Pis. 5-6. 1906.
Massee, George: Revision of the genus Hemileia, Roy. Botan. Garden, Keiv,

Bull. Misc. Inform., 1906:35-42. 1 pi. 1906.
Milesi, M., e G. B. Traverso: Saggio di una monografia del genere Triphrag-

mium, Ann. Mycol., 2(2):143-156. PL 5. 1904.

716 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

DiETEL, P.: Zur Umgrenzung der Gattung Pileolaria Cast, Ann. Mycol., 19(5-

6):300-303. 1921.
Olive, E. W., and H. H. Whetzel: Endophyllum-like rusts of Porto Rico, Am.

J. Botany, 4(l):44-52. Ph. 1-3. 1917.
MoREAU, M. ET Mme. F. : Les Uredinees du groupe Endophyllum, Bull. soc.

botan. France, 66:13-44. 1919.
Cummins, George B.: The genus Prospodium (Uredinales), Lloydia 3(l):l-78.

Figs. 1-12. 1940.

Gymnosporangium

Kern, Frank D.: A biologic and taxonomic study of tlie genus Gymnospo-
rangium, Bull. New York Botanical Garden, 7:391-483. Pis. 151-161. 1911.

Eriksson, Jakob: Die schwedischen Gymnosporangien, ilir Wirtswechsel und
ihre Spezialisierung, Kgl. Svenska Vetenskapsakad. Handl., 59(6):l-82. Pis.
1-4 (two colored). Figs. 1-13. 1918.

Tanaka, Tyozaburo: New Japanese fungi. Notes and translations, XII, My-
cologia, 14(5) :282-295. 1922. (Discusses the Japanese species of Gymno-
sporangium, with a key.)

Prince, Alton Ernest, and Ferdinand Henry Steinmetz: Gymnosporangium
rusts in Maine and their host relationships, Maine Bull, 42(12) :l-46. 1 map.

1940.
HiRATSUKA, Naohide: Gymnosporangium of Japan, Botanical Magazine (Tokyo),

50(597) :481-488, (598) :549-555, (599) :593-599, (600):661-668. Pis. 8-9.

1936; 51(601) :l-8. 1937.
: Japanese species of Gymnosporangium. Botany and Zoology, 7:748-749.

1939. (In Japanese.) (Eleven species are enumerated and a key provided for

their identification.)

Phragmidium and Kuehneola

Arthur, J. C: Relationship of the genus Kuehneola, Bull. Torrey Botan. Club,

44(11) :501-511. 1912.
HiRATSUKA, Naohide: Kuehneola of Japan, /. Japanese Botany, 12:809-815.

Illustrated. 1936. (List of species and key to their determination.)
Dietel, p.: Ueber die Arten der Gattung Phragmidium, Hedivigia, 44:112-132,

330-346. PI. 4. 1905.
Arthur, J. C: North American rose rusts, Torreya, 9(2):21-28. Figs. 1-3. 1909.
Cummins, George B.: Phragmidium species of North America: differential

teliospore and aecial characters, Mycologia, 23(6) :433-445. PI. 32. 1931.
HiRATSUKA, Naohide: Phragmidium of Japan, Japanese J. Botany, 7(3-4) :227-

299. Pis. 3-4. 1935.
: On some new species of Phragmidium, Trans. Sapporo Natural History

Soc, 13(3):134-138. 1934.

Uromyces and Puccinia and Their Segregates

Holway, E. W. D. : North American Uredineae, Parts I-V.P/.s\ 1-54. Minneapolis,
Published by the author, 1905-1924. (Text and illustrations from micro-
photographs of species of Puccinia on various families of host plants.)

Aref'yef, L. A.: Species of the genus Puccinia in the Baltic Province, 85 pp. St.
Petersburg, 1916. (In Russian.)

— : Species of Uromyces in the Baltic Province, Izvyestia i Trudy Sel'sko-

khozyaistovo otdyelya Rizhskovo Politekhnitchevo Instituta, 3:117-156. 1910.
(In Russian.)

LIST 32. UKEDINALES 717

Pole-Evans, I. B.: The South African rust fungi: I. The species of Puccinia on
Compositae. Trans. Royal Soc. South Africa, 5:637-646. 5 pis. 1916.

BisBY, G. R.: Short cycle Uromyces in North America, Bota7i. Gaz., 69(3). -193-
217. PI. 10. 1920.

Ito, Seiya: On the Uredineae parasitic on the Japanese Gramineae, /. Coll. Agr.
Tohuku Imp. Univ., Sapporo, Japan, 3(2):179-362. Pis. 10-12. 1909.

: Uromyces of Japan, /. Coll. Agr. Hokkaido Imp. Univ., Sapporo, Japan,

11(4) :21 1-287. Pis. 7-9. 1922. (Describes 56 species of Uromyces and 3 of
Pileolaria from Japan; also an index of the rust species and a host index.)
Additional notes on Uromyces in Japan, Botanical Magazine (Tokyo),

40(473) :276-280. A^. 1. 1926.
HiRATSUKA, Naohide: Studies on Uromyces fabae and its related species,

Japanese J. Botany, 6(3):329-379. Pis. 16-17. 1933.
Jackson, H. S.: Carduaceous species of Puccinia: I. Species occurring on the

tribe Vernoniae, Botan. Gaz., 65(4):289-312. 1918.

: New or noteworthy rusts on Carduaceae, Mycologia, 14(3) :104-120. 1922.

Petrak, F. : Beitrage zur Kenntnis der auf Achillea vorkommenden Arten der

Gattung Puccinia, Sydowia, Ann. Mycol., 1(1-3) :44-48. 1947.
Schilling, Max: Die Spezialisierung des Puccinia taraxaci Plow., Sydowia, Ann.

Mycol, 3(1-6) :201-233. Figs. 1-20. 1949.
Gaumann, Ernst A. : Zur Kenntnis einiger Umbelliferen-Puccinien, Ber. schweiz.

botan. Ges., 51:143-164. S figs. 1941.
• , und 0. Jaag: Uber Kleinarten aus dem Formenkreis der Puccinia cam-

panulae, Hedwigia, 75(3):121-129. Figs. 1-3. 1935.
PoEVERLEiN, Hermann: Die Saxifraga-Roste Siiddeutschlands, A7in. Mycol.,

35(1) :53-58. 1937. (Contains a key to the species of Puccinia on Saxifraga in

South Germany.)
Spegazzini, Carlos: Breve nota sobre Uredinales berberidicolas sudamericanas,

Revista Chilena de Historia Natural, Pura y Aplicada, 25 :263-279. 2 pis. 1921.
Cummins, George B.: Revisionary studies of the tropical American rusts of

Panicum, Paspalum and Setaria, Mycologia, 34(6) :669-695. Figs. 1-24. 1942.
: The full-cycle Puccinias on Onagraceae in North America, Am. J. Botany,

19(4) :334-339. A>. 1-4. 1932.
-: New species of Puccinia on Lauraceae from China, Bidl. Torrey Botan.

Club, 76(l):31-38. Figs. 1-12. 1949.
BiSBY, G. R.: The Uredinales found upon the Onagraceae, Am. J. Botany,

3(10):527-561. 1916.
Kern, Frank D,: North American species of Puccinia on Carex, Mycologia,

9(4) :205-238. 1917.
: North American rusts on Cyperus and Eleocharis, ibid., 11(3):134-147.

1919.
-: The microcyclic species of Puccinia on Solanum, ibid., 25(6):435-441.

PI. 48. 1933.
Arthur, J. C: The Uredineae occurring upon Phragmites, Spartina, and Arundi-

naria in America, Botan. Gaz., 34(l):l-20. Figs. 1-4. 1902.
Rees, C. C. : The rusts occurring on the genus Fritillaria, A?n. J. Botany, 4(6) :368-

373. Figs. 1-3. 1917.
Orton, C. R.: North American species of AUodus, Mem. New York Botanical

Garden, 6:173-208. 1916.

: Notes on some Polemoniaceous rusts, Mycologia, 11(4):168-180. 1919.

Dietel, p. : tJber Leptopuccinien auf Artemisia-Arten, Ann. Mycol., 39(2-3) :150-

154. 1941.

718 GUIDE TO THE LITERATUEE FOR THE IDENTIFICATION OF FUNGI

Fahrendorff, E.: Ueber die Brachypuccinia der Artemisia- Arten, Ann. MycoL,

39(2-3) :158-203. Illustrated. 1941.
Lindroth, J. J.: Die Umbelliferen-Uredineen, Acta Societatis pro Fauna et Flora

Fennica, 22:(1). 1922.
HoLWAY, E. W. D.: North American Salvia rusts, J. Mycology, 11(4):156-158.

1905.
Lindquist, Juan C: Las Puccinias parasitas de Geranium en la Repiiblica

Argentina, Notas Mus. La Plata, 13:63-71. 1948.
: Uredineas parasitas de Amarantaceas, en la Republica Argentina, ibid.,

13:243-251. P/s. 1-2. 1948.
HiRATSUKA, Naohide: On the microcyclic species of the Pucciniaceae collected

in some mountains of Japan, Trans. Tottori Soc. Agr. Sci., 3:211-253. 1 pi.

1 fig. 1931. (In Japanese.)
VON Tavel, C: Zur Speziesfrage bei einigen AUium-bewohnenden Uredineen, Ber.

schweiz. botan. Ges., 4(1):123-169. Pis. 1-2. 1932.

List 33. Heterobasidiae

(See also List 34, Bourdot et Galzin, C. Re a, Ramsbottom, Donk; List
35, Burt.)

Martin, G. W.: The Tremellales of the North Central United States and adjacent

Canada, Univ. Iowa Studies in Natural History, 18(3):l-88. Pis. 1-5. 1944.

(Includes Tremellales, Auriculariales and Dacrymycetales.)
Coker, Wm. C: Notes on the lower Basidiomycetes of North Carolina, /. Elisha

Mitchell Sci. Soc, 35(3-4) :113-182. PI. 23 (colored) and 30-67. 1920.
Burt, Edward A. : Some North American Tremellaceae, Dacryomycetaceae and

Auriculariaceae, Ann. Missouri Botan. Garden, 8(4):361-396. PI. 3. Figs.

1-6. 1921.
BouRDOT, H., ET A. Galzin: Hymenomycetes de France: I. H^t^robasidi^s, Bull.

soc. mycol. France, 25:15-36. 1909.
: Heterobasidiae nondum descriptae, ibid., 39:261-266. 1924. (Descrip-
tions of various Tremellaceae and of 10 species of Tulasnella and one of

Gloeotulasnella.)
Neuhoff, Walther: Die Gallertpilze (Tremellineae), Die Pilze Mitteleuropas,

2(la-4a; 7a):l-56. Pis. 1-9 (colored), 1-4 (not colored). Leipzig, Werner

Klinkhart, 1935-1938. Uncompleted (?).
: Die Gallertpilze Schwedens (Tremellaceae, Dacrymycetaceae, Tulasnel-

laceae, Auriculariaceae), Arkiv for Botanik, 28A(l):l-57. Pis. 1-8. Fig. 1.

1936. (Includes descriptive keys to all European species of Exidia and

Dacrymyces.)
Teixeira, a. Ribeiro: Himenomicetos Brasileiros: Auriculariales e Dacrymyce-
tales, Bragantia, 5(2):153-186. Pis. 1-14. 1945.
Viegas, a. p.: Alguns Fungos do Brasil: V. Basidiomycetos-Auriculariales,

Bragantia, 5(3) :1 97-21 2. P/s. 1-4. 2 figs. 1945. (Septobasidium.)
KoBAYAsi, Yosio: On the genus Tremella and its allies from Japan, Tokyo

Bunrika Daigaku Science Repts., B, 4(64-65) :l-26. 6 pis. 15 figs. 1938.

LIST 33. HETEROBASIDIAE 719

— : On the genus Holtermannia of Tremellaceae, ibid., 3(50):75-81. 1 pi.
2 figs. 1937. ^

- — •: Fungorum ordinis Tremellalium Studia Monographica: III. On the
Dacrymyces-group, ibid., 4(70):105-128. 3 pis. 4 figs. 1939.
-: Fungorum ordinis Tremellalium Studia Monographica: IV. On the genera

Femsjonia, Guepinia and Calocera from Japan, ibid., 4(74):215-227. 2 pis. 5

^^5. 1939.
Couch, John N.: Septobasidium in the United States, J. Elisha Mitchell Sci. Soc,

51(l):l-77.P/s. 1-44. 1935.
: The genus Septobasidium, ix + 480 pp. Frontispiece and 114 pis. QO figs.

Chapel Hill, Univ. North Carolina Press, 1938.
Rogers, Donald P.: A taxonomic review of the Tulasnellaceae, Ann. My col.,

3(3):181-203. P/s. 6-7. 1933.
: Some noteworthy fungi from Iowa, Univ. Iowa Studies in Natural History,

15(3):9-29. Pis. 1-3. 1933. (Contains keys to the Iowa species of Sebacina

(incl. Bourdotia) and Heterochaetella.)

Notes on the lower Basidiomycetes, ibid., 17(l):l-43. Pis. 1-3. 1935.

(Includes keys to Ceratobasidium, Botryobasidium, and Sebacina, subgenus

Bourdotia.)
VAN DER Byl, p. a.: Suid-Afrikaanse Dacryomycetaceae, Tremellaceae en

Auriculariaceae, Ann. Univ. Stellenbosch, 1A(S):1-14:. Figs. 1-8. 1923.
McGuiRE, J. M.: The species of Sebacina (Tremellales) of Temperate North

America, Lloydia, 4(l):l-43. Pis. 1-5. 1941.
Rick, J.: Dacryomycetaceae Riograndenses, Broteria. Serie Trimestral. Ciencias

Naturals, 5:74-79. 1936.
Brasfield, T. W.: The Dacrymycetaceae of Temperate North America, Am.

Midland Naturalist, 20(1) :21 1-235. Pis. 1-4. 1938.
BoDMAN, Sister Mary Cecilia: The genus Tremellodendron, ibid., 27(1) :203-

216. Pis. 1-3. 1942.
Baker, Gladys E.: A study of the genus Helicogloea, Ann. Missouri Botan,

Garden, 23(1):69-128. Pis. 7-14. 1936.
: Addenda to the genera Helicogloea and Physalacria, Mycologia,

38(6) :630-638. Figs. 1-25. 1946.
Bj0rnekaer, K.: Floristiske Unders^gelser over danske Baevresvampe (Tremel-
laceae), Friesia, 3(2):l-34. 3 figs. 1944.
BoEDJiN, K. B., ET A. Steinmann: Les especes des genres Helicobasidium et

Septobasidium des Indes N^erlandaises, Bull. Jardin Botanique de Buitenzorg,

Ser. Ill, 11(2):165-219. Pis. 14-18. Figs. 1-31. 1931.
Lloyd, C. G.: The genus Naematelia, Mycological Notes, 7:1149-1150. Figs.

2223-2226. 1922.
Olive, Lindsay S.: Notes on the Tremellales of Georgia, Mycologia, 39(1) :90-108.

Figs. 1-16. 1947.
: Taxonomic notes on Louisiana Fungi, I-II, ibid., 40(l):6-20. Figs. 1-3;

(5):586-604. Figs. 1-3. 1948. (Tremellales.)
Martin, G. W., and Edna E, Huber: Notes on the Tremellales of Iowa, with

keys, U7iiv. Iowa Studies in Natural History, 12(4):91-104. 1 pi. 1927.

720 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

List 34. "Hymenomyceteae" : General Works

Smith, W. G. : Synopsis of the British Basidiomycetes, A descriptive catalogue of

the drawings and specimens in the Department of Botany, British Museum,

531 pp. Pis. 1-5. Figs. 1-145. London, 1908.
Cooke, M. C: lUustrations of British Fungi (Hymenomycetes), Vols. 1-2, 1S81-

1883; 3-4, 1884-1886; 5-6, 1886-1888; 7, 1889-1890; 8, 1889-1891. 1198 coL

pis. in all. London, Williams and Norgate.
Rea, Carleton: British Basidiomyceteae. A Handbook to the Larger British

Fungi, xii + 799 pp. Cambridge, Cambridge Univ. Press, 1922.
: Appendix to British Basidiomyceteae. Additions and corrections, Brit.

Mycol. Soc. Trans., 12:205-230. 1927.
-: Appendix II to British Basidiomyceteae, ibid., 17:35-50. PL 17. 1932.

Pearson, A. A., and R. W. G. Dennis: Revised list of British Agarics and Boleti,
Brit.' Mycol. Soc. Trans., 31(3-4) :145-190. 1948.

Bourdot, H., et a. Galzin: Hymenomycetes de France. Heterobasidi^s-Homo-
basidi^s Gymnocarpes. Contribution a la Flore Mycologique de la France,
vol. 1, iv -i- 761 pp. 185^1^8. Sceaux, France, Marcel Bry, 1927.

Heim, Roger: Les champignons. Tableaux d'un monde Strange, 144 pp. 230 pis.
(from photos). 6 colored pis. Paris, Editions Alpina, 1948.

BucHOLTZ, Fedor V. : Illustrated guide to the fungi of Central Russia : I. Hymeno-
mycetes; II. Agaricaceae. Riga, 1909. (In Russian.)

Lindau, Gustav: Die hoheren Pilze: Basidiomyceten mit Ausschluss der Brand-
und Rostpilze, in Kryptogamenflora flir Anfanger, ed. 3, revised by Eberhard
Ulbrich, vii + 497 pp. 14 pis. Berlin, Julius Springer, 1928.

Ramsbottom, John: A Handbook of the Larger British Fungi, 222 pp. 141 ^grs.
London, Trustees of the British Museum, 1923. (Includes all of the genera
and the more important species of British Basidiomyceteae and fleshy
Ascomyceteae.)

DoNK, M. A.: Revisie van de Nederlandse Heterobasidiomyceteae (uitgez.
Uredinales en Ustilaginales) en Homobasidiomyceteae-Aphyllophoraceae, I,
Mededeel. Nederland. Mycol. Ver., 18-20:67-200. 1931.

: Revision der Niederlandischen Homobasidiomyceteae-Aphyllophoraceae,

II, Mededeel. Botanisch Museum en Herbarium van de Rijks Univ. Utrecht,
9:1-278. 1933.

Karsten, Peter Adolph: Kritisk ofversigt af Finlands Basidsvampar (Basidio-
mycetes; Gastero- und Hymenomycetes), 482 pp. Helsingfors, Finska
Vetenskaps-Societaten, 1889. (There are three supplements, as follows:
Tillagg I: 179-230, 1892; II: 157-186, 1893; III: 3-36, 1898. The whole work
is part of a set of several volumes entitled " Bidrag till Kannedom af Finlands
Natur och Folk.")

MoLLER, F. H.: Fungi of the Faeroes: Part I. Basidiomycetes, 295 pp. 3 colored
pis. 134: figs. 1 colored map. Copenhagen, Einar Munksgaard, 1945.

MoFFATT, Will Sayer: The higher fungi of the Chicago region: I. The Hymeno-
mycetes, Chicago Acad. Sci. Natural History Survey Bidl., 7(1):1-156. Pis.

1-24. 1909.
Graham, Verne Ovid: Muslu-ooms of the Great Lakes region, Chicago Academy

of Science Special Bulletin, 5:i-vii, 1-390. PZs. 1-49. 1944. (Hymenomycetes,

and also Gasteromycetes and fleshy Ascomycetes.)
White, Edward A.: A preliminary report on the Hymeniales of Connecticut,

Connecticut State Geological and Natural History Survey Bull. 3:1-81. Pis.

LIST 35. THELEPHORACEAE AND EXOBASIDIACEAE 721

1-40. 1905. (Keys to the genera of Agaiicaceae, Polyporaceae, Thelepho-
raceae, and Clavariaceae.)

Second report on the Hymeniales of Connecticut, ibid., 15:1-70. PZs.

1-2S. 1910. (Keys to the Connecticut species of Agaricaceae.)
Cleland, John Burton: Toadstools and Mushrooms and Other Larger Fungi of

South Australia: Part I. Introduction and the Toadstools and Mushrooms,

pp. 1-178, col. pis. 1-6, Figs. 1-35. 1934; Part II. Polypores, coral fungi and

the remaining Hymenomycetes and the puff-balls, jelly-like fungi, the larger

Ascomycetes and the Myxomycetes, pp. 179-362. Col. pis. 7-10, Figs.

6-79, 1935. Adelaide, British Science Guild (South Australian Branch),

Government Printer.
DE Laplanche, Maurice C: Dictionnaire iconographique de Champignons

sup^rieurs (Hymenomycetes) qui croissent en Europe, Algerie et Tunisie

suivi des tableaux de concordance de Barrellier, Batsch, Battarra, etc., 544

pp. Paris, Paul Klincksieck, 1894.
GiLLET, C. C: Les champignons (Fungi Hymenomycetes) qui croissent en France.

Description et iconographie, proprietes utiles ou v^neneuses, 828 pp. 738

pis. Paris, J. B. Bailliere et fils, 1878-1890.
BiGEARD, Rene, et Henri Guillemin: Flore des champignons superieurs de

France les plus importants a connaitre (comestibles et veneneux), 600 pp.

56 pis. Chalon-sur-Saone, E. Bertrand, 1909. (Describes 1607 species, mainly

Hymenomycetes and Gasteromycetes, but also Tuberales and Pezizales.)
, ET : Flore des champignons superieurs de France. Complement, ou

Tome II, XX + 791 pp. 44 pis. Paris, Paul Klincksieck, 1913. (2200 species

not included in the foregoing work, but covering the same groups of fungi.)
Qu^LET, Lucien: Flore mycologique de la France et des pays limitrophes, xviii +

492 pp. Paris, Octave Doin, 1888. (Includes Basidiomyceteae and some

Ascomyceteae.)
Velenovsky, J.: Ceske houby, 950 pp. 179 figs. Praz. 1920. (In Czech language —

Bohemian.) (All Basidiomycetes and Discomycetes and Tuberales.)
Singer, Rolf: The Laschia-complex (Basidiomycetes), Lloydia 8(3):170-230.

3 pis. 1 fig. 1945.

List 35. Thelephoraceae and Exobasidiaceae

(See also List 34)

Burt, Edward A.: The Thelephoraceae of North America: I. Thelephora, Ann.
Missouri Botan. Garden, l(2):185-228. Pis. 4-5. 1914; II. Craterellus, ibid.,
1(3) :327-350. Ph. 15-17. 1914; III. Craterellus borealis and Cyphella, ibid.,
1(4) :357-382. PI. 19. 1914; IV. Exobasidium, ibid., 2(3) :627-658. PL 21. 1915;
V. Tremellodendron, Eichleriella and Sebacina, ibid., 2(4):731-770. Pis. 26-
27. Figs. 1-7. 1915; VI. Hypochnus, ibid., 3(2):203-241. Figs. 1-30. 1916;
VII. Septobasidium, ibid., 3(3):319-343. Figs. 1-14. 1916; VIII. Coniophora,
ibid., 4(3) :237-269. Figs. 1-19. 1917; IX. Aleurodiscus, ibid., 5(3):177-203.
Figs. 1-14. 1918; X. Hymenochaete, ibid., 5(4):301-372. Pis. 16-17. Figs.
1-32. 1918; XI. Tulasnella, Veluticeps, Mycobonia, Epithele and Lachno-

722 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

cladium, ibid., 6(4) :253-280. PL 5. Figs. 1-15. 1919; XII. Stereum, ibid.,
7(2-3) :81-248. Pis. 2-6. Figs. 1-48. 1920; XIII. Cladoderris, Hypolyssus,
Cymatella, Skepperia, Cytidia, Solenia, Matruchotia, Microstroma, Proto-
coronospora, and Asterostroma, ibid., ll(l):l-36. PL 1. 1924; XIV. Peiiio-
phora, ibid., 12(3):213-357. 1925; XV. Corticium and supplement to the
whole series, ibid., 13(3) :173-354. 3 figs. 1926.
■: Corticiums causing Pellicularia disease of the coffee plant, hypochnose of

Pomaceous fruits, and Rhizoctonia disease, ibid., 5(2) :1 19-132. Figrs. 1-3.

1918.
Rogers, Donald P., and H. S. Jackson: Notes on the synonymy of some North

American Thelephoraceae and other resupinates, Farlowia, 1(2) :263-328.

1943.
Lloyd, C. G.: Synopsis of the genus Cladoderris, Mycological Writings, 4, Figs.

520-530. 1913. (Separate pagination, pp. 1-11.)
: Synopsis of the stipitate Stereums, ibid., 4, Figs. 531-564. 1913. (Separate

pagination, pp. 14-44.)
CoKER, W. C. : Notes on the Thelephoraceae of North Carolina, /. Elisha Mitchell

Sci. Soc, 36(3-4) :146-196. Pis. 1 and 14-35. 1921.
Emmons, C. W. : The Thelephoraceae of Iowa, Univ. Iowa Studies in Natural

History, 12(4):49-90. 2 pis. 1927.
Lentz, Paul L.: The genus Thelephora in Iowa, Proc. Iowa Acad. Sci., 49:175-

ISi.Figs. 1-11. 1942 (1943).
van der Byl, p. a.: Die Suid-Afrikaanse Thelephoraceae, Ann. Univ. Stellenbosch,

7AiS):l-52. Pis. 1-3. 1929.
Massee, G.: a monograph of the Thelephoraceae, /. Linnean Soc. London,

Botany, 25(170) :107-155. 3 pis. 1890; 27(181-182) :95-205. 3 pis. 1891.
Litschauer, v.: Ueber einige Tomentella-Arten aus Schweden und Macedonien,

Ann. MycoL, 39(4-6) :360-378. 7 figs. 1941.
VON HoHNEL, Franz, und Viktor Litschauer: Beitrage zur Kenntnis der

Corticieen, Sitzber. Kaiserlichen Akad. Wiss. Wien Math.-naturw. Klasse,

115:1549-1620. Figs. 1-10. 1906; 116:739-852. Pis. 1-4. Figs. 1-20. 1907;

117:1081-1124. Figs. 1-10. 1908.
Sartory, a., et L. Maire: Synopsis du genre Fistulina Bull., Paris, 1924.
Lentz, Paul Lewis: Some species of Cyphella, Solenia and Porothelium, Proc.

Iowa Acad. Sci., 54:141-152. Pis. 1-2. 1947 (1948).
Pilat, Albert: Beitrage zur Kenntnis der Thelephoraceen : I. Die Cyphellaceen

Bohmens, Ann. MycoL, 22(1-2) :204-218. PL 1. 1924. (Cyphella and Solenia.)
: Zweiter Beitrag zur Kenntnis der tschechoslowakischen Cyphellaceen,

ibid., 23(1-2) :144-173. Figs. 1-23. 1925.
: Monographia Cyphellacearum Cechosloveniae, I-II, Publications de la

Faculte des Sciences de I'Universite Charles a Prague, Nos. 28-29. 1925.
: Zwei neue Arten der Gattung Cyphella aus der Tschechoslowakei,

Hedwigia, 66:261-264. Fig. 1. 1926.
: Ein kleiner Beitrag zur Kenntnis der Gattung Cyphella Fr. in Tschecho-
slowakei, ibid., 67:113-118. PL 1. 1927.
: Ceskoslovensk6 dfevni houby: I. Stereum Pers., Ceskoslovenskd Akademie

ZemSdelskd Sbornik, 5(3):361-420. 3 pis. 2 figs. 1930. (Detailed descriptions

(in Czech) of the 20 species recognized in Czechoslovakia up to 1930.)
• : Monographie der Mitteleuropaischen Aleurodiscineen, Ann. MycoL,

24(3-4) :203-230. P/. 16. 1926.
: Monographie der europaischen Stereaceen, Hedwigia, 70:10-132. Pis.

1-3. Fig. 1. 1930.

LIST 36. CLAVAKIACEAE 723

Rick, J.: Monographia Thelephoracearum resupinatarum riograndensium, Bro-

teria. Serie Trimestral. Ciencias Naturais, 3:31-48, 66-80, 151-173. 1934.
OvERHOLTS, L. 0.: Mycological notes for 1933, Mycologia, 26(6):502-515. Pis.

54-55. Fig. 1. 1934. (Keys and descriptions of species of Corticium, Section

Botryodea.)
: The genus Stereum in Pennsylvania, Bull. Torrey Botan. Club, 66(8) :515-

537. Pis. 14-18. 1939.
Ito, Tokutaro: Symboles ad Mycologiam Japonicam: I. Aleurodiscus; II. Penio-

phora; III. Corticium, Gloeocystidium et Asterostroma; IV. Asterostromella

et Hymenochaete; V. Hymenochaete, Botanical Magazine (Tokyo),

43(513) :460-466, (514):515-524, (516):633-643, 1929; 44(518) :89-93, (519):

151-157, 1930.
LiTSCHAUER, Viktor: Beitrag zur Kenntnis der Gattung Aleurodiscus (mit

besonderer Beriicksichtigung schwedischer Arten), Ann. Mycol., 42(1-2) :1-

23. 1944.
Rogers, Donald P. : Notes on the Lower Basidiomycetes, Univ. Iowa Studies in

Natural History, 17(l):l-43. Pis. 1-3. 1935. (Ceratobasidium, Botryobasid-

ium, and other segregates of Corticium.)
: The genus Pellicularia (Thelephoraceae), Farlowia, 1(1):95-118. Figs.

1-11. 1943.
: The genera Trechispora and Galzinia (Thelephoraceae), Mycologia,

36(1):70-103. Figs. 1-14. 1944.

List 36. Clavariaceae

Coker, W. C: The Clavarias of the United States and Canada, 209 pp. 92 pis.

Chapel Hill, Univ. North Carolina Press, 1923.
: Further notes on Clavarias, with several new species. /. Elisha Mitchell

Sci. Soc, 63(l):43-69. Pis. 1-14. 1947.
Burt, Edward A.: The North American species of Clavaria with illustrations of

the type specimens. Ann. Missouri Botan. Garden, 9(l):l-78. Pis. 1-11. 1922.
Doty, Maxwell S.: Clavaria, the species known from Oregon and the Pacific

Northwest, Oregon State Monographs. Studies in Botany, 7:1-91. Pis. 1-11.

Figs. 1-9. 1944.
: A preliminary key to the genera of Clavarioid fungi. Bull. Chicago Acad.

.Sa. 8(5): 173-1 78. 1948.

Proposals and notes on some genera of Clavarioid fungi and their types,

Lloydia, 11(2):123-138. 1948.
Cotton, A. D., and E. M. Wakefield: A review of the British Clavariae, Brit.

Mycol. Soc. Trans., 6:164-198. 1918.
Fawcett, Stella G. M.: Studies in the Australian Clavariaceae, Proc. Roy.

Soc. Victoria, N.S., 51:1-20. Pis. 1-5. 1938; 265-280. Pis. 17-23. 1939.
Cool, C: Overzicht van de in Nederland groeiende Clavaria-soorten, Mededeel.

Nederland. Mycol. Ver., 16-17:96-159. Pis. 1-5. 1928.
Imai, Sanshi: On the Clavariaceae of Japan, I, Trans. Sapporo Natural History

Soc, ll(l):38-45. 1929. (English with Japanese summary.)
: On the Clavariaceae of Japan, II, ibid., ll(2):70-77. 1930.

724 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Imai, Sanshi: On the Clavariaceae of Japan, III. The species of Clavaria found
in Hokkaido and Southern Saghalien, ibid., 12(1):9-12. 1931.

: On the Clavariaceae of Japan, IV, ibid., 13(4):377-384. 1934.

: On the Clavariaceae of Japan, V. The species found in the Northern

Honshu, ibid., 16(4):209-217. 1941.

On the Clavariaceae of Japan, VI. The species found in the Central

Honshu, ibid., 16(4):244-250. 1941.
VAN DER Byl, Paul A.: Oor enige Suid-Afrikaanse Clavaria-soorte of knots-

swamme, S. African J. Sci., 29:317-323. PL 3. 1932.
Harper, Edward T. : The Clavaria fistulosa group, Mycologia, 10(2) :53-57. Pis.

3-5. 1918.
KiLLERMANN, Seb. : Die Gattungen Typhula und Pistillaria. Kritische Darstellung

und neue Arten, Z. fur Pilzkunde, N.F., 13(4):98-108. 1 fig. 1934.
Remsberg, Ruth E.: Studies in the genus Typhula, Mycologia, 32(l):52-96.

Figs. 1-58. 1940. (Key and descriptions of 16 species.)
Vang, J. : Typhula species on agricultural plants in Denmark, K. Vet.- og Landbo-

hojskole Aarskr. (Copenhagen), 1945:1-46. 18 figs. 1945.
Fetch, T.: British species of Hirsutella, Naturalist, 1932:45-49. 1932.
Linder, David H. : The genus Myxomycidium, Mycologia, 26(4) :332-343. PL 39.

Figs. 1-6. 1934.
Baker, Gladys E.: Studies in the genus Physalacria, Bull. Torrey Botan. Club,

68(5) :265-288. Figrs. 1-105. 1941. (Includes key and descriptions of 13 species.)
: Addenda to the genera Helicogloea and Physalacria, Mycologia, 38(6) :630-

638. Figs. 1-25. 1946.

List 37. Hydnaceae

(See also List 39 for some of the dentate forms among the Polyporaceae. Some
species ascribed to the genus Irpex probably are more correctly placed in that
family.)

Coker, W. C: The Hydnuras of North Carolina, /. Elisha Mitchell Sci. Soc.

34(4):163-197. Pis. 1-29. (1 and 19 colored). 1919.

: Further notes on Hydnums, ibid., 41(3-4) :270-286. Pis. 51-65. 1926.

: New or noteworthy Basidiomycetes, ibid., 55(2):373-387. Pis. 34-44.

1939. (Contains a key to fleshy stipitate species of Hydnum of the Eastern

United States.)
Beardslee, H. C: Notes on the scaly species of the Hydnaceae, Mycologia,

16(6) :255-258. 1924.
Banker, H. J.: A preliminary contribution to a knowledge of the Hydnaceae,

BulL Torrey Botan. Club, 28(4):199-222. 1901.
: A contribution to a revision of the North American Hydnaceae, Mem.

Torrey Botan. Club, 12:99-194. 1906.
: Type studies in the Hydnaceae: I. The genus Manina, Mycologia,

4(5):271-278. 1912; II. The genus Steccherinum, ibid., 4(6) :309-318. 1912;

III. The genus Sarcodon, ibid., 5(1):12-17. 1913; IV. The genus Phellodon,

LIST 38. MERULIACEAE 725

ibid., 5(2):62-66. 1913; V. The genus Hydnellum, ibid., 5(4):194-205. 1913;
VI. The genera Creolophus, Echinodontium, Gloiodon and Hydnodon, ibid.,
5(6):293-29S. 1913; VII. The genera Asterodon and Hydnochaete, ibid.,
6(5):231-234. 1914.

Miller, L. W.: The genera of the Hydnaceae, Mycologia, 25(4):286-302. 1933.
(Gives a key to the recognized genera of this family.)

: The Hydnaceae of Iowa: I. The genera Grandinia and Oxydontia, ibid.,

25(5):356-368. PI. 43. 1933; II. The genus Odontia, ibid., 26(l):13-32. Ph.
2-3. 1934; III. The genera Radulum, Mucronella, Caldesiella and Gloiodon,
ibid., 26(3):212-219. PL 27. 1934; IV. The genera Steccherinum, Auri-
scalpium, Hericium, Dentinuna and Calodon, ibid., 27(4) :357-373. PI. 33.

1935.
Henry, LeRoy K. : A review of the Hydnaceae (Fungi) of Western Pennsylvania,

Ann. Carnegie Museum, 31:19-32. Ph. 1-2. 1948. (Article 3.)
Lloyd, C. G.: The genus Radulum, Mycological Writings, 4, Figs. 961-984. 1917.

(Separate pagination, pp. 1-12.)
BouRDOT, H., ET A. Galzin: Hym^nomycetes de France: V. Hydnees, Bull. soc.

mycol. France, 30:243-280. 1914.
Cejp, K.: Monografie Hydnacei Republiky Ceskoslovensk^. Praze, 1928.
VAN DER Byl, p. a.: Die Suid-Afrikaanse Hydnaceae of Stekelswamme, Ann.

Univ. Stellenbosch, 12A(l):l-9. Figs. 1-11. 1934.
PiLAT, Albert : Revision der zentraleuropaischen resupinaten Arten der Gattung

Irpex Fr., Ann. Mycol, 23(3-6) :302-307. 1925. _
J0RSTAD, Ivar: Norske resupinate Hydnaceer, Friesia, l(l):2-20. 1932.
Wakefield, E. M.: Australian resupinate Hydnaceae, Trans. Proc. Roy. Soc.

Australia, 54:155-158. 1930. (A key to the genera Acia, Grandinia, and

Odontia.)
Bataille, F. : Flore analytique-descriptive des Hydnes terrestres d'Europe, Bull.

trimestr. soc. mijcol. France, 39:201-216. 1924.

List 38. Meruliaceae

(See also some of the more general works on Polyporaceae in List 39 and in
List 34, in some of which the Meruliaceae are not regarded as a separate family.)

Burt, Edward A.: Merulius in North America, Ann. Missouri Botan. Garden,

4(4) :305-362. Ph. 20-22. Figs. 1-39. 1917.
: Merulius in North America. Supplementary Notes, ibid., 6(2):143-145.

1919. ,. „

BouRDOT, H., ET A. Galzin: Hym^nomycetes de France: IX. Merulife, Bull. soc.

mycol. France, 39:96-118. 1923.
Buchwald, N. Fabritius: De danske arter af slaegten INIerulius (Hall.) Fr. med
en saerlig omtale af gruppen Coniophori Fr., Dansk Botanisk Arkiv, 5(6) :1-

46. 1 pi. 1928.
Nikolaieva, T. L.: The genus Merulius in U.S.S.R. Sovietskaia Botanika,
1933(5) :96-l 11. 1933. (In Russian.)

726 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

List 39. Polyporaceae

(The first three references will serve to distinguish the genera in accordance
with the more recent, but not as yet, fully accepted segregations and arrange-
ments. The distinction of species will be found in the references further on in
this list.)

BoNDARZEW, A., UND R. SiNGERi Zur Systematik der Polyporaceen, Ann. Mycol.,

39:43-65. 1941.
CooKE, W. Bridge : A nomenclatorial survey of the genera of pore fungi, Lloydia,

3(2):81-104. 1940.
Singer, Rolf: Notes on taxonomy and nomenclature of the Polypores, Mycologia,

36(l):65-69. 1944.
MuRRiLL, W. A.: Polyporaceae, North American Flora, 9:1-131. 1907-1908.
: Northern Polypores, 64 pp. New York, published by the author, 1914.

(Covers the area of Northeastern United States and Canada included in the

limits for Britton and Brown's Illustrated Flora.)
: Western Polypores, 36 pp. New York, published by the author, 1915.

(Covers the area of Alaska, British Columbia and the Pacific Coast

States.)

: Southern Polypores, 66 pp. New York, published by the author, 1915.

: Tropical Polypores, 113 pp. New York, published by the author,

1915.
: A key to the white and bright-colored sessile Polyporeae of temperate

North America, Torreya, 8(1):14-16, (2):28-29, (6):130-132. 1908.
: Corrections and additions to the Polypores of temperate North America,

Mycologia, 12(l):6-24. 1920.
: Light-colored resupinate Polypores, ibid., 12(2) :77-92, (6) :299-308. 1920;

13(2):83-100, (3):171-178. 1921.
-: Florida resupinate Polypores, ibid., 34(5) :595-596. 1942. (A key to the

resupinate genera found in Florida.)

Lowe, Josiah L.: The Polyporaceae of New York State (except Poria), revised
ed., iV. Y. State Coll. Forestry Syracuse Univ. Tech. Pub. Q0:l-1 28. Illus-
trations 1-2. 1942.

: The Polyporaceae of New York State (The genus Poria), ibid., 65:

1-91. Figs. 1-20. 1946.

: Studies in the genus Poria: II. White and brightly-colored type material,

Lloijdia, 10(l):45-59. 1947.
-: Studies in the genus Poria: IV. Brown type material, ibid., 11(3) :162-170.

Figs. 1-11. 1948.
OvERHOLTS, L. 0., AND J. L. LowE : New species of Poria, Mycologia, 38(2) :202-

212. 2 figs. 1946.
■ : The Polyporaceae of the Middle-Western United States, Wash. Univ.

Studies, 3 (Pt. I, No. l):l-98. P/s. 1-8. 1915.
: The Polyporaceae of Ohio, Ann. Missouri Botan. Garden, 1(1):81-155.

1914.
: The s])ecies of Poria described by Peck, New York State Museum Bull.

205-206:67-166. Pis. 1-23. 1918.
: The species of Poria described by Schweinitz, Mycologia, 15(5) :207-232.

Pis. 21-24. Figs. 1-30. 1923.
: Diagnoses of American Porias, I, ibid., 14(1):1-11. PL 1. Figs. 1-6. 1922;

II, Bull. Torrey Botan. Club, 50(7) :245-253. Pis. 13-14. Figs. 1-3. 1923.
: Diagnoses of American Porias: III. Some additional brown species with

LIST 39. POLYPORACEAE 727

a key to the common brown species of the United States and Canada, My-
cologia, 23(2) :1 17-129. PZs. 12-14. 1931.

The Polyporaceae of Pennsylvania: I. The genus Polyporus, Penna. Agr.

Exp. Sta. Tech. Bull. 298:3-28. Pis. 1-2. 1933; II. The genera Cyclomyces,
Daedalea, Favolus, Fomes, Lenzites and Trametes, ibid., 316:3-16. Figs.
1-12. 1935; III. The genus Poria, ibid., 418:3-64. 1942.

Henry, LeRoy K.: Pore fungi of Western Pennsylvania: I. The more common
small members of the genus Polyporus, Carnegie Museum Botcmij Pamphlet
2:1-16. 4M^s. 1939.

: Pore fungi of Western Pennsylvania: II. The more common larger mem-
bers of the genus Polyporus and some common members of other genera,
ibid. 3:1-15. 37 figs. im2.

: A review of the pileate Polypores of Western Pennsylvania, Ann. Carnegie

Museiim, 28:221-272. Pis. 26-29. 1941. (Article XIII.)

New and noteworthy polypores from Western Pennsylvania, Proc.

Pen7isylvania Acad. Sci., 22:87-93. 1948.
Neuman, J. J. : The Polyporaceae of Wisconsin, Wisconsin Geological and Natural

History Survey Bull. 33, Scientific Series (10):l-206. Pis. 1-25. 1914.
Shope, Paul F.: The Polyporaceae of Colorado, Ann. Missouri Botan. Garden,

18:287-456. Pis. 16-39. 1931.
Baxter, Dow V.: Some Porias from the region of the Lake States, Papers Mich.

Acad. Sci., 6:67-76. Pis. 1-6. 1927; 9:39-46. Pis. 26-29. 1929.
: Some resupinate Polypores from the region of the Great Lakes, ibid.,

15:191-228. Pis. 17-26. 1932; 17:421-439. Pis. 41-50. 1933; 19:305-332.

Pis. 58-65. 1934; 20:273-281. Pis. 55-60. 1935; 21 :243-267. Pis. 30-39. 1936;

22:275-295. Pis. 31-37. 1937; 23:285-305. Pis. 1-9. 1938; 24:167-188. Pis.

1-7. 1939; 25:145-170. Pis. 1-12. 1940; 26:107-121. Pis. 1-7. 1941; 27:139-

161. Pis. 1-11. 1942; 28:215-233. Pis. 1-6. 1943; 29:85-109. Pis. 1-6. 1943

(1944)- 30:175-191. Pis. 1-14. 1944 (1945); 31:117-130. Pis. 1-5. 1945

(1947); 32:189-211. Pis. 1-10. 1946 (1948); 33:9-30. Pis. 1-9. 1947 (1949);

34:41-56. PZs. 1-6. 1948 (1950).
Eriksson, John: The Swedish species of the "Poria vulgaris" group, Svensk

Botanisk Tidskrift, 43(l):l-25. P/s. 1-5. 1949.
Coker, W. C: The United States species of Coltrichia, J. Elisha Mitchell Sci.

sdc, 62(1) :95-107. Pis. 17-22. 1946.
: Notes on Carolina fungi, ibid., 64(2) :287-303. Pis. 37-54. 1948. (Keys

to, and descriptions of, the fleshy stipitate Polypores of North Carolina.)
Nobles, Mildred K.: Studies in forest pathology: VI. Identification of cultures

of wood-rotting fungi, Can. J. Research, C, 26:281-431. Pis. 1-18. 1948.
PiLAT, Albert: Polyporaceae, Monographie des especes europeennes en vue des

especes de I'Asie du Nord, in Atlas des Champignons d'Europe, vol. 3, pt. 1-2,

pp. 1-624. 374 pis. Prag, Kavina et Pilat, 1936-1942.
VAN DER Byl, Paul A. : A contribution to our knowledge of the Polyporaceae of

South Africa, S. African J. Sci., 18:246-293. 1921.
: Descriptions of additional South African Polyporeae, ibid., 21:308-313.

1924.
ToRREND C : Les Polyporac^es du Br^sil, Broteria. Revista de Sciencias Naturaes,

Serie Botdnica, 18:23-43, 121-143. Pis. 1-8. 1920; 20:107-112. 1922; 21:12-

42. Pis. 1-4. 1924.

: Les Polyporac^es stipit^es du Bresil, ibid., 22:5-19. PI. 1. 1926.

■ : Les Poiyporac^es du Brfeil. Le genre Hexagonia, Broteria. Serie Tri-

mestral. Ciencias Naturais, 4:108-120. 1935.
BosE S R.: Descriptions of fungi in Bengal, I, Proc. Indian Assoc. Cultivation

Sci., 4:109-114. Pis. 1-11. 1918; II. ibid., 1918:136-143. Pis. 1-13. 1920.

728 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

BosE, S. R. : Polyporaceae of Bengal, III, Carmichael Medical College, Belgachia,

Bull.l:l-b. 12 vis. 1920.
• •: Polyporaceae of Bengal, IV. Descriptions of some polypores new to

Bengal, ihid. 2, pp. 1-5. 15 p/s. 1921.

: Polyporaceae of Bengal, V, ihid. 3, pp. 20-25. Ph. 1-9. 1927.

: Polyporaceae of Bengal, VI, Proc. Indian Assoc. Cultivation Sci., 1919 :55-

62. Pis. 1-6. 1922.

: Polyporaceae of Bengal, VII, ibid., 1920-21:27-36. Illustrated. 1923.

: Polyporaceae of Bengal, VIII, /. Dept. Sci., Calcutta Univ., 9:27-31.

Illustrated. 1928.

— : Polyporaceae of Bengal, IX, ibid., 9:35-44. Illustrated. '192S.
-: Polyporaceae of Bengal, X, ibid., 11:1-18. Pis. 1-5. 1934.

Lloyd, C. G.: Synopsis of the genus Fomes, Mycological Writings, 4:211-288.

Figs. 570-610. 1915.

: Synopsis of the genus Hexagona, ibid., 3:1-46. Figs. 279-330. 1910.

: Synopsis of the sections Microporus, Tabacinus and Funales of the genus

Polystictus, ibid., 3:49-70. Figs. 336-356. 1910.
: Synopsis of the section Ovinus of Polyporus, ibid., 3 :73-94. Figs. 496-509.

1911.
: Synopsis of the stipitate Polyporoids, ibid., 3:95-208. Figs. 395-500.

1912.
: Synopsis of the section Apus of the genus Polyporus, ibid., 4:291-392.

Figs. 631-706. 1915.
: The tessellate FavoU, Mycological Notes, 7:1156-1157. Figs. 2281-2284.

1922.
EspiNosA BusTOS, Marcial R. : Sobre las especies chilenas del genero Fomes,

Revista Chilena de Historia Natural, Pura y Aplicada, 25:321-343. 7 pis. 3

figs. 1921.
Rick, J.: Polypori riograndenses: I. Mesopodes aut Pleuropodes, Broteria. Serie

Trimestral. Ciencias Naturais, 3(4):180-189. 1934; II. Merisma, ibid., 4:17-

27. 1935; III. Apodes, ibid., 4:84-94. 1935.

: Polysticti riograndenses, ibid., 4:121-138. 1936; 5:171-178. 1936.

Patouillard, N.: Le genre Ganoderma, Bull. soc. mycol. France, 5(3):64-80. Pis.

10-11. 1889.
Haddow, W. R.: Studies in Ganoderma, /. Arnold Arboretum Harvard Univ.,

12(l):25-46. 2 pis. I jig. 1931.
Humphrey, C. J.: A partial revision of the Ganoderma applanatum group with

particular reference to its oriental variants, Philippine J. Sci., 45(4) :483-589.

Pis. l-SQ.Fig. 1. 1931.
, AND SiMEONA Leus: Studics and illustrations in the Polyporaceae, III,

ibid., 49(2):159-184. Pis. 1-12. 1932. (A continuation of the previous paper

on Ganoderma.)
Cunningham, G. H.: The Polyporaceae of New Zealand, Trans. New Zealand

Inst., 58(3) :202-250. 11 pis. 1927.
• : New Zealand Polyporaceae: I. The genus Poria, Bull. Dept. Sci. Industr.

Res. New Zealand 72, 43 pp., 7 pis., 37 figs., 1947; II. The genus Fuscoporia,

ibid. 73, 14 pp., 1 pL, 10 figs., 1948; III. The genus Polyporus, ibid. 74, 39 pp.,

7 pis., 194S; IV. The genus Coriolus, ibid. 75, 10 pp., 2 pis., 1948; V. The

genus Fomitopsis, ibid. 76, 8 pp., 3 pis., 1948; VI. The genus Coltrichia, ibid.

77, 10 pp., 3 pis., 1948. A^II. The genus Inonotus, ibid. 78; 5 pp., 2 pis., 1948;

VIII. The genus Fomes, ibid. 79, 24 pp., 6 pis., 1948. IX. Trametes, Lenzites

and Daedalea, ibid. 80, 10 pp., 4 pis., 1948.

LIST 40. BOLETACEAE 729

Cleland, J. Burton, and Edwin Cheel: Australian fungi: Notes and descrip-
tions: II. The sclerotia-forming polypores of Australia, Trans. Proc. Roy. Soc.
South Australia, 43:11-22. Pis. 1-5. 1919.

, AND : Notes on Australian Fungi: IV. Polyporus, Fomes and

Hexagona, /. Proc. Roy. Soc. Neio South Wales, 51:473-557. 1918.
-, AND L. Rod way: Notes on the genus Poria. I. Roy. Soc. Tasmania: Papers

and Proc, 1928:31-43. 1928; II. The Porias and Poria-like fungi with defi-
nitely coloured hyphae, ibid., 1928:73-86. 1929; III. The Australian Porias
and Poria-like fungi with hyphae not deeply coloured, ibid., 1929:7-24. 1929.
(A continuation of Pt. II.)

Imazeki, Rokuya: Studies in Ganoderma of Nippon, Bull. Tokyo Science Museum,
1:29-52. 15^grs. 1939. (In Japanese with English summary.)

Graff, Paul W.: North American Polypores: I. Polyporus squamosus and its
varieties, Mycologia, 28(2): 154-1 70. 1936; II. Polyporus biennis and its
varieties, ibid., 31(4):466-484. 1939.

Cooke, William Bridge: Oxyporus nobilissimus and the genus Oxyporus in
North America, Mycologia, 41(4):442-455. Figs. 1-10. 1949.

List 40. Boletaceae

(See also some of the references in List 39, Polyporaceae, and R. Singer:
Phylogenie und Taxonomie der Agaricales in List 41.)

MuRRiLL, Wm. a.: Boletaceae, North American Flora, 9:133-161. 1910.

Peck, Charles H.: Boletinus, New York State Museum Bull. 8:74-80. 1889.

: Boletus, iUd., 8:80-150. 1889.

: Strobilomyces, ibid., 8:158-159. 1889.

Gilbert, E. J.: Les Bolets, in Les Livres du Mycologue, vol. 3, 254 pp. 16 pis.

Paris, Librairie E. le Francois, 1931.
Notules sur les Bolets, I. Bull, trimestr. soc. mycol. France, 52 (2) :249-

260. Figs. 1-5. 1936.
-: Notules sur les Bolets, II, ibid., 56(2):120-124. 1940.

Sartory, a., et L. Maire: Les Bolets. Monographic du genre Boletus Dill,

Fascicules 1-2, 512 pp. 3 ?;/s. Nancy, V. Idoux et Cie, 1931.
Kallenbach, Franz: Die Rohrlinge (Boletaceae), in Die Pilze Mitteleuropas,

vol. 1, pts. 1-21, pp. 1-54. Pis. 1-55 {in greater part colored). Leipzig, Werner

Klinkhart, 1926-1941 (?). Uncompleted (?).
Snell, Walter H.: Notes on Boletus, Mycologia, 24(3):334-341. Fig. 1. 1932;

25(3):221-232. 1933; 26(4) :348-359. 1934; 28(l):13-23. (5):463-475. 1936.
: Tentative keys to the Boletaceae of the United States and Canada, Rhode

Island Botanical Club P^Mication, 1:1-25. 1935.

• The genera of the Boletaceae, Mycologia, 33(4) :41 5-423. _ 1941.

New proposals relating to the genera of the Boletaceae, ibid., 34(4) :403-

411. Fig. 1. 1942.

Slipp, Albert W., and Walter H. Snell: Taxonomic-ecologic studies of the
Boletaceae in Northern Idaho and adjacent Washington, Lloydia, 7(l):l-66.
Pis. 1-8. 1944.

730 GUIDE TO THE LITERATUBE FOR THE IDENTIFICATION OF FUNGI

Strauser, M. C: The Boletaceae of Pennsylvania, Proc. Penna. Acad. Sci.,

4:17-24. Pis. 1-2. 1930. (Contains a key to the Pennsylvania species of

Boletaceae.)
Henry, LeRoy K.: A review of the Boletes (fungi) of Western Pennsylvania,

Ann. Carnegie Museum, 30:213-240. Pis. 1-3. 1946. (Article XIII.)
Chiu, Wei-Fan: The Boletes of Yunnan, Mycologia, 40(2):199-231. 1948.
KiLLERMANN, S. : Ueber den Hexenpilz (Boletus luridus Schaff.) und Verwandte,

Kryptogamische Forschungen der Bayerischen Botanischen Gesellschaft, 4 :336-

343. Sfigs. 1919.
LoHWAG, Heinrich: Kritische Bemerkungen zur Luridus-Gruppe, Hedwigia,

63:323-328. 1922.
Smotlacha, Frantisek: Monografie ceskych hub hfibovitych (Boletinei),

Sitzber. kgl. BohmischenGes. Wiss. Math.-naturw. Klasse, 1911(8) :l-73. 1912.
Bonus, Gabor: Von der Gruppe scaber, Acta Mycologica Hungarica, 1(1) :28,

(2):62-65, (3-4) ill 1-1 18. 1944.
Anonymous: A tinorugombak (Boletus-Boletinus-Gyrodon-Strobilomyces) hata-

roz6 kulcsa., ibid., 2(1-2) :50-56. 1945. (Hungarian.) (Key to these 4 genera

and to 47 species of Boletaceae.)
Singer, Rolf: Sur les genres Ixocomus, Boletinus, Phylloporus, Gyrodon et

Gomphidius, Rev. mijcoL, N.S., 3(2):17-53, (3):157-177. PI. 4 (colored). 1938.
: Notes sur quelques Basidiomycetes : IVe Series. 1. Le genre Krombholzia

Karst. ibid., 3(4):187-191. 1938.

The Boletineae of Florida with notes on extralimital species: I. The

Strobilomycetaceae, Farlowia, 2(1):97-141. PI. 1. 1945; II. The Boletaceae
(Gyroporoideae), ibid., 2(2) :223-303. 1945; III. The Boletoideae of Floiida,
Am. Midland Naturalist, 37(1):1-135. Pis. 1-2. 1947; IV. The lamellate
families (Gomphidiaceae, Paxillaceae and J ugasporacesie), Farlowia, 2(4) :527-
567. PZ. 1. 1946.
MuRRiLL, William A.: Florida Boletes, Lloydia, ll(l):21-35. 1948.

List 41. Agaricaceae (in the Broader Sense)

(For the lamellate Boletoid genera see List 40.)

Works (of More or Less Popular Nature) on Edible and Poisonous
Mushrooms.

Atkinson, George F. : Studies of American Fungi. Mushrooms, Edible, Poison-
ous, etc., 275 pp. Pis. 1-76. Figs. 1-223. Ithaca, N. Y., Andrus and Church,
1900.

Hard, M. E.: The Mushroom, Edible and Otherwise, Its Habitat and Time of
Growth, 609 pp. Pis. 1-66. Figs. 1-505. Columbus, Oliio, Ohio Library Co.,
1908.

Gibson, W. H.: Our Edible Toadstools and Mushrooms and How to Distinguish
Them, x -f 337 pp. 30 col. pis. 57 figs. New York, Harper and Brothers, 1895.

Smith, Alexander H.: Common edible and poisonous mushrooms of south-
eastern Michigan, Cranbrook Institute of Science Bull. 14:1-71. Illustrated.
1938.

LIST 41. AGARICACEAE (iN THE BROADER SENSE) 731

Anonymous: Edible and poisonous fungi, ed. 6, Ministry of Agriculture and
Fisheries Bull. 23, 35 pp. 27 col. pis. London, 1945.

RiCHON, Charles, et Ernest Roze: Atlas des champignons comestibles et
v^n^neux de la France et des pays circonvoisins. Text vol., pp. i-xcviii,
1-265. Vol. of pis. pp. i-xii. Pis. 1-72. Paris, Octave Doin, 1888.

Bresadola, G.: Funghi mangerecci e velenosi, ed. 3, 2 vols., 647 pp. 224 pis. 60
figs. Trento Societa Botanica Italiana. 1932, 1934.

Sartory, a., et L. Maire: Les champignons v^ieneux, 251 pp. 10 col. pis. Paris,
1922.

Bavendamm, W. : Wie unterscheide ich die Speisepiize von den Gift- und Bitter-
pilze? Merkbl. Reichsinst. Forst- und Holzw. 7, 59 pp. 38 figs. 1948.

Maublanc, a. : Les champignons comestibles et v^neneux, ed. 2, vol. 2, pp. 108-
144. 96 col. pis. Paris, Paul Lechevalier, 1927.

DujARRic de la Riviere, R., et Roger Heim: Les champignons toxiques.
Caracteres et determination. Toxines, intoxications, th^rapeutique. Aqua-
relles de A. Bessin, 1 vol. in quarto, 60 pp. 8 col. pis., Figs. 1-5. Paris, En-
cyclop^die Medico-chirurgicale Editeur, 1938.

ScHMiDEG, Armand: Mcs experiences d'un quart de siecle et mes experiments
personnels avec des champignons concernant leur comestibilit^. Acta My-
cologica Hungarica, 3(1-4) :33-47, 1946; 4(1-2) :33-52, 1947.

GtJssow, H. T., AND W. S. Odell: Mushrooms and Toadstools. An account of the
more common edible and poisonous fungi of Canada, 274 pp. 128 pis. (2
colored). Ottawa, Division of Botany, Dominion Experiment Farms of
Canada, 1927.

Works on the Taxonomy of the Agaricaceae.

Singer, Rolf. Phylogenie und Taxonomie der Agaricales, Schweizerische Z. fur
Pilzkunde, 17(2):23-28, (3):35-39, (4):52-57, (5):71-73, (6):84-87, (7):97-
101. 1 fig. 1939. (Gives keys to the genera of Boletaceae and Agaricaceae (in
their old sense), based upon the new anatomical and chemical studies.)

: Das System der Agaricales, Ann. Mycol., 34(4-5) :286-378. 1936. (Keys

to the families and genera of the Agaricales (narrow sense) .)

: The Agaricales, Waltham, Mass., Chronica Botanica Company, Pub-
lishers, 1950. (In press.)

Murrill, W. a.: Agaricaceae, North American Flora, 9:162-426. 1910-1916
(keys and descriptions of genera and species of Tribes Chanterellae and
Lactarieae and part of the white-spored Agariceae); ibid., 10:1-348. 1914-
1932 (vol. not completed) (remainder of white-spored forms and the pink- and
brown-spored forms, including Inocybe and Cortinarius by C. H. Kauffman,
and Pholiota and Hypodendron by L. 0. Overholts).

: The Agaricaceae of the Pacific Coast, I, Mycologia, 4(4):205-217. 1912

(white-spored genera); II, ibid., 4(5):231-262. 1912 (white- and ochre-
spored genera); III, ibid., 4(6) :294-30S. PL 77. 1912 (brown- and black-
spored genera).

: The Agaricaceae of Tropical North America, I, Mycologia, 3(l):23-36.

1911 (white-spored genera); II, ibid., 3(2):79-91. 1911 (white-spored genera);
III, ibid., 3(4):189-199. 1911 (white-spored genera); IV, ibid., 3(6):271-282.
1911 (genera with rose-colored spores); V, ibid., 4(2):72-83. 1912 (ochre-
spored genera); VI, ibid., 5(l):18-36. 1913 (ochre-spored genera); VII, ibid.,
10(l):15-33. 1918 (purple-brown- to black-spored genera); VIII, ibid.,
10(2):62-85. 1918 (purple-brown- to black-spored genera).

732 GUIDE TO THE LITEEATTJRE FOR THE IDENTIFICATION OF FUNGI

MuERiLL, W. A.: Dark-spored Agarics: I. Drosophila, Hypholoma and Pilosace,
Mycologia, 14(2) :61-76. 1922; II. Gomphidius and Stropharia,z&id., 14(3) :121-
142. 1922; III. Agaricus, ibid., 14(4):200-221. 1922; IV. Deconica, Atylospora
and Psathyrella, ibid., 14(5) :258-278. 1922; V. Psilocybe, ibid., 15(l):l-22.
1923.

: The rosy-spored Agarics of North America, Brooklyn Botanic Garden

Mem., 1 :334-336. 1918. (Contains a key to the genera of Subtribe Pluteanae.)

Fayod, v.: Prodrome d'une histoire naturelle des Agaricinees, Ann. sci. nat.
Botan., VII Ser., 9:181-411. Pis. 6-7. 1889.

Lange, Jakob E.: Flora Agaricina Danica, 5 vols. Copenhagen, published under
the auspices of the Society for the Advancement of Mycology and the
Danish Botanical Society.
Vol. 1, pp. 1-90. Ph. 1-40. 1935. Amanita, Limacella, Lepiota, Armillaria,

Tricholoma, Clitocybe.
Vol. 2, pp. 1-105. Pis. 41-80. 1936. CoUybia, Marasmius, Mycena, Omphalia,

Pleurotus, Panus, Volvaria, Pluteus.
Vol. 3, pp. 1-96. PZs. 81-120. 1938. Cortinarius, Pholiota, Inocybe, Hebeloma.
Vol. 4, pp. 1-119. Pis. 121-160. 1939. Flammula, Naucoria, Tubaria, Galera,
Bolbitius, Pluteolus, Crepidotus, Paxillopsis, Paxillus, Psalliota, Stro-
pharia, Lacrymaria, Hypholoma, Psilocybe, Panaeolus, Psathyra,
Pseudocoprinus, Coprinus.
Vol. 5, pp. 1-108. PZs. 161-200. Gomphidius, Melanomphalia, Nyctalis, Linia-
cium, Camarophyllus, Hygrocybe, Lactarius, Russula, Cantharellus,
Lentinus, Schizophyllum. (Supplementary notes. Also title pages and
i-xxiv and index for the five volumes.)

Ceruti, Arturo: Fungi analytice delineati iconibus pictis illustrati, vol. 1, pp.
1-276. Pis. 1-35 {in part colored). Torino, 1948. (Latin, English and French.)
(This first volume of the series includes several species of Amanita and of
Russula.)

KtJHNER, R. : Contribution a I'etude des Hym^nomyc^tes et specialement des
Agaricac^es, Le Botanists, 17(1-4) :l-224. 4 pis. S7 figs. 1926.

RoMAGNESi, Henri: Cle pratique pour la determination generique des Agarics
d'Europe, Rev. mijcol., N.S., 2(supplement 1):11-18. 1937.

CosTANTiN, J., ET L. Dufour: Nouvelle flore des champignons pour la determina-
tion facile de toutes les especes de France, ed. 2, 291 pp. 4166 figs. 1 pi. col.
Paris, Paul Dupont, 1895.

Henry, LeRoy K. : Twenty-eight common gilled mushrooms of western Penn-
sylvania, Trillia, 10 :S2-100. PZs. 12-14. 1939.

Heim, Roger: Les Agarics tropicaux a hym^nium tubul^ (Madagascar, Cote
dTvoire, Antilles, Insulinde), Rev. m^jcol., N.S., 10(1-4) :3-61. 4 pis. 32 figs.
1945.

Cleland, J. Burton, and Edwin Cheel: The Hymenomycetes of New South
Wales, Agr. Gaz. New South Wales, 25 :507-515, 885-888, 1045-1049. 2 pis.
1 fig. 1914; 26:325-333. 2 pis. 1915; 27:97-106. 2 pis. 1916. (Apparently all
that appeared. Descriptions are given of the New South Wales species of
Amanita, Amanitopsis, Lepiota, and Armillaria.)

• : Australian Fungi. Notes and Descriptions, I, Trans. Roy. Soc. S. Australia,

42:88-138. Pis. 9-12 (col.). 1918 (brown- purple- and black-spored Agarics) ;
III, ibid., 43:262-315. Pis. 28-29 (col). 1919 (white-spored Agarics, Polypo-
raceae, Hydnaceae, Thelephoraceae, Gasteromyceteae, Ascomyceteae) ; IV,
ibid., 47:58-78. Pis. 1-2 (col.). 1923 (miscellaneous Agaricaceae, Polypo-
raceae, Thelephoraceae, Clavariaceae, Tremellaceae, and Gasteromyceteae) ;

LIST 41. AGARICACEAE (iN THE BROADER SENSe) 733

V, ibid., 48:236-252. 1924 (miscellaneous Agaricaceae, Polyporaceae, Thele-
phoraceae, Gasteromyceteae, and Ascomyceteae) ; VI, ibid., 51:298-306.
1927 (miscellaneous Agaricaceae); VII, ibid., 53:217-222. 1928 (South
Australian species of Cortinarius) ; VIII, ibid., 55:152-160. 1931 (miscellane-
ous Agaricaceae (mainly Amanita and Clitocybe) and new species of Clavaria) .

Krieger, Louis C. C: Field Key to the Genera of the Gill Mushrooms, pp. 1-8.
1 chart. Baltimore, The Norman-Remington Co., 1920.

Kauffman, C. H.: The Agaricaceae of Michigan, Michigan Geological and Bio-
logical Survey Publication 26, Biological ser. 5, Vol. 1, pp. 1-924, Figs. 1-4,
1918; Vol. 2, pp. 1-10, Pis. 1-172, 1918. (Very full descriptions of all species of
Agaricaceae known to occur in Michigan, and in many genera of all species
recognized in Northeastern United States. Illustrated by excellent photo-
graphs.)

Stover, Wilmer G.: The Agaricaceae of Ohio, Proc. Ohio State Acad. Sci.,
5:462-577. 1912.

Smith, Huron H.: Mushrooms of the Milwaukee region, Milwaukee Public
Museum Field Guide No. 1, Botan. ser., 87 pp. 184 ^^s. Colored table. 1931.

Graham, V. 0.: Mushrooms of the Chicago region. Program of the Activities
Chicago Acad. Sci., 4(3) :42-62. Figs. 1-45. 1933.

Imai, Sanshi : Studies on the Agaricaceae of Hokkaido, J. Faculty Agr. Hokkaido
Imp. Univ., Sapporo, 43:1-378. Pis. 1-5. 1938.

Stevenson, John: Hymenomycetes Brittanici. British Fungi (Hymenomycetes) .
Edinburgh, William Blackwood and Sons. Vol. 1, pp. 1-372. Figs. 1-39. 1886
(Agaricus to Bolbitius); vol. 2, pp. 1-336. Figs. 40-103. 1886 (Cortinarius to
Dacrymyces).

Massee, George: British Fungus-Flora. A Classified Text-Book of Mycology.
London, George Bell and Sons. Vol. 1 pp. 1-432, numerous {unnumbered)
figs., 1892 (Basidiomycetes through the purple-spored Agaricaceae); vol. 2,
pp. 1-460, numerous (unnumbered) figs., 1893 (Agaricaceae: Ochrosporae,
Rhodosporae, and part of Leucosporae) ; vol. 3, pp. 1-512, numerous
(unnumbered) figs., 1893 (Remainder of Agaricaceae-Leucosporae : Hypho-
mycetes) ; vol. 4, pp 1-522, numerous (unnumbered) figs., 1895. (Ascomycetes.)

RiCKEN, Adalbert: Die Blatterpilze (Agaricaceae) Deutschlands und der an-
grenzenden Lander, besonders Oesterreichs und der Schweiz, xxiv -\- 480 pp.
Pis. 1-112. Leipzig, Theodor Oswald Weigel, 1915.

■ : Vademecum fiir Pilzfreunde. Zweite vermehrte und verbesserte Auflage,

xxiv + 352 pp. Leipzig, Quelle und Meyer, 1920.

Karsten, p. a.: Rysslands, Finlands och den Skandinaviska halfons Hatts-
vampar, Bidrag till Kdnnedom af Finlands Natur och Folk, 32:1-571, 1879;
37:1-257, 1882.

Michael, Edmund: Fiihrer fiir Pilzfreunde, systematisch geordnet und ganzlich
neu bearbeitet von Roman Schulz, 3 Bande, 144 pp. introductory text. 386
colored plates (with descriptions). Leipzig, Quelle und Meyer, 1927.

Peck, Charles H.: Reports of the State Botanist of New York. (From 1871 up
to about 1913. Descriptions of numerous species of fungi, chiefly Agarics, as
well as monographs of many genera. Many colored illustrations.) Among the
more important are the following:
New York species of Amanita, 33 :38-49. 1880.
New York species of Lepiota, 35:150-164. 1882.
New York species of Psalliota, 36 :41-49. 1883.
New York species of Lactarius, 38:111-133. 1884.
New York species of Pluteus, 38:133-138. 1884.

734 GUIDE TO THE LITERATUKE FOR THE IDENTIFICATION OF FUNGI

New York species of Pleurotus, Claudopus, and Crepidotus, 39 :58-73. 1885.

New York species of Clitopilus, 42 :39-46. 1889.

New York species of Armillaria, 43 :40-45. 1890.

New York species of Tricholoma, 44:38-64. 1891.

New York species of Pluteolus, 46:58-61. 1893.

New York species of Galera, 46 :61-69. 1893.

New York species of Collybia, 49 :32-55. 1896.

New York species of Flammula, 50:133-142. 1897.

New York species of Hygrophorus, 60:47-67. 1907.

New York species of Russula, 60 :67-98. 1907.

New York species of Pholiota, 61:141-158. 1908.

New York species of Lentinus, 62 :42-47. 1909.

New York species of Entoloma, 62:47-58. 1909.

New York species of Inocybe, 63:48-67. 1910.

New York species of Hebeloma, 63:67-77. 1910.

New York species of Hypholoma, 64:77-84. 1911.

New York species of Psathyra, 64:84-86. 1911.

New York species of Clitocybe, 65 :59-89. 1912.

New York species of Laccaria, 65:90-93. 1912.

New York species of Psilocybe, 65:94-105. 1912.

: Cantharellus, New York State Museum Bull. 2:34-43. 1887.

Craterellus, ibid. 2:44-48. 1887.

Oilman, J. C: Illustrations of the fleshy fungi of Iowa, I. The purple-brown
spored Agarics, Proc. Iowa Acad. Sci., 47:83-90. Figs. 1-6. 1940; II. The
white-spored Agarics, ibid., 48:99-115. Figs. 1-14. 1941; III. The black-
spored Agarics, ibid., 49:153-158. Figs. 1-5. 1942; V. The pink-spored Agar-
ics, ibid., 50:159-163. Figs. 1-7. 1943.

Earle, F. S.: The genera of the North American gill fungi, Bull. New York
Botanical Garden, 5(18):373-451. 1909.

: A key to the North American species of Hypholoma, Torreya, 2(2) :22-23.

1902.

: Keys to the North American species of Coprineae, ibid., 2(3) :37-40. 1902.

: A key to the North American genera and species of the Hygrophoreae,

ibid., 2(4):53-54, (5):73-74. 1902.

: A key to the North American species of Russula, ibid., 2(7):101-103,

(8): 117-1 19. 1902.

: A key to the North American species of Lactarius, ibid., 2(9):139-141,

(10):152-154. 1902.

: A key to the North American species of Cortinarius, ibid., 2(11) :169-172,

(12):180-183. 1902.

: A key to the North American species of Stropharia, ibid., 3(2):24-25.

1903.

: A key to the North American species of Lentinus, ibid., 3(3):35-38,

(4):58-60. 1903.

: A key to the North American species of Panus, ibid., 3(6):86-87. 1903.

: A key to the North American species of Pluteolus, ibid., 3(8):124-125.

1903.

— : A key to the North American species of Galera, ibid., 3(9) :134-136. 1903.
-: A key to the North American species of Inocybe, ibid., 3(11):168-170,

(12):183-184. 1903.

Gams, H.: Schliissel fiir die europaischen Familien, Gattungen und wichtigen
Untergattungeu der Agai'icales, Veroffentlichungen der Osterreichischen Myko-
logischen Gesellschaft, 2:24 pp. 1948.

LIST 41. AGARICACEAE (iN THE BROADER SENSE) 735

White-Spored Genera.

DoNK, M.: De geslachten Cantharellus, Craterellus en Dictyolus in Nederland,

Mededeel. Nederland. Mycol. Ver., 16-17:163-183. 1928.
CoKER, W. C: Craterellus, Cantharellus and related genera in North Carolina;

with a key to the genera of gill fungi, /. Elisha Mitchell Sci. Soc, 35(1-2) :24-

28. Pis. 1 {col.)-17. 1919.
Smith, Alexander H., and Elizabeth E. Morse: The genus Cantharellus in

the Western United States, Mycologia, 39(5) :497-534. Figs. 1-13. 1947.
Bataille, Fred:^ric: Flore monographique des Hygrophores, Mem. soc. d' Emu-
lation du Douhs, 8me s^-., 4:131-191. 1909.
Kaufmann, F.: Die in Westpreussen gefundenen Pilze der Gattungen Dermocybe,

Myxacium, Hygrophorus und Nyctalis, Bericht des Westpreussischen Botan-

isch-Zoologischen Vereins, 34:199-233. 1912.
Graham, V. 0.: The genus Hygrophorus in the Chicago region, Trans. Illinois

State Acad. Sci., 23:160-168. 1932.
Smith, Alexander H., and L. R. Hesler: Studies in North American species of

Hygrophorus: I. The subgenus Limacium, Lloydia, 2(l):l-62. Pis. 1-24.

1939; II. ibid., 5(l):l-94. P/s. 1-18. 1942.
CoKER, W. C: The smaller species of Pleurotus in North Carolina, /. Elisha

Mitchell Sci. Soc, 60(l):71-95. Pis. 40-52. 1944.
PiLAT, Albert: Pleurotus Fries, in Atlas des Champignons de I'Europe, Vol. II.

193 pp. 80 pis. 114 text figs. Prag, Kavina et Pilat, 1928.
FoRSTER, Edward J.: Agarics of the United States. Genus Panus, J. Mycology,

4(2-3) :21-26. 1888.
Harper, Edward T.: The species of Lentinus in the region of the Great Lakes,

Trans. Wisconsin Acad. Sci., 20:365-385. Pis. 14-28. 1921.
VAN DER Byl, p. a.: Suid-Afrikaanse Lentinus-Soorte, Ann. Univ. Stellenbosch,

2A{\):l-n. Figs. 1-7.1924.
Malkovsky, Karel M.: tJber die europaischen Arten der. Gattung Panus, Ann.

Mycol, 30(1-2) :10-80. Pis. 1-2. Figs. 1-26. 1932.
Pilat, Albert: Revision der tropischen Lentinus-Arten aus dem Herbar des

Botanischen Museums in Berlin-Dahlem, Ann. Mycol., 34(1-2) :108-140.

1936.
: Monographie des especes europeennes du genre Lentinus Fr., in Atlas des

Champignons de I'Europe, vol. 5, pp. 1-46. 31 pis. Prag, Kavina et Pilat,

1946.
Singer, Rolf: Studien zur Systematik der Basidiomyceten : 1(1). Uber Panus Fr.

und verwandte Gattungen, Beihefte Botan. Centr., Abt. B, 56(1-2) :137-347.

1936. (Gives a key to all the genera of white-spored Agaricaceae with ec-
centric or lateral stipe or with stipe lacking.)
Kauffman, Calvin H.: The genus Armillaria in the United States and its rela-
tionships. Papers Mich. Acad. Sci., 2:53-67. Pis. 5-9. 1923.
Hotson, H. H.: The genus Armillaria in western Washington, Mycologia,

32(6) :776-790. Figs. 1-3. 1940.
Kauffman, Calvin H.: The genus Clitocybe in the United States with a critical

study of all north temperate species, Papers Mich. Acad. Sci., 8:153-204.

7 pis. 1926.
NtJEscH, Emil: Die Trichterlinge. Monographie der Gattung Clitocybe, 279 pp.

St. Gallen, Switzerland, F. Schwald, 1926.
MuRRiLL, W. A. : The genus Clitocybe in North America, Mycologia, 7(5) :256-

2S3. Pis. 164-166. 1915.

736 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Singer, Rolf, and Alexander H. Smith: A monograph of the genus Leuco-

paxillus Boursier, Papers Mich. Acad. Sci., 28:85-132. Pis. 1-8. 1942 (1943).
Metrod, G.: Les Tricholomes, Rev. mycol., N.S., 7(supplement 2):22-50. Figs.

1-86. 1942. (Key to the five genera into which Tricholoma is segregated,

Rhodocybe, Rhodopaxillus, Melanoleuca, Tricholomopsis, and Tricholoma,

and to their species.)

: Les Clitocybes, ibid., 14(supplement) :4-37. Figs. 1-18. 1949.

MuRRiLL, William A.: Florida Tricholomas, Lloijdia, 12(l):62-69. 1949.
Atkinson, George F.: Collybia campanella Peck, and its near relatives in the

eastern United States, New York State Museum Bull, 205-206:61-65. 1918.
CoKER, W. C., AND H. C. Beardslee: The CoUybias of North Carolina, /. Elisha

Mitchell Sci. Soc, 37(1-2) :83-107. PI. 1 (col). Pis. 4-23. 1921.
• , and : The Laccarias and Chtocybes of North Carolina, ibid.,

38(1-2) :98-126. 1922.
Boursier, M.: Note sur le genre Mucidula Pat., Bull, trimestr. soc. mycol. France,

40:332-333. 1926.
Morgan, A. P.: North American species of Marasmius, J. Mycology, 11(5) :201-

212, (6):233-247. 1905; 12(1) :l-9. 1906.
: Descriptive synopsis of Morgan's North American species of Marasmius,

ibid., 12(4):159-162. 1916. (Species key to the foregoing paper.)
Pennington, L. H.: New York species of Marasmius, New York State Museum

Bull, 179:52-79. 1915.
KtJHNER, R.: Etudes sur le genre Marasmius, Le Botaniste, 25(1-2) :57-114. Illus.

1933.
Singer, Rolf: Studien zur Systematik der Basidiomyceten: I (6). Abgrenzung

zwischen Collybia und Marasmius, Beihefte Botan. Centr., Abt. B, 56(1-

2) :157-163. 1936. (Contains key to the subfamily Marasmioideae and to the

sections of the genus Marasmius.)
Petch, T.: a revision of Ceylon Marasmii, Brit. Mycol Soc. Trans., 31(1-2) :19-

U.Pls. 2-4 (col). 1947.
Bataille, F. : Flore monographique des Marasmes d'Europe, 37 pp. Besangon,

1919.
Binder, David H.: The genus Schizophyllum: I. Species of the Western Hemi-
sphere, Am. J. Botany, 20(8) :552-564. Pis. 33-36. Fig. 1. 1933.
Smith, Alexander H.: North American species of Mycena, pp. i — xviii, 1-521.

Pis. 1-99. Figs. 1-56. Ann Arbor, Univ. Michigan Press, 1947.
Beardslee, H. C, and W. C. Coker: The Mycenas of North Carolina, J. Elisha

Mitchell Sci. Soc, 49(1-2) :49-91. Pis. 6-30. 1924.
OoRT, A. J. p.: De Nederlandsche Mycenas, Mededeel Nederland. Mycol Ver.,

16-17:163-183. 1928.
Cejp, Karel: Omphalia (Fr.) Qu6L, in Atlas des Champignons de I'Europe, Vol.

4. 157 pp. 58 pis. Prag. Kavina et Pilat.
Romagnesi, H.: Quelques points de taxonomie: I. Sur un groupe particulier

d'Omphalia; II. Sur le genre Fulvidula, Bull trimestr. soc. mycol France,

58:81-89. 1942.
Sartory, a., et L, Maire: Compendium H.ymenomycetum, II. Lepiota, 512 pp.

Illustrated. Paris, Librairie le Frangois. 1925-1927.
Kauffman, Calvin H.: The genus Lepiota in the United States, Papers Mich.

Acad. Sci., 4:311-344. Pis. 15-18. 1925.
Murrill, William A.: Florida Lepiotas, Lloydia, 12(1):56-61. 1949.
Morgan, A. P.: North American species of Lepiota, J. Mycology, 12(4):154-159,

(5):195-203, (6):242-248. 1906; 13(1):1-18. 1907.

LIST 41. AGAKICACEAE (iN THE BROADER SENSE) 737

Beeli, M.: Fungi Goassensiansi: IX. Genre Lepiota, Bull. soc. roy. botan. Belg. 64,

2me s6t., 14(2):206-219. Pis. 25-27. 1912.
Kalm AR, ZoLTAN : Nouvelle clef pour la determination des Lepiotes de la Hongrie,

Acta Mycologica Hungarica, 4(1-2): 15-30. 1947.
KiJHNER, R. : Recherches sur le genre Lepiota, Bull, trimestr. soc. mycol. France,

52(2) :177-2S8. Figs. 1-9. 1936.
HuYSMAN, H. S. C.: Observations sur le "genre" Lepiota, Mededeel. Nederland.

Mycol. Ver. Leyden, 28:3-54. 12 figs. 1943.

: Un faux Lepiota. Le genre Coolia nov. gen., ibid., 28:54-60. 1 fig. 1943.

Smith, Alexander H., and Rolf Singer: A monograph of the genus Cysto-

derma. Papers Mich. Acad. Sci., 30 :71-124. Pis. 1-5. Figs. 1-12. 1944 (1945).
Smith, Helen V.: The genus Limacella in North America, Papers Mich. Acad.

Sci., 30:125-147. PL I. 1944 (l945).
Sartory, a., et L. Maire: Compendium Hymenomycetum : L Amanita, 447 pp.

26 pis. Numerous text figs. Paris, Librairie le Frangois, 1922-1923.
Atkinson, George F.: Six misunderstood species of Amanita, Mem. Torrey

Botan. Club, 17:246-252. 1918.
Martin, G. W.: Some Amanitas from eastern Iowa, Proc. Iowa Acad. Sci.,

32:205-218. 3 pZs. 1925.
Gilbert, Jean-Eduard: Le genre Amanita Persoon (Amanita s.st. — Amanitopsis

R. — ^Limacella E.). Etude morphologique des especes et vari^t^s. Revision

critique de la syst^matique, 188 pp. Lons-le-Saunier, Lucien Declume, 1918.
: Notules sur les Amanites: Supplement: commentaires et conjectures sur

quelques Amanites mal connues, 23 pp. 1 pi. Paris, published by the author,

1941.
: Notules sur les Amanites: XXX. Amanites d'Europe, 3 pp. Paris, pub-
lished by the author, 1941.
-: Amanitaceae, Iconographia Mycologica, 27(Suppiement l):8-427. 73 col.

pis. 1940-41. (Besides the foregoing the author has published various other
" Notules " on Amanita in Bull, trimestr. soc. mycol. France between 1924 and
1930.)

Vesely, Rudolf: Revisio critica Amanitarum europaearum, Ann. Mycol.,
31(4) :209-304. Pis. 8-25. Figs. 1-7. 1933.

: Amanita, Atlas des Champignons de I'Europe, Serie A, Tome 1, Fas-
cicules 1-5, 80 pp. 40 pis. Prag, Charles Kavina and Albert Pilat, 1934.

Huysman, H. S. C: Opmerkingen en problemen betreffende de taxonomie van
het geslacht Amanita, Mededeel. Nederland. Mycol. Ver. Leyden, 26:1-27.
7 figs. 1942.

Imai, Sanshi: Studies on the Agaricaceae of Japan: I. Volvate Agarics in Hok-
kaido, Botanical Magazine (Tokyo), 47(558) :423-432. 1933. (List with no
descriptions except some species described as new.) II. Lactarius in Hokkaido,
i6tc^., 49(585) :603-610. 1935.

: Studia Agaricacearura Japonicarura, I. ibid., 53(633) :392-399. 1939; II.

ibid., 55(658) :444-451; III. ibid., 55(659) :514-520. 1941. (This is a continua-
tion of Studies on the Agaricaceae of Japan.)

Kalmar, Zoltan: A galocagombdk (Amanita) hatd,roz6 tdblazata. (Hungarian),
Acta Mycologica Hungarica, 1(3-4) :101-1 10. 1944.

Gallfy, Zoltan: Legyolo — Amanita muscaria (L.) Pers. (Hungarian), Acta
Mycologica Hungarica, 3(1-4) :26-28. 1946. (Description of Amanita mus-
caria and six varieties.)

MuRRiLL, W. A.: The Amanitas of Eastern United States, Mycologia, 5(2) :72-86.
Pis. 85-86. 1913.

738 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

MuRRiLL, W. A.: Florida Amanitas, Lloydia, 11(2):99-110. 1948. (Contains a key
to all known species of Venenarius (Amanita) occurring in Florida.)

Seyot, Pierre: Les Amanites et la tribu des Amanit^es. 120 pp. 59 figs. Nancy,
Edit, des Arts Graphiques Modernes, 1930.

Beardslee, H. C: Notes on the Amanitas of the Southern Appalachians: I. Sub-
genus Amanitopsis, in Lloyd, C. G. : Mycological Writings, vol. 1., 7 pp.3
pis. 1902.

HoTsoN, J. W.: The Amanitae of western Washington, Mycologia, 28(l):63-76.
Figs. 1-4. 1936.

BuRLiNGHAM, Gertrude S.: A study of the Lactariae of the United States, Mem.
Torrey Botan. Club, 14:1-109. Figs. 1-15. 1908.

: New or noteworthy species of Russula and Lactaria, Mycologia,

28(3):253-267. Ft>. 1-8. 1936.

Beardslee, H. C., and Gertrude S. Burlingham: Interesting species of Lac-
tariae from Florida, Mycologia, 32(5):575-586. Figs. 1-4. 1940.

MuRRiLL, W. A.: Species of Florida Lactarius Fr., Lloydia, 11(2):86-9S. 1948.
(Contains a key"^ to all of the species of the genus known to occur in
Florida.)

Coker, W. C: The Lactarias of North Carolina, ./. Elisha Mitchell Sci. Sac,
34(l):l-62.P/s. 1 (coL)-40. 1918.

NtJESCH, E.: Die Milchlinge (Pilzgattung Lactarius). Bestimmungsschliissel und
Beschreibung der Milchlinge Mitteleuropas, 50 pp. St. Gallen, Switzerland,
published by author, 1921.

RoMAGNESi, H. : Les lactaires. C16 pratique de determination des especes d'Europe.
Rev. mycol, N.S., 4(suppl6ment 1):8-21. 1939.

■ — ■: Recherches sur les Lactaires de la section des Fuliginosi Konrad, ibid.,

14(2):103-112. i^t>s. 1-3. 1949.

Knauth, B., UND W. Neuhoff: Die Milchlinge (Lactarius), Die Pilze Mitteleu-
ropas, 2(5b, 6b, 9b-12b):l-64(?). Colored pis. 1-14(?). 1 uncolored pi. Leipzig,
Werner Klinkhart, 1937-1943. Uncompleted.

KoNRAD, P.: Les Lactaires. Notes critiques et essai de classification, Bidl. trimestr.
soc. mycol. France, 41(1):160-191. 1935.

Heim, Roger: Les Lactario-Russul^s du Domaine Orientale de Madagascar.
Essai sur la classification et la phylogenie des Ast^rosporales, pp. 1-196. 8 pis.
(4 col.). 59 figs. 2 phylogenetic diagrams. Paris, Laboratoire de Cryptogamie
du Museum National d'Histoire Naturelle, 1937 (1938).

Singer, Rolf: Monographie der Gattung Russula, Hedwigia, 66:163-260. 1 pi.
1926.

: Wie bestimmt man frische Taublinge? Z. fur Pilzkunde, 6(11):169-176.

1927. (Key to the common German species of Russula.)

: Neue Mitteilungen iiber die Gattung Russula, Hedwigia, 68(3-4) :191-

202. 1928.

: Monographie der Gattung Russula, Beihefte Botan. Centr., Zweite Abt.,

49:205-280. 1932.

: Sur la classification des Russules, Bull, trimestr. soc. mycol. France,

41(2):281-304. 1935.

: Supplemente zu meiner Monographie der Gattung Russula, Ann. Mycol.,

33(5-6) :297-352. 1935.

: Notes sur quelques Basidiomycetes, Rev. mycol., N.S., l(2):75-84. PI. 6.

1936. (Contains keys to the more or less green species of Russula in France
and to the species of Russula associated with the birch in France and border-
ing countries, also key to distinguish the genera Phyllotopsis, Dochmiopsis,
Rhodotus, and Octojuga.)

LIST 41. AGAEICACEAE (iN THE BROADEK SENSE) 739

— : Notes sur quelques Basidiomycetes, 2 me. s6r., ibid., l(4):279-293. 1936.
(Contains a key to the yellow species of Russula in Europe.)

New and. interesting species of Basidiomycetes, II, Papers Mich. Acad.

Sci., 32:103-150. PI. 1. 1946 (1948).
ScHAEFFER, JuLius: Russula-Monographie, Ann. MycoL, 31(5-6) :305-516. Pis.

26-27. 1933; 32(3-4) :141-243. Pis. 1-4. 1934.
: Bestimmungstabelle fiir die europaischen Taublinge, Z. fur Pilzkunde,

12(2):48-53, (3):83-91. 1933.
: Le Systeme naturel des Russules, Bull, trimestr. soc. rmjcol. France,

41(2) :263-276. 1935.
: Die Taublinge (Russulae), Die Pilze Mitteleuropas, 3(1-3). pp. (?).

1942-1943. Uncompleted.
-, W. Neuhoff, und W. G. Heiter: Die Russulaceen. Bestimmungstabelle

fiir die mitteleuropaischen Russula- und Lactarius-Arten, Sydowia, Ann.

MycoL, 3(1-6) :150-173. 1949.
Maire, Rene : Les bases de la classification dans le genre Russula, Bull. soc. mycol.

France, 26 :49-125. Figs. 1-6. 1910.
RoMAGNESi, H.: Contribution a I'^tude des Russules de la flore frangaise, Bull.

trimestr. soc. mycol. France, 59:61-71. Figs. 1-4. 1943.
Melzer, v., et Jar. Zvara: Cesk^ holubinky (Russulae Bohemiae), Archiv

prirod. vyzk. Cech. 17:1-126. 25 ^^s. Prag, 1927.
, ET : Cesk^ holubinky (Russulae Bohemiae). Flore monographique

des Russules de Boheme. Avec un tableau analytique des especes. R^sum^,

Bull, trimestr. soc. mycol. France, 44:135-146. 1928. (A French summary of

the preceding book.)
Zvara, Jar.: Russula atropurpurea Kromb. et ses vari^t^s, ibid., 47:44-51. Pis.

1-2. 1931.
Bonus, Gabor: Wie steht es mit der Russula sardonia? Acta Mycologica Hun-

garica, 1(3-4) :123-127. 1944. (Discussion and key for the distinction of the

Russula sardonia-sanguinea-emetica group.)
: A Russuldk-galambgombdk hatdrozo kulcsa. ibid., 3(1-4) :55-71. 1946.

(In Hungarian.) (Contains key with brief descriptions of 92 species and 2

subspecies or varieties of Russula.)
Crawshay, Richard: The spore ornamentation of the Russulas, 188 pp. 48 pis.

London, Bailliere, Tindall and Cox, 1930.
Chiu, Wei-Fan: The Russulaceae of Yunnan, Lloxjdia, 8(l):31-59. 48 figs. 1945.
Winters, Grace: The Iowa species of Russula, Univ. Iowa Studies in Natural

History, 11:5-30. 1 pi. 1926. (Keys to 39 species with descriptions and spore

measurements.)
Bataille, Fr;^d^ric: Flore monographique des Asterosporfe, Lactaires et Rus-
sules, Mem. soc. d' Simulation du Doubs, 8me s6r., 2:163-260. 1907.
Burlingham, Gertrude S.: Spore ornamentation of some American Russulas

and a new species of Lactaria, Mycologia, 34(1):8-12. Figs. 1-8. 1942.
Kauffman, Calvin H.: Unreported Michigan fungi for 1908 with a monograph

of the Russulas of the state, Mich. Acad. Sci. Kept., 11 :55-91. Figs. 1-3. 1909.
Beardslee, H. C: The Russulas of North Carolina, /. Elisha Mitchell Sci. Soc,

33(4):147-199.PZs. 70-111. 1918.

Red- or Pink-spored Genera.

MuRRiLL, W. A.: The genus Lepista, Mycologia, 7(2):105-107. 1915.

Bonus, Gabor: Volvaria. Bocskorosgomb^k hatdrozo kulcsa. Acta Mycologica

Hungarica, 2(3-4) :95-96. 1945. (In Hungarian.) (Key to 10 species of

Volvaria.)

740 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

RoMAGNESi, H.: Quelques observations sur les Rhodophyllus, Bull, trimestr. Soc.

mycol. France, 48(3-4) :306-323. Figs. 1-11. 1932.
: Les Rhodophylles de Madagascar, in Prodrome a une flore mycologique de

Madagascar publie sous la direction de R. Heim, Tome II. 175 pp. 45 figs.

Paris, Laboratoire de Cryptogamie du Museum Nationale d'Histoire

Naturelle, 1941.
CoKER, W. C: North Carolina species of Volvaria, /. Elisha Mitchell Sci. Soc,

63(2) :220-230. 5 pis. 1947.

Ochre- or Rust-spored Genera.

(See also above under White-spored genera for yellow-spored species of Lac-
tarius and Russula.)

Singer, Rolf: Contributions toward a monograph of the genus Crepidotus,

Lilloa 13:59-95. Fig. 1. 1947.
Bataille, FrediSric: Flore analytique des Inocybes d'Europe, Bull. soc. d'His-
toire Naturelle Doubs 18, 27 pp. 1910.
Kauffman, Calvin H. : Studies in the genus Inocybe, New York State Museum

Bull. 223-224:43-60. 1921.
Sartory, a., et L. Maire: Synopsis du genre Inocybe, 246 pp. 2 pis. Paris, E. le

Francois, 1923.
Heim, Roger: Le genre Inocybe, in Encyclop^die Mycologique, vol. 1, pp. 1-429.

Pis. 1-35. Figs. 1-219. Paris, Paul Lechevalier et Fils, 1931.
BouRSiER, J., ET R. KtJHNER: Notes sur le genre Inocybe, Bull, trimestr. soc.

mycol. France, 44(2):170-189. Figs. 1-9. 1928.
KuHNER, R., ET J. Boursier: Notes sur le genre Inocybe: I. Les Inocybe gonio-

spor^es. Bull, trimestr. soc. mycol. France, 48(2):118-161. Figs. 1-31. 1932.
■ : Notes sur le genre Inocybe: les Inocybe goniospor^es (fin), ibid., 49(1) :81-

121. Figs. 32-53. 1933.
Stuntz, D. E. : Studies in the genus Inocybe: I. New and noteworthy species from

Washington, Mycologia, 39(1) :21-55. Figrs. 1-50. 1947.
KiJHNER, R.: Le genre Galera, in Encyclopedic Mycologique, vol. 7, 240 pp. 75

figs. Paris, Paul Lechevalier, 1935.
Metrod, Georges: Description de Galera, Bull, trimestr. soc. mycol. France,

56(1) A6-55. Figs. 1-4. 1940.
Atkinson, George F.: The genus Galerula in North America, Proc. Am. Phil.

Soc, 57:357-374. 1918.
Kauffman, Calvin H. : The genera Flammula and Paxillus and the status of the

American species. Am. J. Botany, 13(l):ll-32. 1926.
RoMAGNESi, H. : Sur quelques groupements naturels d'Agarics ochrospor^s, Rev.

mycol., N.S., 1(4):207-213. 1936. (Includes a key to the genera segregated

from Flammula and Pholiota.)

: Essai sur le genre Tubaria W. Sm., ibid., 5(l):29-43. 1940. '

: Etudes compl^mentaires sur le genre Tubaria et sur deux Naucoria

tubarioides, ibid., 8(3-4) :26-35. 8 figs. 1943.
-: Description de quelques especes d'Agarics ochrospor^s, Btdl. trimestr.

soc. mycol. France, 58:121-149. Figs. 1-15. 1942. (Includes discussion of

Alnicola, Naucoria, Agrocybe, Flammula, Galerina, and Conocybe.)
Overholts, L. 0.: a monograph of the genus Pholiota in the United States, Ann.

Missouri Botan. Garden, 14(2):S7-210. Pis. 8-24. Figs. 1-171. 1927.
Harper, Edward T.: Species of Pholiota in the region of the Great Lakes, Trans.

Wisconsin Acad. Sci., 17:470-502. Pis. 26-55. 1913.
: Species of Pholiota and Stropharia in the region of the Great Lakes, ibid.,

17:1011-1026. PZs. 69-77. 1914.

LIST 41. AGARICACEAE (iN THE BROADER SENSE) 741

: Additional species of Pholiota, Stropharia and Hypholoma in the region

of the Great Lakes, ibid., 18:392-431. Pis. 11-24. 1916.
Singer, Rolf: Studien zur Systematik der Basidiomyceten : I (8). Die Gattung

Pholiota ist kiinstlich, Beihefte Botan. Centr., Abt. B, 56(1-2) :165-174. 1936.

(Contains a key to tlae European species of Agrocybe and to the modern

segregates of the Friesian genus Pholiota.)
Bataille, Frederic: Flore monographique des Cortinarius d'Europe, Exir.from

Bull. soc. d'Histoire Naturelle Doubs, I vol., 162 pp. 1912.
Kauffman, Calvin H.: The genus Cortinarius with key to the species, J. My-
cology, 13(1) :32-39. Pis. 93-100. 1907.
Smith, Alexander H.: Studies in the genus Cortinarius I, Contributio7is from the

Univ. Michigan Herbarium, 2:1-42. Pis. 1-12. Ann Arbor, Univ. Michigan

Press, 1939.
: New and unusual Cortinarii from Michigan, with a key to the North

American species of subgenus Bulbopodium, Bull. Torrey Botan. Club,

69(3):44-64. Figs. 1-7. 1942.
MuRRiLL, W. A.: Some Florida gill fungi, /. Elisha Mitchell Sci. Sac, 55 :361-372.

1939. (Contains a key to the Alachua County species of Cortinarius.)
Henry, R.: Etude de quelques Cortinaires du groupe des Scauri. Deux especes

nouvelles, Bull, trimestr. soc. mycol. France, 51(1) :34-101. P/s. 1-2. Figs. 1-10.

1935.

: Etude de quelques Cortinaires, ibid., 51(2):205-241. 2 figs. 1935.

: Revision de quelques Cortinaires, ibid., 53(l):49-80. 1937.

: Description de quelques dermocybes du groupe "Anomaliae" Fr., ibid.,

53(2):143-164. 1937.

: Suite a I'^tude du genre Hydrocybe, ibid., 56 :85-119. Figs. 1-6. 1940.

: Quelques Cortinaires " Hinnulo'ides " (Telamonias, Hydro-telamonias et

Hydrocybes hinnuloides), ibid., 57:17-35. 1941 (1942).

: Cortinaires nouveaux ou rares de la fiore frangaise, ibid., 59 : 52-60. 1943.

: Essai d'une cU dichotomique analytique provisoire destinee a faciliter

r^tude des Cortinaires du groupe des Scauri, Rev. mycol. , N.S., 8(suppl6ment

a2):l-56. 1943.
: Essai d'une cl6 dichotomique provisoire destinee a faciliter I'^tude des

Cortinaires de groupe des Phlegmacia (Cliduchi et Elastici), ibid., 10(sup-

pl^ment a 2):44-82. 1945 (1947).

Purple-spored Genera.

Harper, Edward T.: Species of Hypholoma in the region of the Great Lakes,
Trans. Wisconsin Acad. Sci., 17:1142-1164. Pis. 72-84. 1914.

Parker, Charles S.: A taxonomic study of the genus Hypholoma in North
America, Mycologia, 25(3):160-212. Pis. 26-31. Figs. 1-2. 1933.

Smith, Alexander H.: Studies of North American Agarics, I, Contributions from
the Univ. Michigan Herbarium, 5:1-73. Pis. 1-32. 1941. (Includes a discus-
sion of Psathyrella.)

: Studies in the genus Agaricus, Papers Mich. Acad. Sci., 25:107-138.

Pis. 1-10. Figs. 1-4. 1939 (1940).

HoTSON, J. W., and D. E. Stiintz: The genus Agaricus in western Washington,
Mycologia, 30(2) :204-234. Figs. 1-10. 1938.

Mendoza, J. M., AND SiMEONA Leus-Palo: a revision of the genus Psalliota in
the Philippines, Philippine J. Sci., 72:337-345. 8 pis. 1940.

CoKER, W. C: The Chapel Hill species of the genus Psalliota, /. Elisha Mitchell
Sci. Soc, 43(3-4) :243-256. Frontis. Pis. 38-46, 48. 1928.

742 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Kalmar, Zoltan: A csiperke (Psalliota) nemzetseg europai fajai, Acta Mycologica
Hungarica, 3(1-4) :10-22. 1946. (In Hungarian.) (Key and descriptions of 16
species of Psalliota (Agaricus).)

Black-spored Genera.

(See also Purple-spored genera.)

Massee, George: A revision of the genus Coprinus, Ann. Botany, 10(38) :123-

IM.Pls. 10-11. 1896.
Kuhner, R., et M. Josserand: Description de quelques especes du groupe de

Coprinus plicatilis (Curt.) Fr., Bull, trimestr. soc. mycol. France, 50(l):53-63.

Figs. 1-5. 1934.
RoMAGNESi, H.: Les Coprins, Rev. mycol., N.S., 6(suppl^ment a l):20-35. 1 pi.

1941.
: fitude de quelques Coprins (2me serie), ibid., 10(5-6) :73-89. 8^grs. 1945

(1947).
Morgan, A. P.: North American species of Agaricaceae, Melanosporae, /. My-
cology, 13(2):53-62, (4):143-153, (6):246-255. 1907; 14(l):27-32, (2):64-75.

1908.
Kauffman, Calvin H.: The genus Gomphidius in the United States, Mycologia,

17(3):113-126. PZs. 12-14. 1925.
Singer, Rolf: The genus Gomphidius Fries in North America, Mycologia,

41(4):462-489. Figs. 1-3. 1949.
Sartory, a., et L. Maire: Synopsis du genre Gomphidius, 23 pp. Paris, 1922(?).

, ET : Synopsis du genre Chitonia. Paris, 1923.

NiJESCH, E.: Die schwarzsporigen Blatterpilze der Kantone St. Gallon und

Appenzell. Bestimmungsschliissel, Artenbeschreibung mit kritischen Bemerk-

ungen und Fundverzeichnis, Jahrh. d. St. Gallischen Nalurwiss. Ges. 57(2):

141-169. 1920-1921.

List 42. Gasteromyceteae — General Works

(See also Mattirolo in List 17 and Graham in List 34.)

CoKER, W. C, and John N. Couch: The Gasteromycetes of the Eastern United
States and Canada, ix + 201 pp. 123 pis. Chapel Hill, Univ. North Carolina
Press, 1928.

: The Gasteromycetes of North Carolina, /. Elisha Mitchell Sci. Soc,

38(3-4) :23 1-243. PZs. 71-83. 1923.

Morgan, A. P.: North American Fungi. The Gasteromycetes, /. Ciyicinnati Soc,
Natural History, 11:141-149. Pis. 2-3. 1883; 12:8-22, 163-172. Pis. 1-2.
1890; 14:5-21, 141-148. Pis. 1, 2, 5. 1891-1892.

Johnson, Minnie May: The Gasteromyceteae of Ohio: Puffballs, bird's-nest
fungi and stinkhorns, Ohio Biological Survey Bull., 4(7):273-352. 1929.

Kambly, Paul E., and Robert E. Lee: The Gasteromycetes of Iowa, U7iiv. Iowa
Studies in Natural History, 17(4):121-185. Pis. 9-11. 1936.

Henry, LeRoy K. : A review of the Gasteromycetes (fungi) of Western Pennsyl-
vania, Ann. Carnegie Museum, 30:339-362. Pis. 1-4. 1947. (Article XX.)

LIST 43. HYMENOGASTRALES, SCLERODERMATALES, ETC. 743

Kauffman, C. H.: Unreported Michigan fungi for 1907, with an outline of the
Gasteromycetes of the state, Mich. Acad. Sci. Rept., 10:63-84. 1908. (In-
cludes a key to the genera and species.)

Moffat, Will S.: The higher fungi of the Chicago region: II. The Gastero-
mycetes, Chicago Acad. Sci. Natural Histonj Bull., 7(2) :l-24. Pis. 1-26. 1923.

GiLMAN, Joseph C.: Illustrations of the fleshy fungi of Iowa: VII. Some common
puff balls, Proc. Iowa Acad. Sci., 52:113-119. Figs. 1-6. 1945; VIII. The
stinkhorns, ibid., 53:147-151. Figs. 1-5. 1946; IX. Further Gasteromycetes,
ibid., 54:131-137. Figs. 1-7. 1947 (1948).

Massee, George: A monograph of the British Gasteromycetes, Ann. Botany,
4(13):1-103. P^s. 1-4. 1891.

Fries, Thore C. E.: Sveriges Gasteromyceter, Arkiv for Botanik, 17(9):l-63.
Figs. 1-43. 1921. (Keys to orders, families, and genera, and descriptions of
species of all Gasteromycetes known to occur in Sweden. Every genus and
most species illustrated.)

Lloyd, C. G.: The genera of the Gasteromycetes, Mycological Writings, 1 (1898-
1905). (Separate pagination, pp. 1-24.) Pis. 1-11. Jan. 1902. (Key to all the
genera (American) and good illustrations.)

Cunningham, G. H.: The Gasteromycetes of Australia and New Zealand,
XV + 236 pp. Pis. 1-37. 1 text fig. Dunedin, New Zealand, published by the
author, 1944.

KiLLERMANN, S.: Bayerische Gasteromyceten, Kryptogamische Forschungen.
Bayerische Botanische Gesellschaft. Erforschungen der Heimischen Flora,
7:498-512. 2 pis. 1926.

ViEGAs, A. P.: Algunos fungos do Brasil: X. Gastromicetos, Bragantia, 5(9):583-
595. 1945.

Lange, Morten: Macromycetes. I. The Gasteromycetes of Greenland, Meddalel-
ser om Gr^inland, 147(4) :l-32. Illustrated. 1948.

List 43. Hymenogastrales, Sclerodermatales, etc.

Hesse, R.: Die Hypogaen Deutschlands. Natur, und Entwicklungsgeschichte

sowie Anatomie und Morphologie der in Deutschland vorkommenden Triif-

feln und der diesen verwandten Organismen nebst praktischen Anleitungen

beziiglich deren Gewinnung und Verwendung, vol. 1, pp. 1-149. Pis. 1-13.

Halle a. S., Ludw. Hofstetter, 1891. (Vol. 2 contains the Tuberales.)
Bataille, F.: Flore analytique et descriptive des Hymenogastrac^es d'Europe,

Bull. soc. mycol. France, 39:157-196. 1923.
Soehner, Ert: Die Formenkreis von Hymenogaster tener Berk, et Br., Hedwigia,

64:192-202. Figs. 1-15. 1913. (Key and description of six species and varieties

of Hymenogaster of the H. tener series.)
: Hymenogasterstudien. Das Formenkreis um Hymenogaster verrucosum

Buch., Hechvigia, 81(3-4): 162-1 92. PL 4. 1943.
Lloyd, C. G.: The Hymenogastraceae. The Octaviana group. The genus Arc-

angeliella, Mycological Notes, 7:1138-1143. Figs. 2152-2176. 1922.
Velenovsky, J.: Les especes tch^ques du genre Rhizopogon Fr., Mykologia,

8:89-94.1^^.1931.

744 GUIDE TO THE LITEKATUKE FOR THE IDENTIFICATION OF FUNGI

Zeller, Sanford M., and Carroll W. Dodge: Rhizopogon in North America,

Ann. Missouri Botan. Garden, 5(1) :l-36. Pis. 1-3. 1918.

, AND : Gautieria in North America, ihid., 5(2) :133-142. PL 9. 1918.

, AND : Arcangeliella, Gymnomyces and Macowanites in North

America, ihid., 6(1) :49-59. Figs. 1-3. 1919.
, AND — : Leucogaster and Leucophlebs in North America, ihid.,

11(4):389-410. PZ. 11. 1924.
— , AND : Hysterangium in North America, ihid., 16(l):83-228. Pis.

1-3. 1929.

, AND : New species of Hydnangiaceae, ihid., 22:365-373. 1935.

, AND : Elasraomyces, Arcangeliella, and Macowanites, ihid..

23(4):599-638. 1936.
-, AND : Melanogaster, ihid., 23(4):639-655. 1936.

Dodge, Carroll W.: Alpova, a new genus of Rhizopogonaceae, with further

notes on Leucogaster and Arcangeliella, Ann. Missouri Botan. Garden,

18 :457-464. PZ. 40. 1931.
, AND Sanford M. Zeller: Hydnangium and related genera, ihid.,

23(4):565-598. 1936.
Zeller, Sanford M.: New and noteworthy Gasteromycetes, Mycologia, 31(1) :1-

32. Figs. 1-54. 1939. (Includes a key to the families of the Hysterangiales.)
: Further notes on fungi, ihid., 33(2):186-214. Figs. 1-17. 1941. (Mostly

Hymenogastrales.)
: North American species of Galeropsis, Gyrophragmium, Longia and

Montagnea, ihid., 35(4):409-421. 1 fig. 1943.

List 44. Lycoperdales, including Tulostomataceae and

Podaxaceae

Verwoerd, Len: Suid-Afrikaanse Lycoperdaceae en Nidulariaceae, Ann. Univ

Stellenhosch, 3A(l):l-45. Figs. 1-14. 1925.
Lloyd, C. G.: The Lycoperdons of the United States, Mycological Writings,

2 (1905-1908); Mycological Notes, 20:221-238. Pis. 41-67. 1905.
Massee, George: A monograph of the genus Lycoperdon, /. Roy. Microscop.

Soc, 1887(5) :701-727.P/.s. 12-13. 1887.
Peck, Charles H. : New York species of Lycoperdon, Report of the State Botanist

{New York), 32:58-72. 1879.
Lohman, M. L. : The Iowa species of Lycoperdon, Univ. Iowa Studies in Natural

History, 12(4):5-28. 2 pis. 1927.
Massee, George A.: A revision of the genus Bovista, J. Botany, 26(4):129-137.

PI. 282. 1888.
Lloyd, C. G.: Key to American species of Catastoma, Mycological Notes, 7:1167-

1168. 1922.
: The genus Bovistella, Mycological Writings, 2 (1905-1908); Mycological

Notes, 23:277-287. Pis. 86-89. 1905.
Long, W. H., and David J. Stoufper: Studies in the Gasteromycetes: II. Bovi-

stina, a new genus, Mycologia, 33(3):270-273. 1 fig. 1941.

LIST 44. LYCOPERDALES, INCLUDING TULOSTOMATACEAE 745

Zeller, Sanford M.: Representatives of the Mesophelliaceae in North America,

Mycologia, 36(6) :627-637. Figs. 1-6. 1944. (Keys to the Suborder Lyco-

perdineae, separating the three families Lycoperdaceae, Mesophelliaceae, and

Geastraceae. Also a key to the 4 genera of Mesophelliaceae, Abstoma,

Radiigera, Mesophellia, and Castoreum, and a monograph of the genus

Radiigera.)
Pole-Evans, I. B., and A. M. Bottomley: On the genera Diplocystis and

Broomeia, Trans. Roy. Soc. S. Africa, 7:189-192. Pis. 19-22. 1919.
Lloyd, C. G.: The Geasters, Mijcological Writirigs, 1(1898-1905). Figs. 1-80.

1902. (Separate pagination, pp. 1-44.)
Morgan, A. P. : The North American Geasters, Am. Naturalist, 18(10) :963-970.

Figs. 1-12. 1884.
DeToni, J. B.: Revisio monographica generis Geastris Mich, e tribu Gastero-

mycetum. Rev. mycoL, 9(34):61-77, (35):125-133. PZs. 62-63. 1887.
Destree, Caroline E.: Revision des Geaster observfe dans les Pays-Bas,

Nederlandsch Kruidkundig Ar chief, ser. 2, deel 6, stuk. 3. 488 pp. Illustrated.

Nijmegen, 1894.
Longnecker, William M.: The Geasters of Iowa, Univ. Iowa Studies in Natural

History, 12(4):29-47. 2 pis. 1927.
Coker, W. C.: The Geasters of the United States and Canada, J. Elisha Mitchell

Sci. Soc, 39(3-4): 170-221. Pis. 18-36. 1924.
Ponce de L:^on, Patricio : Contribuci6n al estudio de los Gasteromicetos Cubanos :

I. El g^nero Geastrum en Cuba, Rev. soc. cubana Botan., 3(3):63-70. ^ figs.

1946.
Reisner, Otaker: Les especes du genre Geaster Mich, en Boheme, Travaux

Mycologiques Tchecoslaviques, 1:1-9. 1 pi. 1924. (Reprinted from Mxjkologia,

1, 1924.)
Willis, J. H.: The Geastrae or "earth-stars" of Victoria, Victorian Naturalist,

51(5):115-124. PL 24. S figs. 1934.
Long, W. H.: Studies in the Gasteromycetes : XL The genera Trichaster and

Terrostella, Mycologia, 37(5):601-608. Figs. 1-4. 1945.
, AND David J. Stouffer: Studies in the Gasteromycetes: XVI. The

Geastraceae of the Southwestern United States, ibid., 40(5) :547-585. Figs.

1-52. 1948.
Lloyd, C. G.: The Tylostomeae, Mycological Writings 2 :(1905-190S). Pis. 10, 11,

20, 28, 74-85. 1906. (Separate pagination, pp. 1-28.)
White, V. S.: The Tylostomaceae of North America, Bull. Torrey Botan. Club,

28(8) :42 1-444. Pis. 31-40. 1901.
van der Byl, Paul A. : The genus Tulostoma in South Africa, Trans. Roy. Soc.

S. Africa, 9(2):185-186. PL 9. 1921.
Fries, Thore C. E.: Sveriges Tulostoma-arter, Botan. Notiser, 1921(1) :33-36.

Fig. 1. 1921.
Petri, L.: Sul valore diagnostico del capillizio nel genere "Tylostoma" Pers.,

Ann. MycoL, 2(5):412-438. PL 6 (col.). 25 text figs. 1904.
Massee, George: A monograph of the genus Calostoma, Ann. Botany, 2(5) :25-

45. PL 3. 1888.
BuRNAP, Charles Edward: Notes on the genus Calostoma, Botan. Gaz.,

23(3):180-192.P^. 19. 1897.
Lloyd C. G.: The genus Mitremyces, Mycological Writings, 2(1905-1908).

Mycological Notes, 30:238-243. Pis. 8, 9, 68, 69. Fig. 87. 1905.
Long, W. H., and 0. A. Plunkett: Studies in the Gasteromycetes: I. The genus

Dictyocephalos, Mycologia, 32(6):696-709. Figs. 1-13. 1940. (Contains also a

key to the genera of Family Tulostomataceae.)

746 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OP FUNGI

Long, W. H., and David J. Stoupfer: Studies in the Gasteromycetes : VII. The

genus Schizostoma, ihid., 35(l):21-32. Figs. 1-8. 1943.
, AND Sultan Ahmad: The genus Tylostoma in India, Farloivia, 3(2):225-

267. Figs. 1-25. 1947.
: Studies in the Gasteromycetes: XV. Notes on new or rare species of

Tylostoma, Lloydia, 10(2) :1 15-135. 19 ^^s. 1947.
-: Studies in the Gasteromycetes: XVII. Two interesting species from

Argentina, Lloydia, ll(l):57-59. Figs. 1-4. 1948. (Schizostoma argentinense
and Broomeia congregata.)

Rea, Paul Marshall: Fungi of Southern California, I, Mycologia, 34(5) :563-
574. 3 figs. 1942. (A study of Battarrea with a key to the two groups of
species.)

Long, W. H.: Studies in the Gasteromycetes: VIII. Battarrea laciniata, My-
cologia, 35(5):546-556. Figs. 1-6. 1943.

List 45. Nidulariales, including Arachniaceae

White, V. S.: The Nidulariaceae of North America, Bull. Torrey Botan. Club

29(5) :251-280. P/s. 14-18. 1902.
Lloyd, C. G.: The Nidulariaceae, Mycological Writings, 2:(1905-1908). Figs.

1-20. 1906. (Separate pagination, pp. 1-32.)
KiLLERMANN, S.: Die Nidularia Fr.-Gruppe, Kryptogamische Forschungen her-

ausgegeben von der Kryptogamenkommission der Bayerischen Botanischen

Gesellschaft zur Erforschung der heimischen Flora, 2(2):194— 198. PI. 5. 1931.
Long, W. H.: Studies in the Gasteromycetes: III. The Family Arachniaceae,

Mycologia, 33(4):350-355. Figs. 1-7. 1941. (Includes descriptions of a new

genus, Araneosa, and a key to the seven species of Arachnion recognized by

the author.)

List 46. Phallales

Fischer, E.: Untersuchungen zur vergleichenden Entwicklungsgeschichte und

Systematik der Phalloiden, I, Denkschrift der Schweizerischen Naturfor-

schenden Gesellschaft, 32(1):1-103. Pis. 1-6. 1890; II, ibid., 33(1):1-51. 3 pis.

5 figs. 1893; III, ibid., 36(2):l-84. 6 pis. 4 figs. 1900.
Lloyd, C. G.: Synopsis of the known Phalloids, Mycological Writings, 3:(1909-

1912). \07 figs. 1909. (Se])arate pagination, pp. 1-96.)
Fetch, T.: The Phalloideae of C'eylon, Ann. Royal Botanical Garden Peraderiiya,

4:139-184. 1908.
Martin, G. W.: Notes on Iowa fungi. 1928, Proc. Iowa Acad. Sci., 36:127-130.

1 pi. 1929. (Descriptions of the Iowa species of Mutinus.)
BoEDiJN, K. B.: The Phallineae of the Netherkmds East Indies, Bull. Jardin

Botaniqv^ de Buitenzorg, ser. Ill, 12:71-103. 12 figs. 1932.

LIST 47. SPHAEROPSIDALES 747

Long, W. H.: The Phalloideae of Texas, /. Mycology, 13(3) : 102-1 14. Pis. 102-106.

1907.
, AND David J. Stouffer: Studies in the Gasteromycetes : XVIII. The

Phalloids of the Southwestern United States, Lloydia, ll(l):60-76. Figs.

1-21. 1948.
KoBAYASi, Y. : Revisions of several species of Clathraceae, J. Japanese Botany,

13(4):262-270. Illustrated. 1937. (Japanese, with Latin diagnoses of species.)
Lopes, J. Pinto: Lysurus sulcatus (Cke. et Massee) Cunn. e a ordem das Phal-

lales em Portugal, Bol. soc. Broteriana, 19(2a ser., Pt. I) :125-162. 8 pis. 1944.

List 47. Sphaeropsidales

Bender, Harold B.: The genera of Fungi Imperfecti: North American species
and hosts with particular reference to Connecticut, 3 parts, 2000 pages.
Unpublished thesis, Yale University, (June) 1931.

: The Fungi Imperfecti: Order Sphaeropsidales. With keys and references

for the genera, 52 pp. North Woodbury, Conn., published by the author,
1934.

Grove, W. B.: British stem- and leaf-fungi (Coelomycetes). A contribution to our
knowledge of the Fungi Imperfecti belonging to the Sphaeropsidales and
Melanconiales : I. Sphaeropsidales. To the end of the Sphaerioideae which
have colourless or nearly colourless spores, xx + 488 pp., 31 figs., 1935; II.
Remainder of Sphaeropsidales and the Melanconiales, xii + 407 pp., 102
figs., 1937; Cambridge, Cambridge Univ. Press.

voN Hohnel, Franz: Zur Systematik der Sphaeropsideen und Melanconieen,
Ann. Mycol., 9(3) :258-265. 1911.

: System der Fungi Imperfecti Fuckel: I. Histiomyceten; II. Synnemato-

myceten, Mykologische Untersuchungen und Berichte, l(3):301-369. 1923.

Unamuno, p. Luis M. : Enumeracion y distribucion geografica de los Esferopsidales
conocidos de la peninsula Iberica y de las Islas Baleares. Familia Esferioid-
aceos, Memorias de la Academia de Ciencias Exactas, Fisicas y Naturales de
Madrid. Serie de Ciencias Naturales, 4:1-457. 1933. (A list of all species
known from the area of the work with no descriptions. Also a host index.)

Martin, George: The Phyllostictas of North America, /. Mycology, 2(2):13-20,
(3) :25-27. 1886.

Ellis, J. B., and B. M. Everhart: The North American Phyllostictas with
descriptions of the species published up to August, 1900, 79 pp. 1900.

Anderson, P. J.: Index to American species of Phyllosticta, Mycologia, 11(2) :66-
79. 1919. (Additions to the species of Phyllosticta that have been recorded
for North America since the publication of the preceding article together
with a complete host index for the now known North American species.)

Seaver, Fred. J.: Phyllostictales. Phyllostictaceae (pars). North American Flora,
6:1-84. 1922. (The genus Phyllosticta.)

DaCosta, G. C, and B. B. Mundkur: A revision of the Genus Phyllosticta in
India, Proc. Nat. Inst. Sci. India, 14(2):55-63. 1948.

CiFERRi, R. : Notae mycologicae et phytopathologicae, ser. I, No. 1-11, Ann.
Mycol., 20(1-2) :34-53. PI. 1. 1922. (Contains a conspectus of the species of
Phyllosticta on Acer and on Cydonia.)

748 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Tehon, L. R., and E. Y. Daniels: Notes on the parasitic fungi of Illinois: III,
Mycologia, 19(3):110-129. PL 11. 1927. (Contains, aside from descriptions of
new species, keys to the five known species of Phyllosticta parasitic on
Syringa vulgaris, to the four known species of Phyllosticta parasitic on
Solidago, to the four known species parasitic on Nymphaea advena, to the
three known species on Plantago, to the two species known on Sassafras, to
the five species known on Chenopodium, to the four species of Septoria
known on Festuca, and to the three species of Cercospora known on Setaria.)

, and G. L. Stout: Notes on the parasitic fungi of Illinois: IV, ibid.,

21(4):180-196. PL 13. 1929. (Contains a key to distinguish five genera of the
Family Stigmateaceae, Order Hemisphaeriales; also keys to the American
species of Phyllosticta on Rubus, to the three known species attacking
Asparagus officinalis, to the two species of Cercospora on Sarabucus, to eight
species of Diplodia occurring on Acer.)

: Notes on the parasitic fungi of Illinois: V, ibid., 25(4) :237-257. PL 34.

1933. (Contains keys to the 15 species of Phyllosticta known in America on
Quercus, to the 6 species of Stagonospora reported on Scirpus, and to the 10
species of Marssonina reported on Salix.)

Notes on the parasitic fungi of Illinois: VI, ibid., 29(4) :434-446. Figs. 1-9.

1937. (Contains keys to 4 species of Phyllosticta on Plantago, 10 species of
Macrophoma on grasses, and 5 species of Septoria on Quercus.)

Grove, W. B.: The British species of Phomopsis, Roy. Botan. Gardens, Kew. BuU.
Misc. Inform., 1917(2) :49-73. Pis. 1-2. 1917.

: Species placed by Saccardo in the genus Phoma, ibid., 1919(4) :177-201.

Figs. 1-23. (10):425-445. Figs. 1-6. 1919; 1921(4) :136-157. Figs. 1-8. 1921.
(Discusses and gives descriptions of many species placed by Saccardo in
Phoma but which must be transferred to other genera, e.g., Phomopsis,
Dendrophoma, Dothiorella, Cytospora, Diplodia, Camerosporium, Rhabdo-
spora, Gloeosporium, Colletotrichum, etc.)

: The British species of Cytospora, ibid., 1923(1) :l-30. 1923.

-: The British species of Ceuthospora and Cytosporina, ibid., 1923(10) :353-

359. 1923.
Gutner, L. S.: Materialien zu einer Monographic der Gattung Cytospora, Acta

Instituti Botanici Academiae Scientiarum U.S.S.R., ser. II, Fasc. 2, pp. 411-

484. Figs. 1-66. 1934. (Russian, with German summary.)
von Hohnel, Franz: Fragmente zur Mykologie: 973. tJber Myxosporella populi

Jaap, Sitzber. kaiserlichen Akad. Wiss. Wien Math, naturiv. Klasse, 125 :76-80.

1916. (Gives key to the genera of the family Sclerophomaceae.)
Sydow, H. und p. : Scleropycnis, ein neurer Gattungstypus unter den hyalosporen

Sphaeropsideen, Ann. MycoL, 9(3) :277-278. Figs. 1-4. 1911.
Jaczewski, a. L. : Monographie du genre Sphaeronema, Notwelle Memoire de la

Societe Imperials des Naturalistes de Moscou 15, 112 pp. Illustrated. 1898.
Petrak, F. : Mykologische Notizen 225: tJber einige Pleurostromella Neben-

fruchtformen von Cucurbitariaceen, A^in. MycoL, 21(3-4) :2 15-224. 1923.
Diedicke, H.: Die Gattung Phomopsis, Ann. MycoL, 9(l):8-35. Pis. 1-3. 1911.

(Besides a discussion of the known species of this genus and of the structural

cliaracters, tlie author gives a key distinguishing Phomopsis, Plenodomus,

Dothiopsis, Sclerophoma, and Sclerotiopsis.)

LIST 47. SPHAEROPSIDALES 749

— : Die Gattung Plenodomus Preuss, ibid., 9(2):137-141. PI. 8. 1911.

- — : Dothiopsis, Sclerophoma und Sclerotiopsis, ibid., 9(3) :279-285. PI. 15.

1911.

— : Die Gattung Asteroma, ibid., 9(5) : 534-548. PL 18. 1911.
— : Myxofusicoccum, nov. gen. Sphaeropsidearum, ibid., 10(l):68-72. Figs.

1-5. 1912.

Die Abteilung Hyalodidymae der Sphaerioideen, ibid., 10(2):135-152.

1912.
Davis, J. J. : North American Ascochytae, Trans. Wisconsin Acad. Sci., 19(2) :655-

670. 1919.
Swift, Marjorie E. : A new species of Chaetomella on rose, Mycologia, 22(4) :165-

168. Fig. 1. 1930. (Includes a key to the described species of Chaetomella with

spore measurements and hosts.)
DiEDicKE, H.: Die braunsporigen Sphaeropsideen, Ann. Mycol., ll(l):44-53.

1913. (A discussion of a few genera of this group.)
Petrak, F.: Mycologische Notizen, V, 187. Coniotliyrinula n.g., Ann. Mycol.,

21(1-2) :2-8. 1923. (Descriptions of genera segregated from Coniothyrium,

and key.)
Petrak, F., and H. Sydow: Die Gattungen der Pyrenomyzeten, Sphaeropsideen

und Melanconieen: Teil I. Die phaeosporen Sphaeropsideen und die Gattung

Macrophoma, Repertorium Specierum Novarum Regni Vegetabilis Beihefte,

42(1-3) :1-551. 1926-1927.
Diedicke, H.: Die Gattung Septoria, Ann. Mycol., 10(5) :478-487. 1912.
Martin, George: Enumeration and description of the Septoriae of North

America, /. Mycology, 3(4) :37-41, (5) :49-53, (6) :61-69, (7) :73-82, (8) :85-94.

1887.
Uppal, B. N.: a provisional list of the species of Septoria from Iowa, Proc. Iowa

Acad. Sci., 32:189-199. 1925. (List with descriptions and spore measure-
ments, also host index.)
Garman, Philip, and F. L. Stevens: The genus Septoria presented in tabulation

with discussion. Trans. Illinois Acad. Sci., 13:176-219. 1920.
Sprague, Roderick: The genus Phaeoseptoria on grasses in the Western Hemi-
sphere, Mycologia, 35(4):483-491. Figs. 1-2. 1943.
: The status of Septoria alopecuri and some related species, ibid., 35(3) :259-

263. Fig. 1. 1943.
LiNDER, David H. : New species of Sphaeropsidales and Melanconiales, Mycologia,

35(5) :495-502. 1 fig. 1943.
Fetch, T.: British Nectrioideae and allied genera, Brit. Mycol. Soc. Trans.,

26(1-2) :53-70. 1943.
Diedicke, H.: Die Leptostromaceen, Ann. Mycol, 11(2):172-184. Figs. 1-10.

1913. (A discussion of some of the genera and species of this family and of the

Pycnothyriaceae segregated from it.)
Arnaud, G.: Les Ast&inees, Ann. ecole nat. agr. Montpellier, N.S., 16:1-288. Pis.

1-53. Figs. 1-22. 3 maps. 1918. (Pages 205 to 220 and pis. 46-50 are devoted

to Asterinoid pycnidioid fungi. A key is given to the genera and these are

described.)
ViEGAS, A. P.: Algunos fungos do Brasil: XL Fungi imperfect!, Bragantia,

5(12):715-780. 1945. (Sphaeropsidales.)

750 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OP FUNGI

List 48. Melanconiales

Ellis, J. B., and B. M. Everhart: The North American species of Gloeosporium,

/. Mycologxj, 1(9):109-119. 1885.

: North American species of Cylindrosporium, ihid., 1(10):126-128. 1885.

: Additions to Cercospora, Gloeosporium and Cylindrosporium, ihid.,

3(2):13-22. 1887.
Grove, W. B.: The British species of Melanconium, Roy. Botan. Gardens, Kew.

Bull. Misc. Inform., 1918:161-178. 1 plate. 1918.
Klebahn, H.: Beitrage zur Kenntnis der Fungi Imperfecti: III. Zur Kritik

einiger Pestalozzia-Arten, Mycolog. Centr., 4(1):1-19. Figs. 34-38. 1914.
Guba, E. F.: Monograph of the genus Pestalotia de Notaris, I, Phytopathology,

19(3) :91-232. PL 4. Figs. 1-7. 1929; II, Mycologia, 24(4):355-397. Figs. 1-4.

1932. (Part II contains a key to all the species whose descriptions are dis-
tributed in Parts I and II.)
Steyaert, R. L.: Contribution a I'^tude des Pestalotia du Congo beige, Bull.

Jard. Bot. Brux., 19(2):173-186. 3 pis. 1948.
Shen, C. I.: Species of Pestalozzia and Monochaetia in China, Contributions of

the Biological Laboratory of the Scientific Society of China, Botanical Series,

7(5):131-141. 2 figs. 1932.
Duke, Maud M.: The genera Vermicularia Fr. and CoUetotrichum Cda., Brit.

Mycol. Soc. Trans., 13(3-4) :156-184. 1 pi. U figs. 1928.
Edgerton, C. W. : The Melanconiales, Trans. Am. Microscopical Soc, 31:243-

265. Figs. 1-9. 1912.
VifiGAS, A. p.: Algunos fungos do Brasil: XII. Fungi imperfecti — Melanconiales,

Bragantia, 6(l):l-37. 11 pis. 2 figs. 1946.

List 49. Moniliales : Moniliaceae

(For the asporogenous (i.e., imperfect) yeasts see List 30.)
Actinoinycetes.

(This group of organisms is often considered to be intermediate between the
Bacteria and the Fungi. Other students consider them to be Imperfect Fungi, and
in that case they should be placed in or near to the Moniliaceae. They are accord-
ingly so placed in this list.)

Breed, Robert S.; E. G. D. Murray; and A. Parker Hetchens: Bergey's
Manual of Determinative Bacteriology, ed. 6., xvi + 1529 pp. Baltimore,
The Williams and Wilkins Co., 1948. (Pages 875-980 are devoted to Actino-
mycetales, divided into three families: Mycobacteriaceae, Actinomycetaceae,
and Streptoniycetaceae. Aside from the foregoing, most recent and extensive
work the following papers may prove helpful.)

Dreschler, Charles: Mori)hology of the genus Actinomyces, Botan. Gaz.,
67(l):65-83, (2):147-168. Pis. 2-9. 1919.

Waksman, S. a., and R. E. Curtis: The Actinomyces of the soil, Soil Sci.,
1:99-134. PZs. 1-3. Fig. 1. 191G.

LIST 49. moniliales:moniliaceae 751

: Cultural studies of species of Actinomyces, ibid., 8:71-207. Pis. 1-4.

1919.
DucHE, Jacques: Les Actinomyces du groupe albus, in Encyclop^die Mycol-

ogique, vol. 6, pp. 1-375. Pis. 1-4. Figs. 1-32. Paris, Paul Lechevalier et Fils,

1934.
Breed, R. S., and H. J. Conn: The nomenclature of the Actinomyceteae, /.

J5ac<., 4:583-602. 1919.
Erikson, D.: Morphology, cytology and taxonomy of the Actinomycetes, Ann.

Rev. Microbiol., 3. 1949.
Jensen, H. L. : Contributions to our knowledge of the Actinomycetales, I, Proc .

Linnean Soc. New South Wales, 56:79-98. 1931; II, ibid., 56:345-370. Pis.

19-20. 1931 (recognizes two families: Proactinomycetaceae and Actinomy-

cetaceae with 3 and 2 genera, respectively. Gives morphological and cultural

descriptions of about 20 species of Actinomyces and 6 of Proactinomyces) ; III.

Further observations on the genus Micromonospora, ibid., 57:173-180.

Illustrated. 1932; IV. The identity of certain species of Mycobacterium and

Proactinomyces, ibid., 57(5-6) :364-376. Illustrated. 1932.
DE Mello, F., et J. F. St. Antonio Fernandes: Revision des champignons

appartenants au genre Nocardia, Mem. Asiatic Soc. Bengal, 7:103-138. 1919.
Baldacci, E. : Revisione di alcune specie del genere Actinomyces, Mycopathologia,

1(1) :68-76. 1938.
: Introduzione alio studio degli Attinomiceti, ibid., 2:84-106. Pis. 13-15.

1939.
: Contributo alia sistematica degU Attinomiceti, IV. Sull' Actinomyces

melanosporus Kr., Atti. ist. botan. "Giovanni Briosi" e lab. crittogam. univ.

Pavia, ser. 10:321-329. 3 ^^s. 1938.
: Die Svstematik der Actinomyceten, Mycopathologia, 4(l):60-84. 1947.

Moniliaceae.

Constantin, J.: Les Muc^din^es simples. Mat^riaux pour I'histoire des champig-
nons, vol. 2, viii + 210 pp. Figs. 1-190. Paris, Librairie Paul Klincksieck,
1888. (Keys to the families and genera of the Moniliales (not to species).)

Fragoso, Romualdo Gonzales: Estudio sistemdtico de los Hifales de la flora
espanola, Mem. real acad. dene. Madrid. Ser. cienc. nat., 6:1-377. 85 figs.
1927.

Linder, David H.: A contribution towards a monograph of the genus Oidium,
Llo^jdia, 5(3) :165-207. 7 pis. 1942.

Berkhout, Christine Marie : De schimmelgeslachten Monilia, Oidium, Oospora
en Torula, pp. 1-77. Pis. 1-4. Doctor's Thesis, University of Utrecht.
Scheveningen, Edauw and Johannissen, 1923.

Pinkerton, M. Elizabeth: A comparative study of conidial formation in Cephal-
osporium and some related Hyphomycetes, Ann. Missouri Botan. Garden,
23(l):l-68. Pis. 1-6. 1936. (Keys to the human-parasitic species of Cephalo-
sporium and to related saprophytic genera, Clonostachys, Gliocladium,

etc.)
Baldacci, E.; R. Ciferri; e E. Vaccari: Revisione sistematica del genere

Malbranchea, Atti ist. botan. "Giovanni Briosi" e lab. crittogam. univ. Pavia,

ser. IV, 11:75-103. 15 figs. 1938.
SiMOES Barbosa, Frederico a.: Subsidios para o estudo parasitologico do

genero Hyalopus Corda, 1938, 62 pp. 6 pis. Recife, Imprensa Industrial, 1941.
Thom, Charles: The Penicillia, xiii + 644 pp. 98 j^^s. Baltimore, Williams and

Wilkins Co., 1930.

752 GUIDE TO THE LITEKATURB FOR THE IDENTIFICATION OP FUNGI

Raper, Kenneth B., and Charles Thom: A manual of the Penicillia, i-ix, 1-875

pp. 10 Colored plates. 172 Figs. Baltimore, Williams & Wilkins Co. 1949.
, AND Dorothy I. Fennell: New species of Penicillium, Mycologia,

40(5) :507-546. Figs. 1-11. 1948.
BiouRGE, P.: Les moisissures du groupe Penicillium Link, La Cellule, 33:1-331.

Col. pis. 1-13. Pis. 1-23. 1923. (An attempt at a monograph of this difficult

genus.)
Sopp, Olav Johan-Olsen: Monographie der Pilzgruppe Penicillium. Mit beson-

derer Berlicksichtigung der in Norwegen gefundenen Arten, Videnskapsel-

skapets-Skrifter I. Mat.-naturv. Klasse, 1912(11) :l-208. Pis. 1-23. Fig. I.

1912.
Westling, R. : Uber die griinen Spezies der Gattung Penicillium. Versuch einer

Monographie, Arkiv for Botanik, 11(1):1-156. Figs. 1-81. 1911.
WoLTJE, Wilhelm: Unterscheidung einiger Penicillium-species nach physiol-

ogischen Merkmalen, Centr. Bakt. Parasitenk., Zweite Abt., 48:97-130. 1918.
Shih, You-Kuang: The Penicillium from Wuchang, Central China, Trans.

Sapporo Natural History Soc, 14(4) :286-296. PL 12. 1936.
NiETHAMMER, Anneliese : Zur Systematik der Pilzgruppe Penicillium Link: L

Mitteilung: Die Untersektion Radiata (in der Sektion Velutina der Asym-

metrica), Arch. Mikrobiol., 14(l):46-62. Q figs. 1948.
Petch, T.: Gliocladium, Brit. Mycol. Soc. Trans., 22:257-263. Figs. 1-2. 1938-39.
Thom, Charles, and Kenneth B. Raper: A Manual of the Aspergilli, ix + 373

pp. 7 col. pis. 7Qfigs. Baltimore, Williams and Wilkins Co., 1945.
Blochwitz, Adalbert: Die Gattung Aspergillus: I. Neue Spezies, Diagnosen,

Synonyme, Ann. Mycol., 27:205-240. PI. 3. 1929. (Includes a key to the

recognized species of the genus.)
: Die Gattung Aspergillus: IL Neue Spezies, Synonyme und Nachtrage,

ibid., 31(1-2) :73-83. 1933.
: Die Gattung Aspergillus: III. Neue Spezies, Varianten und Mutanten

der Konidienfarbe, Synonyme und interessante Standorte, ibid., 32(1-2) :83-

89. 1934.
■: Die Gattung Aspergillus: IV. Neue Arten, Synonyme, Varianten und

Mutationen, ibid., 33(3-4) :238-250. 1935.

Neill, J. C: The mould fungi of New Zealand: I. The genus Penicillium, Trans.
Proc. Roy. Soc. New Zealand, 67:101-112. Pis. 22-24. 1937; II. The genus
Aspergillus, ibid., 69:237-264. 1939. (Keys and descriptions and illustrations
of all species of Penicillium recognized in New Zealand and keys and descrip-
tions of all 18 species of Aspergillus recognized as valid, of which 12 are
known in New Zealand. Sixteen doubtful species are also described.)

Shih, Y. K.: A taxonomic study of the genus Aspergillus around Wuchang,
Central China (Hyphomycetes), Lingnan Sci. J., 15(3):365-378. PI. 16.
(4):607-612. 1936.

MossERAY, Raoul: Les Aspergillus de la section "Niger" Thom and Church,
La Cellule, 43(2):203-28(3. 4 pis. 1934.

Ellis, J. B., and B. M. Everhart: North American species of Ramularia with
descriptions of the species, /. Mycology, l(6):73-83. 1885.

, and : Supplementary notes on Ramularia, ibid., 1(8):102. 1885.

, and : Additions to Ramularia and Cercospora, ibid., 4(1) :l-7. 1888.

SuMSTiNE, David Ross: Studies in North American Hyphomycetes: I. The genera
Rhinotrichum and Olpitrichum, Mycologia, 3(2):45-56. Pis. 37-39. 1911;
II. The tribe Oosporeae, ibid., 5(2):45-6i. Pis. 82-84. 1913.

Hansford, C. G.: The genus Eriomycopsis, Bothalia, 4(2) :464-472. 17 figs. 1942.
("Hyphomycetea Mucedinea, Macronemea.")

LIST 50. moniliales:dematiaceae 753

WoLLENWEBER, H. W. : Ramularia, Mycosphaerella, Nectria, Calonectria. Eine
morphologisch-pathologische Studie zur Abgrenzung von Pilzgruppen mit
cylindrischen und sichelformigen Konidienformen, Phytopathology, 3(4):197-
242. Pis. 20-22. 1913.

List 50. Moniliales : Dematiaceae

LiNDER, David H.: New Venezuela Fungi Imperfecti, Mycologia, 29(6):656-664.
Qfigs. 1937. (Includes a key for the differentiation of all the 11 known species
of Periconia that possess rough globose spores.)

Mason, E. W.: On species of the genus Nigrospora Zimmermann recorded on
Monocotyledons, Brit. Mycol. Soc. Trans., 12:152-165. PI. 15. 1927.

: Annotated account of fungi received at the Imperial Mycological Insti-
tute, List II, Fascicle 3(special part):101-144. Figs. 31-44. 1941. (A careful
study of the genera Monotospora Sacc. and Monotospora Corda, Torula
Pers., Gliomastix Guegen, Catenularia Grove, Sporocybe Fr., sensu Bon-
orden, Zygosporium.)

GoiDANicH, Gabriele: II genere di Ascomiceti Grosmannia G. Gold., Boll. staz.
patol. vegetale, N.S., 16(l):26-60. PI. 1. Figs. 1-19. 1936. (Description of the
genus Scopularia and its species connected with the ascomycetous genus
Grosmannia.)

VAN Beyma thoe Kingma, F. H.: Beschreibung der im Centraalbureau voor
Schimmelcultures vorhandenen Arten der Gattungen Phialophora Thaxter
und Margarinomyces laxa, nebst Schllissel zu ihrer Bestimmung, Antonie van
Leeuwenhoek J. Microbiol. SeroL, 9(1-2) :51-76. Figs. 1-15. 1943.

Baldacci, Elio: Un nuovo genere di micete parassito del pioppa PoUaccia
radiosa (Lib.) Baldacci e Ciferri. Revisione dei g. Stigmella e Stigmina, Atti
ist. botan. "Giovanni Briosi" e lab. crittogam. univ. Pavia, ser. IV, 10:55-72.
bfigs. 1938.

Neergaard, Paul: Danish species of Alternaria and Stemphylium. Taxonomy.
Parasitism. Economical significance, Communications from the Phytopatho-
logical Laboratory of J. E. Ohlsens Enke, Copenhagen, pp. 1-559. Figs.
1-157. Copenhagen, Einar Munksgaard, Publisher, 1945.

Groves, J. W., and A. J. Skolko: Notes on seed-borne fungi: I. Stemphylium,
Can. J. Research, C, 22 :190-199. Illustrated. 1944; II. Alternaria, ibid., 22 :217-
234. Illustrated. 1944; III. Curvularia, ibid., 23:94-104. Pis. 1-7, 1945.

Elliott, John A.: Taxonomic characters of the genera Alternaria and Macro-
sporium. Am. J. Botany, 4(8) :439-476. Pis. 19-20. 9 graphs. 1917.

Young, P. A. : Tabulation of Alternaria and Macrosporium, Mycologia, 21(3) :155-
166. 1929.

Drechsler, Charles: Some graminicolous species of Helminthosporium, J.
Agr. Research, 24:641-740. Pis. 1-33. 1923.

Stevens, F. L.: Some meliolicolous parasites from Porto Rico, Botan. Gaz.,
65(3) :227-249. Pis. 5-6. Figs. 1-5. 1918. (Contains keys to the Porto Rican
species of Arthrobotryum and Helminthosporium that occur on Meliola.)

Hughes, S. J.: Studies on microfungi. II. The genus Sporoschisma Berkeley &
Broome and a redescription of Helminthosporium rousselianum Montague,

754 GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Commonwealth Mycological Institute Mycological Papers, 31 :l-33. PL 1. Figs.

1-9. 1949.
Jacques, J. Emile: Studies in the genus Heterosporium, Contribs. inst. botan.

univ. Montreal No. 39:1-46. Pis. 1-6. 1941.
Linder, David H. : A monograph of the Helicosporous Fungi Imperfecti, Ann.

Missouri Botan. Garden, 16(3) :227-388. Pis. 12-31. Figs. 1-17. 1929.

: The genus Helicoceras, ibid., 18(1) :l-7. PL 1. 1931.

: Brief notes on the HeUcosporeae with descriptions of four new species,

ibid., 18(1):9-16. PZ. 2. 1931.
Ellis, J. B., and B. M. Everhart: Enumeration of the North American Cerco-

sporae, /. Mycology, l(l):17-24, (2):33-40, (4):49-56, (5):61-65. 1885.
,'and — : Additions to Cercospora, Gloeosporium and Cylindrosporium,

ibid., 3(2):13-22. 1887.

AND — ■ — — •: Additions to Ramularia and Cercospora, ibid., 4(1) :l-7. 1888.

Overholts, L. a.: Species of Cercospora on Smilax in the United States, Ann.

Missouri Botan. Garden, 14(4) :425-432. Pis. 40-41. 1927.
LiENEMAN, Catherine: A host index to the North American species of the genus

Cercospora, Ann. Missouri Botan. Garden, 16(l):l-52. 1929.
SoLHEiM, William G.: Morphological studies of the genus Cercospora, Illinois

Biological Monographs, 12(1) :l-84. Pis. 1-4. 1929. (Divides the genus into 21

sections listing the species under each and describing in detail the more

important species.)
, AND F. L. Stevens: Cercospora Studies: II. Some tropical Cercosporae,

Mycologia, 23(5) -.363-405. Figs. 1-12. 1931.
Davis, R. H. : The Cercospora leaf spot of rose caused by Mycosphaerella rosicola,

ibid., 30(3) :282-298. Figs. 1-7. 1938. (Contains a key to the four species of

Cercospora known to grow on Rosa.)
Savulescu, Trian, et T. Rayss: Les Cercospora parasites des feuilles de vigne

en Palestine, Rev. path, vegetale entomol. agr. France, 22(3):1-19. Pis. 1-6.

1935.
Ray, W. Winfield: Notes on Oklahoma Cercosporae, I, Mycologia, 33(2) :174-

177. 1941; II, ibid., 34(5) :555-562. 1942.
MtJLLER, Albert S., y Charles Chupp: Las Cercospora de Venezuela, Bol. soc.

venezolana cienc. nat., 8(52):33-59. 1942. (List and host index of 176 species

of Cercospora including descriptions of 29 species new to science.)
, Y : Cercosporae de Minas Geraes, Arquiv. inst. biol. vegetal,

l(3):213-220. 1935.

Y : Una segunda contribuigao a 's Cercosporae de Minas Geraes,

i6id., 3(l):91-98. 1936.
Chupp, Charles: Cercosporae. Reprinted from C. E. Chardon and R. A. Toro:

Mycological Exploration of Venezuela, pp. 241-255. Monographs of the Univ.

Puerto Rico, Series B, No. 2. 1934.
: Cercospora species and their host genera, Dept. Plant Pathol., New York

(Cornell) Agr. Expt. Sta. Mimeographed Circ, 23 pp. 1937.
-, AND David H. Linder: Notes on Chinese Cercosporae, Mycologia,

29(l):26-33. 1 fig. 1937.
Tai, T. L.: Cercosporae of China, I, Bull. Chinese Botan. Soc, 2:45-66. 5 pis.

1936; II, IJoydia, ll(l):36-56. Figs. 1-15. 1948.
ViEGAS, A. P.: Alguns fungos do Brasil: Cercospora, Bol. soc. brasil. agron. (Rio

de Janeiro), 8(1):1-160. 86 p/.s. 18 ^^s. 1945.
Yamamoto, Wataro: Cercospora from Formosa, I, Trans. Sapporo Natural

History Soc., 13:139-143. Figs. 1-3. 1934.

LIST 51. moniliales:tuberculariaceae, stilbellaceae 755

— : Cercospora-Arten aus Taiwan (Formosa), II, /. Tropical A^r., 6:599-608.
Figs. 1-4. 1934.

List 51. Moniliales : Tuberculariaceae, Stilbellaceae
and Mycelia Sterilia

Fusarium.

Appel, O., und H. W. Wollen WEBER : Grundlagen einer Monographie der
Ciattung Fusarium (Link), Arheiten aus der Kaiserlichen Anstalt fiir Land-
und Forshvirtschaft, 8:1-207. Pis. 1-3. Figs. 1-10. 1910. (This work is the
basis for most of the subsequent systematic work on this genus.)

WoLLENWEBER, H. W. : Fusaria autographice delineata, Ann. MijcoL, 15(1-2) :1-
56. 1917. Supplementary to this, 659 separate sheets of drawings.

: Fusarium-Monographie. Fungi parasitici etsaprophytici, I, Z.Parasitenk.,

3(3):269-516. Figs. 1-71. 1931; II, Centr. Bakt. Parasitenk. Abt. II, 106(5-
7):104-135, (8-10):171-202. 71 figs. 1943.

UND O. A. Reinking: Die Fusarien, ihre Beschreibung, Schadwirkung

und Bekam])fung, 355 pp. 9^^ figs. Berlin, Paul Parey, 1935.
Shkkbakoff, C. ]).: Fusaria (jf potatoes, dornell Univ. Agr. Expt. Sta. Mem.,

6:89-270. Col. pis. 1-7. Figs. 1-51. 1915.
Reinking, O. A., and H. W. Wollenweber: Tropical Fusaria, Philippine J.

Sci., 32(2):103-252. 6 pis. (2 colored). 47 figs. 1927.
DoiDGE, Ethel M.: Some South African Fusaria, Bothalia, 3(3):331-483. Pis.

1-4. Figs. 1-48. 1938.
BuGNicouRT, Francis: Les Fusarium et Cylindrocarpon de ITndochine, These de

doctorat es sciences, Paris. 208 pp. 10 pis. 36 figs. Paris, Paul Lechevalier,

1939.
Jamalainen, E. a.: Ueber die Fusarien Finnlands, I, Staatl. Landwirtsch. T er-

s'uchstdtigk. Veroffentlich., 122, 22 pp. 5 figs. Helsinki, 1943.
Snyder, W. C. : Notes on Fusaria of the section Martiella, Zentr. Bakt. Parasitenk.,

Abt. II, 91(8-10) :163-184. Illustrated. 1934.
^ AND H. N. Hansen: The species concept in Fusarium with reference to

section Martiella, Am. J. Botany, 28(9) :738-742. 1941.

^jjD ; The species concept in Fusarium with reference to Discolor

and other sections, ibid., 32(10) :657-666. 1945.
Miller, J. J.: Cultural and taxonomic studies on certain Fusaria: II. The tax-
onomic problem in Fusarium with particular reference to section Elegans,
Can. J. Research, C, 24:213-223. 1946. (Keys to 13 species based on the
features expressed on potato dextrose agar.)

Other Tuberculariaceae.

Hughes, S. J.: Studies in microfungi: I. The genus Fusariella Saccardo, My-
cological Papers. The Commonwealth Mycological histitute, 28:1-11. Figs. 1-5.

1949.

Preston, N. C: Observations on the genus Myrothecumi Tode: I. The three
classic species, Brit. Mycol. Soc. Trans., 26(3-4) :158-168. 2 pis. Text figs.
1943.

75G GUIDE TO THE LITERATURE FOR THE IDENTIFICATION OF FUNGI

Stevens, F. L., and Nora Dalby: New or noteworthy Porto Rican fungi, Mtj-
cologia, 11(1) :4-9. Pis. 2-3. 1919. (Gives a generic key to the Tuberculari-
aceae — Scolecosporae.)

Stilbellaceae.

VAN Zinderen-Bakker, E. M.: Stephanoma tetracoccum, Ann. MycoL, 32(1-
2):101-104. 1 fig. 1934. (Descriptions of all known species of the genus.)

Fetch, T. : The genus Endocalyx Berkeley and Broome, Ann. Botany, 22(87) :389-
400. P/. 24. 1908.

Mycelia Sterilia.

HoTsoN, J. W.: Notes on l)ull)iferous fungi with a key to the described species,
Botan. Gaz., 64(4) :265-284. Pis. 21-23. Figs. 1-6. 1917. (Chiefly Fapulospora.)

Shaw, F. J. F., and S. L. Ajrekar: The genus Rhizoctonia in India, Mem. Dept.
Agr. India. Botanical Series, 7:177-194. Pis. 1-6. 1915.

Matz, J.: The Rliizoctonias of Porto Rico; J. Dcpt. Agr. Porto Rico, 5:1-31. Pis.
1-28. 1921.

INDEX

Boldface folios indicate main discussion of topic.

Abe, 24

Abies, 395, 403, 405, 494
Absidia, 165
oerulea, 160
glnuca, 160, 163, 164
septula , 164
spi7iosa, 160
Abstoma, 553
Acaulopage, 179
Acer, 235, 589
rubrum, 235
Acervulus, 231, 263, 333, 334
Achlya, 106, 107, 108, 109, 110, 112,
130, 142
flagellata, 110
glomerata, 34

oblongata, var. globosa, 104
polyandra, 104
racemosa, 111
Achlyogeton, 95
Achlyogetonaceae, 50
/I cm, 482
Acrasiales, 23, 29, 30, 35, 36, 631

key to genera, 38
.4rr«s/s, 30
Actinodothis, 319, 354
Actinomyces, 179, 585, 603

6o?;is, 585
Actinomycetaceae, 585
Actinomycetales, 585
Actinonema, 231
Adams, 387

Aecial priniordivim, 386, 390
Aecidiospore, 381
Aecidium, 408, 572

berberidis, 18
Aeciosporo, 381, 388

origin of chains, 392
Aecium, 399

caeomoid, 400, 407, 408
cornute, 399, 407

Aecium — {Continued)
cupulato, 399, 400, 407
diffuse, 399, 408
hyphoid, 399, 400, 407
secondary, 399
types, 400
uredioid, 407

Aethalium, 28

Agaricaceae, 9, 10, 11, 12, 193,
469, 470, 480, 484, 494,

497, 498, 500, 501, 502,

505, 506, 507, 509, 510,
534, 540, 541, 543

keys to genera, 520
115, Agaricacces, 470

Agaricales, 335, 464, 469, 471,

498, 534
origin, 652

Agaricum, 11
fungus, 10

Agaricus, 10, 11, 502, 504, 507,
542
arvensis, 196, 498, 502, 509
campestris, 196, 369, 374, 498,

506, 509, 589
rodmani, 502

Ajello, 63

Alhuginaceae, 8, 127, 135, 136,
key to genera, 145

Albugo, 135, 139, 142
bliti, 136, 137
Candida, 135, 137, 139
i pomoeae-panduranae 138
portulacae, 133, 137, 138
tragopogonis, 137

Aleuria umbrina, 586

Aleuriospore, definition of, 575

Aleurodiscus, 475
amorphus, 474

Algae, blue-green, 217
green, 217

Alina, 317

Alisma, 65
757

372, 467,
495, 496,
503, 504,
511, 532,

473, 495,

509, 540,
501,502,

138, 139

758

INDEX

Alismataceae, 421

Alkaloids, 505

AUantosphaeriaceae, 279, 281, 282, 284

Allen, Miss, 14, 166, 312, 385, 388, 389,

390, 391, 397
AUomyces, 4, 42, 78, 79, 81, 82, 84, 86, 91,
107

arbusculus, 85

cystogenus, 81, 86

javanicus, 85
Alnus, 243
Alpova, 539, 547
Alternaria, 194, 596

tenuis, 597
Alternation of generations, 79, 85, 86
Amanita, 10, 502, 503, 508

caesarea, 9, 502

muscaria, 503, 509

phalloides, 503

rubescens, 502

verna, 503
Amaranthus, 137
Amazonia, 309, 319, 354
Ames, L. M., 264, 268, 269, 277, 329, 375,

391
Ames, Miss, 14, 488
Ammiaceae, 143

Amoeba, 24, 178, 179, 182, 591, 593
Amoebidiaceae, 182
Amoebidium, 181, 182
Amorphomyces falagriae, 214
Amphiloma, 218
Amphisphaeriaceae, 278
Amygdalus persica, 243, 582
Ancylistes, 100, 175
Ancylistidaceae, 100, 175
Andrus, 389, 390
Angiocarpy, 496
Angiospermae, 2, 126, 127
Anisogametes, 6
Anisogamy, 143
Anisolpidiaceae, 70, 71
Anisolpidium, 71

ectocarpii, 70, 71
Anisomyxa, 35
Annulus, 505, 509

simple, 509
Anteriorly uniflagellate fungi, key to

order, 43
Antherid, 6, 106

aniphigynous, 132, 133

diclinous, 129

hypogynous, 129

monoclinous, 129

paragynous, 132
Anthoceros, 419

Anthophyta, 2, 126, 127, 309, 382
Anlkostoma, 282, 284

Anthurus borealis, 544
Antibiotics, 9, 15, 602
Aphanodictyion, 115

papillatum, 115
Aphanomyces, 108, 110, 112, 114, 115

acinetophagus, 113

exoparasiticus, 113

phycophilus, 113
Aphanomycopsis, 102
Aphragmiuiti, 130
Aphyllophoracees, 470
Aphyllophorales, 464, 469, 471
Apinis, 115, 116
Apium graveolens, 598, 600
Aplanea, 110, 634
Aplanogametos, 6
Aplanospore, 5, 151
Apobasidium, 454
Apodachlya, 116, 119

brachynema, 107
Apodachyella, 116, 117
Apogamous development, 213
Apophysis, 57
Apostemidium, 230
Apothecium, 193, 196, 202, 210, 220, 262

angiocarpic, 204

gymnocarpic, 204
Appendage, 314, 315
Apple, bitter rot, 280

scab, 8
Appressoria, 310, 317
Arachniotus aureus, 324, 330

trachyspermus, 329

trisporus, 329, 330
Araiospora, 106, 117, 119

pulchra, 118
Arcangeliella, 532, 540
Archicarp, 205, 215
Archilegnia, 116
Arcyria, 28

Armillariella mellea, 4, 501, 502, 508, 602
Arnaud, 271, 308, 309, 314, 319, 320, 321,

322, 357
Aronescu, 269
Arrhcnatherum elaiius, 416
Arrliytidia, 451
Arthrobotrys, 589

conoides, 592
Arthrobotryum, 317
Arthronia, 220
Arthropoda, 3, 180
Artlmr, 381, 396, 399, 402, 407, 425
Arundinelia, 182
Arundinula capitata, 181, 182
Arundinulaceae, 182
Aschersonia, 580
Ascobolus, 226, 227

carbonarius, 224, 225, 226

INDEX

759

Ascobolus — {Contin ued)
fiirfuraceus, 220
immersus, 227
niagnificus, 225, 227
stercorarius, 204
Ascocarp, 236, 237
Ascochyta, 579
dianthi, 578
pi si, 298, 580
Ascocortieiaceae, 241, 243, 039
Ascocorticium, 243, 458, 510, 647
Ascogenous hyphao, 208, 209, 210, 211,

220, 222, 313
Ascogonium, 203, 210, 220, 264
Ascohvmeniales, 205, 271
Ascoidea, 330, 338, 340, 352, 353

riibescens, 336, 340
Ascoideaceae, 338, 339, 340
Ascoloculares, 205, 271
Ascomycetales, 278

Ascomyceteae, 3, 6, 12, 14, 143, 172, 193,
200, 366, 368, 371, 380, 424, 455,
456, 458, 466, 510, 572, 570
Floridean ancestry, 030, 637
origin, 635
Ascophanvs, 111
airnetts, 195
granulatus, 211
granuliformis, 204
Ascosclerodenna, 330
Ascospore, discharge, 226

hat-shaped, 336, 337, 340, 341
Ascosporogenesis, 342
Asciis, 145
bilabiate, 225
dehiscence, 202
bilabiate, 202
inoperculate, 202
operculate, 202
development, 367
inoperculate, 219, 227
operculate, 227
Aseroe, 544

Ashby and Nowell, 350
Ashbyn, 345, 351

gossypii, 345
Ashworth, Miss, 397
Asparagus officinalis, 395
Aspergillaceae, 194, 323, 324, 325, 326,
329
key to genera, 356
Aspergillales, 239, 272, 274, 275, 277, 292,
299, 322, 331, 334, 586, 587
key to families, 355
Aspergillus, 11, 168, 323, 325, 326, 336,
573, 586, 587, 589, 602, 603
glaucus, 328
niger, 587

A spergillus — {Contin ued)

niveo-glaucus, 587

versicolor, 587
Aspidotus, 446
Aster, 405

Asteraceae, 140, 143, 421
Asterina, 294, 320

carneUiae, 294
Aster odon, 482
Asterophlyctis, 55
Asterosporales, 473
Asterostroina, 475
Asterostromella, 475
Astomella, 311, 314
Astraeus, 556

hygrometricus, 554, 555
A strot helium, 285
Atanasoff, 202
Atichia, 322

millardeti, 321
Atichiaceae, 308, 309, 321, 322, 334

key to genera, 355
Atkinson, 170, 212, 334, 452, 486, 498,

500, 507, 540, 039, 041
Auricularia, 10, 11, 438, 439, 443, 453, 510

auricvla-judae, 443

auricularis, 443, 444
Auriculariaceae, 9, 440, 442, 444
Am-iculariales, 197, 379, 381, 384, 430,
437, 438, 444, 446, 453, 455, 456,
556, 584

key to families and genera, 458
A uriscalpium, 482
Avena fatua, 416

sativa, 395, 409, 416, 418
Avocado, anthracnose, 280
Ayers, 151
Azygospore, 170, 174, 175, 176, 177

Bachmann, Miss, 220, 221, 224

Backus, 231

Bacteria, 2, 585

Baker, Miss, 439, 442

Baker, Mrak and Smith, 145

Ballistospore, 348, 349

Baranetzky, 329

Barnett, H. L., 378, 439, 453, 542

Barrage sexuel, 160, 377

Barrett, 98, 99

Bartlett, 66

de Bary, 1, 13, 22, 23, 133, 139, 237, 244,

311, 328, 334, 394, 633
Basidial nest, 539

primordium, 440
Basidiobolaceae, 175
Basidiobolus, 172, 174, 175, 178, 635, 636

ranarum, 174, 176

760

INDEX

Basidiomyceteae, 3, 6, 14, 160, 193, 195,
212, 237, 242, 313, 349, 350, 366,
436, 572, 602
Basidiophora, 140

entospora, 138
Basidiospore, 350, 366, 381

color, 468

discharge, 369

shape, 408

structure, 532
Basidium, 366, 381

chiastic, 470, 479, 495, 531

development, 367
' shape, 465

stichic, 469, 479, 531

tuning fork type, 458
Basisporium gallarum, 594
Battarrea, 532, 534, 557

digueti, 557

phalloides, 557
Bauch, 14, 378, 413, 414, 416, 417, 419
Bauhm, 9, 10
Baur, 220, 221
Baxter, 488
Bdellospora, 179
Bell morel, 228

Bender, 572, 576, 580, 581, 583, 584
Benjamin, 168
Bennett and Murray, 379
Bensaude, Mile., 14, 372, 373, 375
Berberis vulgaris, 18, 385, 388, 394, 395
Berdan, Miss, 64, 100
Bergman, 312

Berlese, 136, 138, 277, 279, 311, 346
Bernard, 543
Bertrandia, 501

Bessey, C. E., 196, 197, 380, 551
Bessey, E. A., 284
Beta vulgaris, 577, 598
Betts, 226, 352
Betula, 491

Biflagellatae, 35, 95, 030
Biflagellate fungi, key to orders, 43

zygotes, 52
Biggs, Miss, 201, 339
Biologic forms, 8
Bi polarity, 501
Bird's nest fungi, 548
Bisby, 15
Bishop, 106

Bjerkandera adusla, 488
Black scurf, 5
Blackinan, 286, 387, 391
Blackwell, Miss, 84, 131
Blackvvell, Watorhouse and Thompson,

135
Blakeslea, 155, 167

irispora, 158, 160

Blakeslee, 14, 157, 160
Blastocladia, 78, 79, 82, 84

globosa, 84

pringsheirmi, 83, 84
Blastocladiaceae, 79, 82, 83, 85
Blastocladiales, 49, 64, 78, 79, 81, 86, 119,
143, 633, 634

key to families and genera, 91

sexual reproduction, 79
Blaslodudidla, 78, 79, 82, 91

asperosperma, 83

cystogena, 81, 83, 84, 86

simplex, 82

stomophila, 82

stiihenii, 82, 84

variabilis, 82
Blastodendrion, 347
Blastomyces, 346
Blepharoplasts, 24
Blizzard, 506, 507
Blumer, 316
Boedijn, 236

Boedijn and Steinmann, 439, 442
Boletaceae, 9, 10, 11, 196, 464, 467, 469,
470, 473, 484, 495, 496, 497, 498,
507, 509, 511, 532

key to subfamilies and genera, 518
Boletales, 473
Boletineae, 496
Boletinus porosus, 497
Boletopsis, 489
Boletus, 10, 11, 12

colossus, 495

edulis, 495, 497

felleus, 497

luridus, 497

sphaerosporus, 496, 497
Bondarzew and Singer, 488, 489, 494, 516
Bonnier, 217
Boss, 414, 415
Bolryobasidium, 467, 473
Boiryodiplodia, 579, 580
Botryosphaeriaceae, 299
Botryotinia, 232

Juckeliana, 211, 212, 232
Botrylis, 8, 11, 232, 589

cinerea, 233, 575
Boudier, 227, 327
Bourdot and Galzin, 481, 484, 515
Bovista, 10, 552, 553

pliimbea, 552
Brand Fungi, 379
Brassica, 34, 47
Brassicaceae, 31, 141
Brefeld, 14, 152, 154, 159, 232, 328, 334,
380, 414, 418, 420, 444, 445, 450,
456, 474, 639
Brefeld and P'alck, 411

INDEX

7G1

Bremia, 141

laducae, 138, 139, 141

Brevilegnia diclina, 107, 634

Briosi and Cavara, 578, 579, 581, 590

Brodie, 195, 311, 368, 370, 371, 377

Brovius, 316

Broomeia, 553

Brown, A. M., 382, 388

Brown, H. B., 268

Brown, W. H., 210

Brunswik, 374, 378

Brussels sprouts, 34

Brj/opsis, 100

Bucholtz, 170, 171, 172, 237, 540

Buddin and Wakefield, 443

Budlia pundifoniiis, 220

Buisman, 277

BuUer, 155, 166, 195, 196, 226, 270, 348,
349, 367, 369, 370, 371, 373, 386,
390, 444, 493, 498, 500, 501, 510

BiilUra, 348, 349

von Bl'uen, 143, 144

Burgeff, 160

Burkholder, 333

Burnianniaceae, 2

Burnap, 557

Burt, 473, 475

Butler, 130, 132

Cabbage, 47
Cadophora, 576

obscura, 57(5
Caeoma, 408, 572
Cain, 338
Calbovista, 553
Caldesiella, 482
Calicium parietinuni, 220
Callitrichc, 34
Callose, 3, 181, 193, 350
Calocera, 451

cornea, 369
Calodon, 482
Calostoma, 556

cinnaharinum, 557
Calostoinataceae, 554, 556
Calvatia, 552

gigantea, 7, 196, 551, 552

sculpta, 553
Camp, 25
Caiiavalia, 334
Candida, 347

albicans, 347
Cannabis, 141
Canter, Miss, 54
Cantharellaeeae, 473, 476, 477, 479

key to genera, 512

Cantharellales, 480
Cantharellus, 470, 476, 499

floccosus, 477
Capillitium, 25, 325, 422, 531, 550
Capnodiaceae, 308, 309, 318, 319, 320

key to genera, 355
Capnodium, 320

citri, 320

salicinum, 320
Curdamine bellidifolia, 410
Carex, 419

canescens, 410
Carleton, 382
Carpogenic cell, 213
Carpomyceteae, 192, 379

coloring, 197

fruiting structures, 196

nomenclature, 197

structure, 192
Caryoganiy, 371
Cassytha, 2
Castanca dentata, 282
Castoreum, 553
Catastoma, 552

circuniscissuDi, 552, 553
Catenaria, 64, 78, 81

allotnycis, 81

anguillulae, 81
Catenariaceae, 79, 81
Catenarioideae, 64
Catenariopsis, 71
Catenoniyces, 45, 63, 64

persicinus, 64
Cutemilaria, 576
Caulocystidiuni, 466
Cayley, Miss, 24, 25
Cellular hypha, 3
Cellulin, 78, 87, 104, 116
Cellulose, 3, 26, 45, 134, 151, 181, 182,
193

reaction, 94, 98, 104, 127, 142, 143

wall, 100, 145
Cellis, 141

occidentalis, 310
Centrosome, 208

Cephalospniium, 277, 576, 585, 586, 600
Cephalothecium, 589

roseum, 589
Ceracea, 451
Ceramium, 100
Ceratiomyxa, 26

fndiculosa, 27
Ceratobasidiaceae, 455
CeratobasidiuDt, 455, 457, 466, 473

coniigerum, 456

sterigmaticLirn, 457
Ceratostomataceae, 277
Ceralostomclla, 267, 277, 278, 598

762

INDEX

Ceratostomella — (Continued)

coerulea, 267

-pluriannulata, 267
Cercosphaerella, 298
Ccrcospora, 573, 597

apii, 598

beticola, 598

cerasella, 298, 598

liriodendri, 265

zeae-maydis, 595
Cercosporella, 597, 598
Cerinomyces, 451
Ceriomyces, 468, 487
Cetraria islandica, 223
Ceuthospora, 578

ahietina, 579
Chaetocladium, 155, 167
Chaetomiaceae, 274, 275, 276, 323
Chaetomium, 275

aterrimum, 276

bostrychoides, 268

kunzeanum, 268
Chalara, 277, 576
Chalaropsis, 576
Chalk-brood, 352
('hamaeayce, 385
(,'liuniomxia, 532, 540, 543
Chara, 34
Chaze, 237

Cheilocystidium, 466, 499
Chiastobasidial development, 437, 451,

464, 477, 480, 499
Chilemyces, 314
Chitin, 3, 26, 31, 35, 45, 50, 134, 142, 143,

150, 151, 193
Chlamydopus, 558

meyenianus, 558
Chlamydospore, 5, 110, 143, 170, 171,
172, 179, 195, 329, 349, 380, 468,
487
Chlorochytrhim, 630
Chloivciboria aeruginosa, 197, 232
Chlorococcum, 217
("lilorophyceae, 215, 217
Chlorophyllum escidenlum, 504
Choanephora, 155, 167

conjunda, 158
Choanephoraceao, 158, 167, 168

key to genera, 185
Chondrogaster, 532
Christeuberry, 164
('hristensen, 409, 415
Clhristman, 391, 392, 399, 401
Cliruniosporium, 591
Chroococcus, 217
Chrysomyxa, 405
Chrysopsora, 384
Chrysothrix nolilangere, 193

Chupp, 33
Chytridiaceae, 54

Chytridiales, 6, 44, 49, 69, 78, 81, 95, 97,
116, 119, 143, 145, 197, 632, 633,
635

composition of cell wall, 45

eucarpic, monocentric, 54, 61

haploid chromosome number, 46

kej^ to families and genera, 71

life history, 45

phylogeny, 631

polycentric, 61
eucarpic, 60
Chj/tridiochlon's, 59
Chytridiuni, 59
Ciboria, 232
Ciborinia, 232
Ciccinobohis cesatii, 311
Cichoriaceae, 143
Ciferri and Redaelli, 347
Cintractia fischcri, 410
Circinella, 165

minor, 164

umbellata, 164
Citromyces, 326
CitrulluH vulgaris, (iOO
Citrus, 280, 334, 581
Cladochytriaceae, 61, 64, 69, 81, 635
Cladochytrium, 61

replicatum, 61

tenue, 63
Cladonia, 217
Cladophora, 100
Cladosporium, 194, 595

cucumerinunt, 595

fulvurn, 595, 596

herbarum, 595

Clamp connections, 195, 237, 349, 350,

372, 373, 380, 383, 409, 410, 415,

419, 421, 439, 448, 450, 453, 455,

468, 470, 487, 501, 508, 530, 557

development, 372

prevalence, 374

whorls, 373, 374
Clathraceae, 10, 534, 543, 544, 545
Clathrus, 10, 11, 544

ruber, 543, 544
Claussen, 207, 209, 210, 352
Claustula, 545
Claustulaceae, 545
Clavaria, 10, 11, 12, 451, 455, 479, 480

pistillaris, 479
Clavariaceae, 9, 11, 469, 470, 477, 478,
479, 483, 490, 511, 602

key to genera, 513
CLavariadelphus pistillaris, 478, 479
Clavariella, 480

subbotrytis, 479

INDEX

763

Clavariopsis aquatica, 574
Claviceps, 5, 289

purpurea, 286, 289, 290
Clavicipitaceae, 285, 287, 289, 290
Clavicorona, 480
Clavochytrium, 82
Clavulina, 480
Clemencet, 330
Clements, 196, 486
Clinton, 133, 418, 426
Clitopilus, 532, 543
Clonostachys, 588
Closterium, 175
Clusius, 9
Clypeus, 280
Cobb, 543

Coccidiascus legeri, 345
Coccidioides immitis, 145
Coccidioidomycosis, 145
Coccomyces, 231, 582
Cochlonema, 179
Coelomomyces, 78, 79

lativittatus, 81
Coelomomycetaceae, 79, 81
Coemansia, 156, 169

aciculifera, 161

erecta, 161
Coenocyte, 3, 151

tubular, 3, 192
Coenocytic hypha, 3, 127
Coker, 108, 111, 478, 479
Coker and Beers, 496, 518
Coker and Braxton, 115
Coker and Couch, 109, 115, 549, 554, 555
Coker and Matthews, 108, 110
Cokeromyces, 168

recurvatus, 168
Coleoptera, 215
Coleosporium, 384, 399, 402, 405, 408, 439

solidaginis, 401, 405

sonchi-arvensis, 405
Collema, 215, 216, 219, 222, 264

crispum, 221

pulposum, 220, 221
Collemodes, 219, 222, 224

bachmannianum, 220, 221
Colletotrichum, 280, 581, 582

gloeosporioides, 581, 599

malvarum, 583
CoUey, 383, 387, 391, 392
Collybia radicata, 502

velutipes 508
Coltricia perennis, 489
Columella, 152, 153, 154, 531

percurrent, 537, 540
Compatibility, 371

factors, 453

factors governing, 376

Compatibility — (Continued)

mutual, 269
Completoria, 175
Compositae, 143
Conard, 540
Conidiobolus, 172, 174

brefeldianus, 173
Conidiophore, 5

monopodia], 142

sympodial, 142
Conidium, 5, 126, 127, 129, 130, 137, 142,
155, 167, 169, 170, 172, 175, 193

acrogenous development, 194

alpha, 283

basigenous development, 194

beta, 283

catenulate, 142

chain, 194

endogenous, 326, 576

oidial mode of formation, 5
Coniophora, 475, 484

cerebella, 373

puteana, 373, 374
Coniophorella, 475, 484
Coniosporium, 591
Coniothyrium, 579

fuckelii, 579
Conjugate division, 208, 372, 373, 380
Conjugation tube, 95, 100, 101, 104, 107,

108, 112, 117, 126, 129, 132, 144
Connectives, 326
Cook, M. T., 50
Cook, W. R. I., 34
Cook and Nicholson, 97
Cook and Schwartz, 32, 34
Cooke, W. B., 488, 489, 516
Cooper, 107
Cooper and Porter, 132
Coprinus, 370, 374, 378, 466, 470, 499,
500, 507, 509, 510, 541

atramentarius, 500, 502, 505

comatus, 502, 504

ephemerus, 375

ephemerus forma bisporus, 375

fimetarius, 377

lagopus, 370, 371, 373, 376

micaceus, 502

sterquilinus, 373, 500
Copromyxa, 30
Corallofungus, 10
Coralloid, structure, 542

type, 533
Coralloides, 10, 11
Corda, 12
Cordyceps, 289, 598

militaris, 289

ophioglossoides, 289
Coremium, 194, 598

764

INDEX

Coriolellns serialis, 487
Coriolopsis trabea, 487
Coriolns versicolor, 489
Corner, 203, 204
Cornu, 88, 99, 129
Cortex, 216, 312, 313

primary, 239

secondary, 239
Corticiae, 458

Corticium, 338, 473, 475, 482, 489, 494,
510, 511

incrustans, 473

solani, 602

vagum var. solani, 465, 475, 602
Cortina, 505
Coryneum, 582

beijerinckii, 582
Cotner, 78, 88
Cotton, 322, 476, 479
Couch, 53, 56, 57, 64, 69, 78, 79, 81, 82,
98, 100, 102, 107, 113, 114, 115,
158, 173, 175, 440, 441, 443, 445,
447, 449
Couch and Dodge, 81, 82
Couch and Whiffen, 82, 83
Craigie, 14, 370, 388, 390
Cranberry, 53
Craterelhis, 476
Cronartium, 392, 394, 399, 402, 405

comptoniae, 387

flaccidum, 405

quercuum, 405

ribicola, 383, 386, 387, 390, 395, 409
Crosses, illegitimate, 377
Crotalia cintractiae-fischeri, 410
Crozier, 243, 373
Crucibulum, 548

vulgare, 531, 549
Cruciferae, 31
Cryptococcus, 346
Cryptogamia, 11

algae, 11

fungi, 11
Cryptoporus volvatus, 471
Ctenomyces, 329
Cucumis melo, 141

sativus, 141, 596
Cucurbitariaceae, 278, 299
Cucurbitaria, 299
Cudonia, 234

Cummins, 399, 401, 407, 540, 541, 545
Cunninghamella, 155, 167, 168
Curtis, Miss, 51, 53
Cuscuta, 2
Cutter, 160, 162
Cyathus, 548

stercoreus, 549

striaius, 531, 548

Cylindrosporium, 231, 582
Cymadothea trifolii, 595
Cyphella, 476, 484
Cystidium, 473, 487, 499, 537, 541

trabecular, 466, 499, 500

types of, 466
Cystobasidium, 441
Cystogenes, 86
Cystopage, 179
Cystopus, 135
Cystosori, 96, 97
Cystospora, 35, 402
Cytogamy, 210, 268, 371
Cytospora, 282, 578
Cyttaria, 225, 236

gunnii, 236
Cyttariaceae, 236

Dacrymyces, 366, 368, 449, 451, 457

deliqiiescens, 450

lutescens, 450
Dacrymycetaceae, 369, 450, 473

key to genera, 460
Dacrymycetales, 379, 437, 438, 449, 450,

458
Dacryomitra, 451
Dacryopinax, 451
Dactylaria, 591

brochophaga, 593
Dadylella, 591

ellipsospora, 592

tylopaga, 593
Dade, 278, 576
Daedalea, 494, 495

confragosa, 494
Daedaleopsis, 494
Dahlia, 589
Daldinia, 284

concentrica, 284
Dale, Miss, 328, 330
Daleomyces philUpsii, 225
Dangeard, 14, 30, 101, 160, 207, 210, 212,
312, 323, 328, 334, 368, 450, 452,
641
Daphnia, 345
Darwin, 227
Dasyscypha, 232
Dasyspora foveolata, 399, 400

gregaria, 399
Davis, 416
Debaryomyces, 344
De Ferry de la Bellone, 237, 373
Delacroix, 579
De Lamater, 324, 330
Dematiaceae, 346, 584, 591, 593, 594,
595, 597, 599

INDEX

765

Dematiaceae — {Continued)

key to amerosporous genera, 618
key to dictyosporous genera, 620
key to didymosporous genera, 619
key to phragmosporous genera, 620
scolecosporous key, 621

Dendrogaster, 532

Dendrophoma, 577
convallariae, 578

Dendryphium, 596

Dennis, 327

Dentinum, 482

Dermateaceae, 230

Dermatitis verrucosa, 595

Derx, 328, 348

Desmidiaceae, 172, 175

Desmids, 54

Development, angiocarpic, 506, 507, 511,

531, 532

gymnocarpic, 506, 507, 511, 531, 532
pseudoangiocarpic, 506, 507, 511, 531,

532, 535, 540
unipileate, 540

Dianthaceae, 418
Dianthus caryophyllus, 395, 600
Diaporthaceae, 281, 282, 283, 286
Diaporthe, 282

arctii, 283, 580, 582
Diasporangiuvi, 135
Diatrypaceae, 280
Diatrype, 282, 285

virescens, 279
Diatrypella, 282
Dihlepharis, 88
Dicaryon, 208, 237
Dicaryon cells, 350

hypha, 241

phase, 4, 6, 383

stage, 380
Dicheirinia, 408

binata, 407
Dickinson, 413, 414, 415, 416
Dicotyledoneae, 2
Dicr anaphora, 153, 163, 166, 635

fidva, 157
Dictydium cancellatum, 28
Didyocephalos, 558

attenuatus, 558

curvatus, 558
Didyophora, 544

duplicata, 544, 545

indusiata, 544, 546
Didyostelium, 30

discoideum, 29

mucoroides, 29
Didyuchus, 106, 110, 114

missouriensis, 115

polysporus, 1 15

Diddens, 346

Diddens and Lodder, 336, 359
Didymellina iridis, 596
Didymium difforme, 24, 27

squamulosum, 25, 26
Didymodadium, 590

Dietel, 379, 396, 402, 403, 407, 418, 420
Dillenius, 10
Dimorphism, 106, 109
Diplanetism, 104, 106
Diplocarpon, 230

earlianum, 231

soraueri, 580
Diplodadium, 589
Diplocystis, 553
Diplodia, 580
Diploid colony, illegitimate, 342

nucleus, 161
Diploidization, 7, 370, 487
Diplophlydis, 60

intestina, 60
Dipodascus, 336, 338, 352, 353

albidus, 339, 340

iininudeatus, 201, 339, 341, 641
Diptera, 215
Discella, 581 "
Disciseda, 552

Candida, 552, 553
Discomycetes, 16, 200, 215, 227, 273, 292

key to orders, 244
Discula, 280, 581
Disjunctor, 137, 193, 388
Disk lichens, 215
Dispira, 168

americana, 159

cornuta, 151
Dissophora, 170

decumbens, 169
Doassansia, 411, 421

sagittariae, 420
Doassansiopsis, 421

martianoffiana, 420
Dobbs, 157, 166

Dodge, B. O., 14, 224, 225, 268, 269, 270,
386, 391, 398, 415, 549, 586, 639
Dodge, B. O., and Gaiser, 398
Dodge, C. W., 9, 330, 347, 348, 540
Doidge, Miss, 320

Donk, 475, 480, 481, 486, 489, 490, 494
Dothideaceae, 290, 291, 309
Dothideales, 271, 290, 295, 458, 595

key to families, 302
Dothiora, 296
Dothioraceae, 295
Doty, 480, 513
Douglas, 507
Dous and Ziegenspeck, 3
Dowding, Miss, 268, 270

766

INDEX

Drayton, 14, 211, 224, 226, 233, 391
Drechsler, 129, 172, 178, 179, 180, 182,

297, 585, 589, 591, 593
Drepanopeziza rihis, 230
Drosophila, 15, 345
Dryodon, 483
Dudresnaya, 268
Duemling et al., 603
Durand, 227, 233, 234
Dutch elm disease, 278

Earthstars, 553
Eccrina, 181
Eccrinaceae, 181, 182
Eccrinales, 150, 180, 182, 634

key to families, 187
Echinodontium, 482, 484, 494
Edocarpus, 70
Ectostroma, 281
Edrogella, 104
Ectrogellaceae, 94, 104
Eddins, 416
Edgerton, 268
Edson, 128, 130
Eel grass, 30
Eftimiu, 480
Eichleriella, 455
Eidam, 176, 329
Elaphomyces, 330
Elaphomycetaceae, 325, 330, 334

key to genera, 357
Elasmomyces, 532, 534, 540
Elaters, 557
Elfving, 217
ElUott, E. W., 24
EUiott, J. A., 266, 267
Ellis and Everhart, 277, 283, 318, 326
Ellison, 24, 32, 631
Elrod and Blanchard, 484
Elrod and Snell, 496
Elsinoe, 334, 583

ampelina, 334

piri, 334

veneta, 333, 334
Elvela, 12
Emerson, 85, 86
Emmons and Carri6n, 595
Empusa, 17, 175
Endochytrium, 60
Endocochlus, 178, 179

aster oldes, 180
Endoconidia, 182, 576
Endoconidium, 576
Endodesmidium, 54
Endogenous oospore, 89
Endogonaceae, 165, 170, 171, 172

Endogonaceae — (Continued)

key to genera, 186
Endogone, 163, 171, 172

fasciculata, 170

lactiflua, 170, 171, 172

malleola, 170, 171

occidentalis, 170, 171

pisiformis, 171

reniformis, 171

sphagnophila, 170, 171, 172
Endomyces, 336, 337, 340, 344, 345, 352,
353, 637

magnusii, 338
Endomycetaceae, 336, 337, 338, 345, 350
Endomycoideae, 345
Endomycopsis, 337, 345

albicans, 337, 347
Endo-operculate exit tubes, 64
Endophyllum, 396, 398

euphorhiae-sylvaticae, 369, 398

sempervivi, 398
Endospore, 137, 201, 508
Endosporeae, 26, 27
Endothia, 282

parasitica, 282
Englerula effusa, 318
Englerulaceae, 308, 318, 319

key to genera, 354
Englerulaster, 320
Enterobryus, 181
■ elegans, 181
Entoloma, 504, 532
Entomophthora, 172, 175, 177
fresenii, 176

grylli, 177

muscae, 174, 176

sepulchralis, 176
Entomophthoraceae, 174, 176
Entomophthorales, 17, 100, 143, 150, 172,
173, 178, 180, 192, 193, 634, 635,
636

key to genera, 186
Entomosporium, 231

macvlatum, 580
Entophlyctaceae, 54, 59, 60, 61
Entophlyctis, 59, 60

vaucheriae, 60
Entostroma, 281
Entyloma, 410, 419
calendulae, 419
dahliae, 421
Eocronartium, 438, 440

muscicola, 440, 442
Epibasidium, 379, 381, 437, 438, 439,
440, 441, 443, 448, 453, 456, 457,
464
Epichloe, 287
Epicoccum, 601

INDEX

767

Epicoccuni — (Continued)

nigrum, 599

oryzae, 601
Epiplasm, 200, 208
Epispore, 137, 193, 200, 508
Epithecium, 238, 239, 273, 291
Epithele, 475
Eremascaceae, 337
Eremascoideae, 345
Eremascus, 345

albus, 337

fertilis, 337, 338
Eremosphaera, 630
Ereviothecium, 345, 351

ashbyii, 345
Ericaceae, 405
Eriksson, 141, 382
Erinaceus, 11, 12
Eriocladus, 480
Erodium cicutarium, 50
Erysiphaceae, 12, 194, 308, 309, 311, 313,
314, 315, 316, 317, 320, 324

key to genera, 353
Erysiphales, 16, 272, 307, 320, 321, 322,
323, 324, 587

key to families, 353
Erijsiphe, 307, 314

dehor acearum, 316, 317

graminis, 8, 309, 310, 311, 316, 317, 382

polygoni, 311, 312
Euallomyces, 85, 86, 91
Eubasidiae, 346, 378, 379, 390, 436, 437,

457, 458, 464, 473
Eucarpic, 42

genera, 46
Euglena, 50, 57, 99
Eumyceteae, 197
Euphorbia, 385
Eurotium, 325, 326

herhariorum, 328

insigne, 327, 587
Eury'chasma, 104
Eurychasinidium, 104
Eutypella, 282

Evolutionary tendencies, 142
Exciple, 219

proper, 219

thalloid, 219
Excipula, 581
Excipulaceae, 581

key to genera, 608
Excipulum, 203, 210, 219
Exidia, 453, 454
Exoascales, 241
Exoascus, 211, 243
Exobasidiaceae, 469, 480, 481, 594

key to genera, 511
Exobasidium, 464, 465, 468, 470, 480

Exobasidium — [Continued)

rhododendri, 480

vaccina, 480, 481

vexans, 480
Exogenous oospore, 89, 90

sperm, 214
Exospore, 144

Exosporeae, suborder, 26, 27
Exosporium, 601

Fabian, 336

Fabraea maculata, 580

Fagus silvatica, 340

Fair child, 176

Falck, 202, 226, 278, 485, 594

Famintzin and Woronin, 27

Fawcett, G. L., 582

Fawcett, H. S., 579

Fayed, 499, 508

Fedorintschik, 32, 34

Femsjonia, 451

Fenner, Miss, 167

Fermentation, 335

alcoholic, 337
Fimetaria, 275

fimicola, 195, 266
Fimetariaceae, 264, 269, 274, 275
Fink, 245

Fischer, Alfred, 99, 130
Fischer, Eduard, 237, 238, 239, 324, 325,
331, 422, 423, 532, 533, 534, 536,
539, 540, 545, 546, 547, 550, 553,
554, 559, 560, 565, 657
Fischer, G. W., 416, 417, 421
Fischer, G. W., and Holton, 416
Fistulina, 11, 484, 495

hepatica, 484
Fistulinaceae, 469, 473, 476, 484

key to genera, 515
Fitzgerald, 577
Fitzpatrick, H. M., 46, 127, 134, 143,

163, 175, 352, 440, 442, 531
Fitzpatrick, R. E., 241, 242
Flagellata, 630

Flagellum, tinsel type, 24, 46, 69, 71, 94,
105

types of, 629

whiplash type, 24, 45, 46, 69, 78, 94, 97,
98, 105
Flemmg, 602, 603
Flerov, 409, 415
Flint, 166
Flor, 382, 419
Florey, 603

Florideae, 2, 100, 195, 203, 212, 215, 220,
222, 262, 268, 424, 639

768

INDEX

Flowering plants, 126

Foex, 310

Fo7nes, 10, 11, 491, 492, 494

fomentarius, 492, 493

officinalis, 196

pinicola, 16
Fomiiopsis, 492

officinalis, 486, 492
Form families, 576

genus, 573
Forsberg, 389
Fragaria, 298, 601
Fraser, Miss, 224, 320
Fraxinus, 240, 497
Freeman and Johnson, 382
Frey, 266, 298
Fries, Elias, 12, 16, 227, 379, 469, 481,

482, 507
Fries, Nils, 531
Fuligo septica, 23, 26, 28
Fungi, definition, 2

genetics of, 9

holocarpic endobiotic, monocentric, 94

key to major groups, 18

number of, 1

phylogeny, 628

pigments, 197

saprophytic, 7
Fungi Imperfect!, 6, 12, 16, 263, 346, 572

key to form orders, 575

key to genera, 604

key to orders and families, 603
Fungi lamellati, 11

porosi, 11

pulverentes, 11

ramosi, 11
Fungoides, 10
Fungus, 9, 10, 11, 510
Fungus chitin, 634
Funiculus, 549

Fusarium, 7, 197, 287, 586, 600
Fusicladium dendriticrirn, 298
F usicoecum, 280

Gadd and Loos, 480
Gallowaya, 384
Gametangia, 32
Gamete, 6

male, 6
Ganoderma, 493

applanatum, 196, 486, 492, 493

curtisii, 493

lucidum, 493
Gasterella, 532, 534, 537, 539

lutophila, 538
Gasterellaceae, 537

Gasterellopsis, 532, 534, 537, 541
Gasteromyceteae, 16, 379, 454, 464, 471,
472, 501, 509, 511, 530

key to genera, 561

key to orders and families, 560

phylogenetic system, 656

phylogeny, 651
Gasiroboletus, 496
Gastrosporiaceae, 539
Gastrosporium, 539

Gaumann, 112, 141, 167, 212, 291, 292,
296, 299, 334, 381, 441, 442, 470,
480, 481, 641
Gaumann and Dodge, 157, 440, 488
Gauthier, Miss, 178
Gautieria, 532, 540, 543

graveolens, 531

plumbea, 542
Geaster, 11, 554
Geasteroides, 554
Geasteropsis, 554

Geastraceae, 534, 550, 553, 554, 555
Geastrum, 550, 554

rufescens, 554
Geitler, 216
Gelasinospora, 269, 275

tetrasperma, 270
Gemmae, 111, 126, 370
Genea, 238

cubispora, 17

hispidula, 238
Genistella, 178

ramosa, 177
Genistellaceae, 177, 178, 180
Geoglossaceae, 233
Geoglossum, 203, 234

glabrurn, 233
Geographic races, 416, 487
Geolegnia, 110
Geopyxis, 228

cacabus, 196, 225
Geotrichoides, 347
Germ pore, 508

Germination by repetition, 349, 350, 457
Gibberella, 287

saubinetii, 287

zeae, 287, 600
Giesenhagen, 48, 594
Gilbert, E., 454
Gilbert, E. J., 518
Gilbert, E. N., 437
Gilbert, Frank A., 24
Gilbert, H. C, 26
Gilkey, Miss, 17, 201, 225, 238, 239, 255,

331
Glaziella, 172
Gleba, 531

coralloid, 536

INDEX

769

G leba — {Continued)
lacunar, 536
multipileate, 536
unipileate, 536
Gliodadium, 327, 576, 587, 588

penicillioides, 587
Glischrodermataceae, 565
Gloeocystidium, 466, 473
Gloeocystidium, 475
Gloeoglossum, 234
Gloeophyllum, 494

saepiarium, 378, 487
Gloeosporium, 280, 573, 581, 582, 583
populi-albae, 581
ribis, 230, 582
Gloeotulasnella, 455, 456
Glomerella, 280, 573

cingulata, 268, 279, 280, 582
Glotzia centroptili, 178
Gnomonia, 573

veneta, 280
Gnomoniaceae, 279, 280, 573, 582
Goidanich, 601
Goldring, 167
Gomphidiaceae, 496
Gomphus, 476

floccosus, 477
Gonapodya, 78, 86

siliquaeformis, 87
Gonatobotryum, 589
Gonatorrhodiella, 589

parasitica, 588
Gonatorrhodurn, 589
Gonidia, 216

Gonirnochaete horridula, 182
Gonytrichum, 589
Goplana, 402, 408

dioscoreae, 384, 407
Gossypium, 345, 350, 600
Graff, 318, 320
Gramrnothele, 482
Grandinia, 482
Graphiola, 422, 423
phoenicis, 422
thaxteri, 422
Graphiolaceae, 411, 421, 422 '

key to genera, 427
Graphium, 278, 598
rigidurn, 599
wZmi, 278, 598
Greathouse, 277
Green, Miss, 226
Gregory, 139, 140, 166
Greis, 211, 237, 266, 268, 550
Griffiths, 264
Gr^yoto, 489

berkeleyi, 490
Grossularia, 310, 316, 395, 405

Grove, 166, 396
Groves and Skolko, 596, 597
Guepinia, 451
Guepiniopsis, 450, 451
Guignardia, 296, 598

bidwellii, 296
Guilliermond, 338, 342, 343, 344, 345,

349, 350, 351, 352
von Gutenburg, 50
Guttulina, 30

Gwynne-Vaughan, 207, 210, 211, 307
Gymnoascaceae, 324, 325, 328, 329

key to genera, 357
Gymnoascus reessii, 329
Gynvnoconia, 408

peckiana, 386, 389, 393, 395, 398, 408
Gyrnnosporangium, 11, 383, 388, 396, 408

clavariaeforme, 387

juniperi-virginianae, 395, 408

nootkatense, 396

sabinae, 394
Gyrocephalus, 455
Gyrodon meruUoides, 497, 498
Gyromitra, 228

esculenta, 229
Gyrophana, 485

lacrymans, 485
Gyrophora muhlenbergii, 218
Gyrophragmium, 534, 542

Hadley, 336

Hadromycosis, 589

Hadrotrichum, 594

Haerangiomycetes, 278

Haerangium, 278

Hainesia lythri, 601

Hanna, 14, 378, 388, 409, 412, 415, 421

Hanna and Popp, 416

Hansenula, 336, 340, 344

Hanson, 56, 60, 64

Haplographium, 591

Haploid nucleus, 161

Haplosporangium, 170

bisporale, 170
Haplosporella, 579
Haploirichuni, 586
Harder, 45, 79
Harpella, 178

rnelusinae, 177
Harpellaceae, 177, 178, 180
Harper, 14, 207, 209, 210, 311, 312, 313,

380
Harpochytrium, 59

viride, 59
Harter and Zaumeyer, 382
Hartig, 403

770

INDEX

Harvey, 110
Harveyella mirabilis, 2
Hashioka, 316
Hasselbring, 599
Hatch, 85

Hausschwamm, 485

Haustorium, 7, 132, 139, 196, 212, 217,
307

binucleate, 383

dicaryon, 383

monocaryon, 383

uninucleate, 383
Hedgcock, 599
Hedrick, 245
Heim, 196, 312, 313, 464, 471, 473, 495,

501, 507, 508, 509, 534, 540
Helianthus, 388

armuus, 395
Helicohasidium, 439, 443

candidum, 443

compadum, 442

purpureum, 439, 443
Helicocephalum, 172
Helicoceras, 591
Helicogloea, 439, 443

lagerheimi, 442
Helicovia, 591

perelegans, 593
Helicomyces, 591

scandens, 593
Helicoon auratum, 593
Helicostylum, 166
Heliscus, 575
Helminthosporiuni, 317, 596

teres, 296, 297, 596
Helotiaceae, 231, 234
Helotiales, 230
Helotium, 71, 231

schimperi, 419
Helvella, 228
Helvellaceae, 228, 233
Helvellales, 233
Hemiascus, 353
Hemibasidii, 379
Hemigaster, 532, 536, 540

Candidas, 535
Hemigastraceae, 535, 536
Hemisphaeriaceae, 295
Hemisphaeriales, 271, 272, 291, 292, 307,
319, 321, 322, 354, 580

key to families, 303
Hepaticae, 557
Hericium, 477, 483

coralloides, 483
Herpobasidium filicinum, 439, 441
Herrell, 603

Heterobasidiae, 16, 346, 349, 378, 379,
381, 436, 458, 464, 473, 487, 504

Heterobasidiae — (Continued)

and Teliosporeae, summary, 457

origin, 644
Heterochaeie, 455
Heterocontae, 629, 632
Heteroecism, 394
Heterosporeae, 350
Heterosporium, 596

gracile, 596
Heterotextus, 451
Heterothallic species, 106, 157, 162, 316,

352, 375, 376, 501
Heterothallism, 157, 226, 227, 375, 376
Higgins, 205, 207, 231, 265, 582
Higginsia, 582
Higher Fungi, 4, 143, 192

key to classes, 197

reproduction, 194, 196
Hirmer, 374

Hirneola auricula-judae, 443
Hirneolina, 455
Hirschhorn, 416

Hirschioporus abietinus, 487, 489
Hobsonia, 591

mirabilis, 593
Hoehnelomyces, 444
Hoerner, 141
von Hohnel, 205, 256, 271, 274, 292, 321,

322
Hohnk, 135
Holcus lanatus, 34
Holobasidiae, 437
Holobasidium, 455
Holocarpic, 42
Homma, 314, 316
HomothaUic species, 106, 157, 160, 162,

163, 376, 376, 501
Honey, 193

Honeydew, 307, 308, 320
Honig, 31

Hook, 208, 209, 211, 220, 243, 373
Hopkins, 151, 193
Hordeum sativum, 418
Hormiscium, 591
Hormodendron, 595
Hormones, 107
Horn, 104, 112
Hotson, 329, 330
Howard, 23, 24, 26
Howard and Currie, 25
Hugon, 504
Humaria, 227

granulata, 211
Humarina, 227
Humulus, 141

lupulus, 316
Huneycutt, Miss, 115
Huttig, 414, 415

INDEX

771

Hyalopsora, 403
Hyaloria, 451, 453, 454, 534
europaea, 453
pilacre, 453
Hyaloriaceae, 453
Hybrids in fungi, 15
Hydatinophagus, 113
Hydnaceae, 11, 469, 475, 477, 480, 481,
483, 484, 494, 511
key to genera, 514
Hydnangiaceae, 534, 540, 543, 550
Hydnangium, 532, 540
Hydnellum, 482
Hydnes, 481
Hydnochaete, 482
Hydnocystis, 238
Hydnotria, 17

ciibispora, 17
Hydnum, 12, 482, 602

imbricahivi, 483
Hydrodictyon, 71
Hydrophora, 165
Hymenella, 576
Hymenial cavities, 531, 537
Hymenium, 203, 379, 436, 464, 465,
499
angiocarpic, 471, 495, 505
gymnocarpic, 471, 495, 505
monascous, 295
primordium, 505
pseudoangiocarpic, 471, 495, 505
structure, 465
Hymenochaete, 466, 475, 487

cacao, 476
Hymenogaster, 532, 537
rehsteineri, 537, 538
tener, 538
Hymenogastraceae, 534, 537, 538, 539,

640, 550
Hymenogastrales, 535, 536, 538, 546,

550, 552
Hymenomyceteae, 12, 25, 379, 381, 458,
464
phylogeny, 651
Hymenoptera, 215
Hypha, 3

acrogenous, 243, 313
dematioid, 320
flexuous, 390
laticiferous, 499
moniliform, 320
perisporioid, 320
receptive, 391
Hyphal bodies, 172, 176
Hyphochytriaceae, 70, 71
Hyphochytriales, 46, 69, 95, 632, 633

key to families and genera, 74
Hyphochytrium catenoides, 70, 71

Hyphochyiriu m — - {Continued)

hydrodictii, 71

infestans, 71
Hyphomyceteae, 583
Hyphopodium, 317, 318, 319, 320
Hypobasidium, 381, 437, 438, 439, 440,

441, 443, 448, 456, 457, 464
Hypochnus, 467, 475

solani, 475
Hypocopra, 275
Hypocreaceae, 272, 285, 287
Hypocreales, 203, 271, 272, 275, 285, 299,
309, 320, 322, 580, 599, 600

key to families, 301
Hypoderma, 234
Hypodermataceae, 234, 240
Hypodermii, 16, 379
Hypomyces, 287
Hypostroma, 294
Hypothecium, 203, 219, 241
Hypoxylo7i, 268, 284, 287

marginatum, 283
Hysterangiaceae, 534, 542, 544
Hysterangium,, 543
Hysteriaceae, 240
Hysteriales, 215, 239, 278, 294, 597, 639

key to genera, 256
Hystermm insidens, 240
Hysterographium fraxini, 240

minutum, 240

Iceland moss, 223
Imperfect fungi, 572
Incompatibility, 269, 371

genetic factors, 375
Indusium, 544
Ingold, 574, 575, 616
Inoperculatae, 46, 225, 230
Inoperculate series, 45
International Botanical Congresses, 15
Introduction, 1
Ipomoea batatas, 138, 151
Irene, 319

echinata, 318
Irenina, 354
Irenopsis, 354
Iris, 596

Irpex, 477, 481, 482, 484, 486, 494
Irpiciporus, 482, 494
Isaria, 598
Isoachlya, 110
Isogametes, 6
Isogamy, 143
Itajahya, 544
Itersonilia, 348, 349, 350
Iwadara, 601
Ixechinus, 495

772

INDEX

Jaapia, 484

Jackson, 338, 396, 424, 441, 639, 644

J acobsonia, 236

Jaczewski, 635

Jahn, 26

Jane, 59

Jenkins, 334

Johnson, 417

Jola, 438, 439, 441

j'avensis, 442
Jolivette, Miss, 166
Jones, P. M., 34
Jones, S. G., 224, 235
Jones and Drechsler, 66, 67
Jones and Torrie, 139
Josserand, 504

Juel, 242, 438, 457, 469, 532, 536, 640
Jugasporaceae, 496
Juniperus, 408

virginiana, 395

Kabatiella, 481

Kaiser, 419

Kanouse, Miss, 108, 170, 171

Karakulin, 481

Karling, 34, 35, 39, 58, 61, 63, 69, 70, 74,

95, 96, 99, 100, 102
Karlingia, 45

rosea, 57
Karsten, 494, 508
Karyogamy, 210
Keene, Miss, 160
Keilin, 82
Keitt, 270
Kemper, 373, 374
Kevorkian, 107
Kharbush, 211, 480
Kickxella, 156, 169
Kickxellaceae, 156, 161, 169

key to genera, 186
Killermann, 454, 474, 476, 479, 481, 488,

507
Killian, 266, 288, 293, 298, 422, 423
Klebahn, 132, 298, 575, 583
Kloeckera, 347
Kluyver, 349
Kniep, 14, 85, 372, 373, 375, 376, 378,

409, 413, 416
luiy, 223
Kolk, 409
Kordyana, 480
Krafczyk, 155
Kudo, 35
Kiihner, 193, 372, 466, 496, 506, 507

Kunkelia nitens, 385, 386, 389, 396, 398,

408, 415, 648
Kusano, 47, 48, 52, 53, 55
Kusanoopsis, 357

Laboulbeniales, 8, 193, 195, 202, 210, 212,
222, 262, 264

key to families, 244
Labyrinthomyxa, 31, 35
Lahyrinthula, 30, 31, 35

macrocystis, 30, 31
Labyrinthulales, 23, 29, 30, 35, 36, 631

key to genera, 39
Lachnea scuteUata, 227
Lachnocladium, 480
Ladariopsis, 532
Ladarius, 193, 501, 508, 532, 540

deliciosus, 502

volemus, 502
Ladiica saliva, 582
Lacunar type of sporocarp, 533
Laetiporus sulphur ezis, 490, 491
Lafar, 9
Lagena, 100, 102

radicicola, 103
Lagenidiaceae, 96, 100, 101, 103, 145, 175

key to genera, 120
Lagenidiales, 31, 35, 94, 107, 108, 119,
126, 143, 630, 632, 633

key to families and genera, 119
Lagenidium, 100, 102, 175

giganteum, 102

rabenhorstii, 101
Lagerheim, 88, 640
Lager heim and Patouillard, 454
Laibach, 89
Lamb, 397
Lambertella, 232
Lamellae,- 495
Langford, 270
Lanopila, 552

bicolor, 552
Lanopilaceae, 553
Larix, 232, 405, 492, 497
Lasiobotrys, 317
Latex, 501
Laticiferous hyphae, 501

tubes, 540
Latrostium, 71
Leach and Ryan, 409, 417
Leaf curl, 243
Lecanorales, 200, 210, 215, 225, 230, 264,

271, 272, 273, 291
Lecidea platycarpa, 218
Ledingham, 32, 33, 34, 35
Leger and Duboscq, 177, 178, 181, 182

INDEX

773

Leger and Gauthier, 177, 178
Lehfeldt, 370, 390
Leidy, 181
Lembosia, 294
Lemna, 71
Lemonniera, 575
Lendner, 160, 161, 164
Lentinus tigrinus, 506
Lenzites, 494, 495
betulinus, 494
saepiaria, 378, 487, 494
trabea, 487
Leontodon taraxacum, 17
Leotia, 234

chlorocephala, 233
Lepiota, 502
helveola, 504
morgani, 504
naiicina, 502

procera, 16, 502, 504, 509
rachodes, 504
Leptolegnia, 110, 114

caudata, 107, 114
Leptomitaceae, 95, 108, 111, 116, 119

key to genera, 122
Leptomitales, 35, 108, 630
Leptomitus, 116

ladeus, 111, 116
Leptoporus adustus, 378, 488
Leptosphaeria, 292, 296

coniothyrium, 579
Leptostroma, 580
Leptostromataceae, 234, 580

key to genera, 607
Leptothyrium acerinum, 579

pomi, 580
Leucoconis, 314
Leucocoprinus molybdites, 504

procerus, 502
Leucogaster, 539, 547
Leucoporus arcularius, 488

brumalis, 488
Leucosporae, 507

key to centrally stipitate genera, 521
Leucostoma, 282
Leveillula, 310, 314

taurica, 309
Levine, 507
Levisohn, 174
Liagora tetrasporifera, 643
Lichen, 11
Lichen acids, 218
aquatic, 218
crustose, 218
foliose, 218
fruticose, 218
pendent, 218, 219
Lichens, 9, 16, 194

Lichens — (Continued)

apothecial, key to families and genera,
245
Lichtenstein, 181, 182
Lichtheimia, 184 »

Lieneman, Miss, 265
Ligniera, 34
Lilliputia, 327, 331, 587

insignis, 327
Liniacinia, 320
Lindau, 227, 236, 273, 280, 307, 572, 588,

599, 604, 612, 622
Lindegren, 14, 269, 271, 335, 342, 346
Linder, 156, 161, 169, 315, 458, 590, 591,

593, 611, 612, 645, 646, 648
Lindfors, 396
Lindsay, 222

Ling- Young, 157, 160, 162, 164
Linnaeus, 10, 11, 15
Linum usitatissimum, 390, 405, 600
Liriodendron tulipifera, 265
Liro, 217, 410, 413, 414, 421
Lister, 23, 28

Literature cited, 18, 39, 75, 92, 123, 146,

187, 198, 257, 303, 360, 427, 461,

524, 567, 624, 658

Literature for identification of fungi, 660

Agaricaceae, Black-spored genera, 742

General, 730

Ochre- or Rust-spored genera, 740
Purple-spored genera, 741
Red- or Pink-spored genera, 739
White-spored genera, 735
Ascomyceteae, miscellaneous, 682
Aspergillales (Plectascales), 707
Asporogenous yeasts, 707
Atichiaceae, 705
Blastocladiales and Monoblephari-

dales, 675
Boletaceae, 729
Capnodiaceae, Englerulaceae, Tricho-

thyriaceae, Atichiaceae, 705
Chytridiales, 673
Clavariaceae, 723
Dothideales, 700
Eccrinales, 680
Englerulaceae, 705
Entomophthorales, 678
Erysiphaceae, 703
Exobasidiaceae, 721
Gasteromyceteae, General Works, 742
General Works, 662
Hemisphaeriales, 702
Heterobasidiae, 718
Host indexes, bibliographies, etc., 668
Hydnaceae, 724

Hymenogastrales, Sclerodermatales,
etc., 743

774

INDEX

Literature for identification of fungi —
{Continued)
"Hymenomyceteae" (General Works).
720
• Hyphochytriales, 675
Hypocreales, 699
Hysteriales, 695
Laboulbeniales, 682
Lagenidiales, 676
Lecanorales and Pyrenulales, 683
Lycoperdales (incl. Tulostomataceae

and Podaxaceae), 744
Melanconiales, 750
Meliolaceae (Perisporiaceae), 704
Meruliaceae, 725
Moniliales: Dematiaceae, 753
Moniliaceae, 751

Tuberculariaceae, Stilbellaceae, and
Mycelia Sterilia, 755
Monoblepharidales, 675
Mucorales, Entomophthorales, 678
Mycetozoa, 672
Myriangiaceae, 706

Nidulariales (incl. Arachniaceae), 746
Peronosporales, Protomycetales, 677
Pezizales Operculatae and Inopercu-

latae (incl. "Phacidiales"), 688
Phallales, 746

Phycomyceteae, general works, 673
Polyporaceae, 726
Protomycetales, 677
Pseudosphaeriales, 701
Pyrenulales, 683
Saccharomycetales and Asporogenous

yeasts, 707
Saprolegniales (incl. Leptomitales), 676
Sclerodermatales, 743
Sphaeriales, 695
Sphaeropsidales, 747
Taphrinales, 694
Thelephoraceae and Exobasidiaceae,

721
Trichothyriaceae, 705
Tuberales, 693
Uredinales, 712
Ustilaginales (incl. Grapliiolaceae),

709
Zoopagales, Eccrinales, 680
Litmus, 197, 219, 223
Ljungh, 196
Lloyd, 543

Lodder, Miss, 336, 346, 347, 348, 359
Lohman, 240

Lohwag, 242, 349, 532, 533, 543
Lohwag and Follmer, 484
Long, 554

Long and Plunkett, 558
Long and Stouffer, 544, 558

Longula, 542

texensis, 542
Lophiostoma, 278
Lophiostomataceae, 278, 279
Lophium mytilinum, 240
Lophodermium, 235
Lorenz, 531
Lower Fungi, 192, 196
Lunidospora, 575

curvida, 574
Lupo, Miss, 268
Lutman, 409
Luttrell, 293
Luzula midtiflora, 413
Lychnis, 411

Lycoperdaceae, 9, 11, 12, 534, 550, 551, 553
Lycoperdales, 535, 550
Ly coper don, 10, 11, 12, 533, 552

giganteum, 17

poculiforme, 18

pyriforme, 551
Lycopersicon, 350, 580

esculentum, 133, 600
Lysipenicillium, 327
Lysurus australiensis, 544

Maassen, 352

Macbride and Martin, 23, 27, 36
Macrochyirium, 60
Macrophoma, 577, 578, 580
Macrospore, 181, 182
Macrosporium, 596
Magnusia, 327

nitida, 329
Magnusiella, 243
Mains and Jackson, 382
Mainsia, 407
Maire, 469, 473, 477
Malaceae, 408

Malen^on, 193, 393, 540, 558
Malus, 280, 316, 334

sylvestris, 395
Mangifera, 280

indica, 581
Mangin, 151, 321, 322
Mango, anthracnose, 280
Manina, 483
Manuel, Miss, 345
Marasmius, 498

oreades, 501, 502
Marssonina, 231

pmiattoniana, 582
MartenseUa, 156, 169
Martin, 1, 437, 443, 450, 451, 453, 455,

456, 457, 458, 400, 461, 549
Mason, 481, 575, 594
Massee, 386

INDEX

775

Massospora, 172, 177

Matruchot, 587

Matthews, Miss, 128, 130

Mattirolo, 237, 273

Maublanc and Malengon, 558

McCranie, 81, 85, 86

McCuUough, 336

McDonough, 140

McGuire, 478

McWhorter, 217

Medicago saliva, 230

Medulla, 216

Megachytrium, 63, 64

Meiosis, 107, 137, 144, 160, 161, 162, 371

Meiotic division, 22

Melampsora, 402, 405

euphorbiae, 405

farlowii, 405

helioscopiae, 401

lini, 382, 385, 390, 405, 409

medusae, 405

rihesii-purpureae, 405
Melampsoraceae, 402, 403, 405, 407

key to genera, 425
Melampsoridium betulinum, 401
Melanconiaceae, 581, 583

key to genera, 609
Melanconiales, 575, 581, 583
Melanconidaceae, 280
Melanin, 197
Melanogaster, 539, 547
Melanogastraceae, 539, 540
Melanospora, 278, 285

chionea, 276
Melanosporaceae, 275, 276
Melanosporae, 507

key to centrally stipitate genera, 524
Melhus, 135, 139
Melin and Nannfeldt, 576
Meliola, 292, 308, 309, 317, 319, 320, 321

circinans, 318

corallina, 318
Meliolaceae, 308 314, 317, 318, 319

key to genera, 354
Meliolineae, 319
Mellor, Miss, 218
Melogrammataceae, 280
Meruliaceae, 475, 484, 485, 487

key to genera, 515
Merulius, 485, 486

domesticus, 485

lacrymans, 485
Mesophellia, 553
Mesophelliaceae, 553
Mesospore, 408
Mestas, 237
Metaphyses, 273, 292
Metula, 587

Mez, 633, 634
Micheli, 10, 11, 510
Microconidia, 233, 264, 297
Microglossum, 234
Micromonospora, 585
Micromyces, 53, 54

longispinosus, 53

zygogojiii, 53
Micromycopsis, 53, 54

cristata, 45, 54
Micropeltis, 295
Microsphaera, 314

alni, 311, 317

herheridis, 315

quercina, 314
Microspore, 181, 270
Microstroma, 481
Microthyriaceae, 292, 293, 294, 307, 319,

320, 321
Microthyrium, 294
Middleton, 127, 130
Milbrath, 138, 141
Mildew, powdery, 8, 309
Milesia, 403
Miller, Julian, 203, 272, 274, 277, 285,

292, 331, 332
Miller, L. W., 482, 514
von Minden, 49, 86, 99, 134, 135
Mindeniella, 108, 117, 119
Mitrula, 234
Mittman, Miss, 267
Mix, 244
Modicella, 171
Molds, blue, 325

green, 325

herbarium, 325

pink, 327

sooty, 320, 349
MoUer, 216, 220, 454
Mollisia, 230

Mollisiaceae, 230, 320, 573, 580, 582
Monascus, 323, 326, 327

ruber, 324
Monilia, 347

Moniliaceae, 574, 583, 584, 586, 587, 588,
590, 591, 592, 593, 597, 598, 599

key to amerosporous genera, 612
to hyalodidymous genera, 615
to phragmosporous genera, 615
to scolecosporous, 616
Moniliales, 240, 574, 575, 576, 583, 588

key to endosporeae, 610
to form families, 583
to helicosporous genera, 611
to staurosporous, 616
Monilinia, 193, 194, 232

fructicola, 232
Monoblepharella, 80, 89, 91

776

INDEX

Monoblepharella— (Continued)
elongata, 90
mexicana, 90
taylori, 90
Monoblepharidaceae, 86, 87, 88, 89, 90,

635
Monoblepharidales, 49, 78, 79, 86, 95,
119, 143, 633, 634

key to family and genera, 92
Monoblepharis, 3, 78, 79, 86, 88, 90, 91

insignis, 88, 89

macrandra, 88, 89

polymorpha, 89
Monocaryon phase, 4, 6, 383

stage, 380
Monocentric Chytridiales, 42, 46
Monocotyledoneae, 2
Monomorphic zoospore, 106
Monoplanetic zoospores, 106
Monospora, 336
Monosporella bicuspidata, 345
Monotropa, 2
Monsma, 104
Montagnea, 534, 642
Montagnellaceae, 291
Montemartini, 600
Morchella, 10, 11, 12, 203, 211, 228

conica, 229
Moreau, 160, 212, 222, 373, 391, 398
Morels, 228

Morenoella quercina, 293
Morgan, 553
Morse, Miss, 553, 560
Mortierella, 152, 163, 170, 172

rostafinskii, 152
Mortierellaceae, 152, 169, 172

key to genera, 185
Mosquitoes, 78, 81
Mougeotia, 53
Mounce, Miss, 14, 375
Mounce and Macrae, 378, 487
Mucor, 11, 12, 151, 152, 153, 163, 165,
166, 167, 168, 170, 352

hiemalis, 160, 162

mucedo, 154, 161, 162

proliferus, 153

ramannianus, 171

rouxianus, 151

type, sexual process, 162
Mucoraceae, 5, 154, 157, 164, 165, 170

key to genera, 183
Mucorales, 12, 69, 143, 160, 151, 152, 156,
157, 172, 175, 178, 179, 180, 192,
193, 226, 337, 346, 586, 634, 635

key to families and genera, 183
Mucronella, 482
Multipileate type, 533
Mundkur, 132, 411, 419

Murphy, 133

MurriU, 481, 488, 489, 496, 519
Musca domestica, 176
Musci, 218

Muscus Saxatilis vel Lichen, 9
Mushroom, 505
Mutinus, 544
caninus, 544
ravenelii, 544
Mycelia, self-sterile, compatible, 268
Mycglia Sterilia, 576, 601

key to genera, 623
Mycelium, 3
coenocytic, 42, 104, 150
color, 4
dicaryon, 383
diploidization, 371
intercellular, 385
intracellular, 385
monocaryon, 368, 370, 383
primary, 4, 368, 453
secondary, 4, 368, 453
Mycena, 193, 470, 501
Mycenastrum, 552

Mycetozoa, 1, 2, 10, 12, 14, 16, 22, 23, 45,
94, 192, 197, 557
ancestry, 36
key to orders, 36
phylogeny, 630
Mycobacteriaceae, 585
Mycoderma, 347
Mycogone perniciosa, 589
Mycology, 1
history, 9
medical, 9
technical, 9
Mycosphaerella, 210, 296, 298, 578, 591,
639
cerasella, 298, 598
fragariae, 16, 298, 591
pinodes, 298, 580
sentina, 298
tabifica, 298
tulasnei, 595
tulipiferae, 205, 206, 265
Mycosphaerellaceae, 280, 296
Mycotoruleae, 347
Mycotoruloideae, 34(), 347
Mycotypha, 155, 167
Mykes, 9
Myriangiaceae, 322, 325, 332, 333, 334

key to genera, 357
Myriangiales, 272, 292, 299, 309, 320,

331, 583
Myriangina, 357
Myrianginella, 357
Myriangium, 325, 332, 334
curtisii, 333

INDEX

777

Myriangium — {Continued)

duriaei, 202, 333
Myrioblepharis, 86

paradoxa, 86
Myriodiscus, 236
Myriogormm, 338
Myriostorna, 554
Myxamoeba, 22, 24
MyxocoUybia, 508
Myxogastrales, 23, 30, 35, 36

key to families and genera, 36
Myxogastres, 23
Myxomycetes, 1, 23
Myxophyceae, 215, 217
Myxosporium, 581, 582
Myzocytium, 100, 102

proliferum, 101

vermicola, 101

Nabel, 45, 79, 151

Nannfeldt, 203, 205, 212, 223, 227, 230,
231, 234, 240, 251, 254, 271, 272,
274, 275, 277, 291, 323, 582
Napicladiurn, 596
Narasinhan, 132
Naumov, 163, 166
Nauss, Miss, 35
Nectaromyces, 346
Nectaromycetaceae, 346
Nectria, 287

chmaharina, 287, 289, 600
Nectriaceae, 287, 289
Nectrioidaceae, 580
Neergaard, 596
Nematodes, 78, 79, 589, 591
Nematospora, 335, 344

coryli, 345

lycopersici, 345

phaseoli, 345
Nematosporangium, 130
Nematosporoideae, 345
Neovossia indica, 411, 419
Nephrochytrium, 60
Nero, 9

Net-plasmodium, 30, 31
Neuhoff, 438, 457
Neurospora, 15, 269, 275, 391

crassa, 271

sitophila, 269, 270

tetrasperma, 268, 270
Newcomer and KenKnight, 585
Newton, 375, 376, 389
Nidularia, 548
Nidulariaceae, 535, 548, 549
Nidulariales, 535, 548
Nidulariopsis, 549, 550, 555

van Niel, 349
Nienburg, 286
Nigropogon, 532
Nigrospora, 594

oryzae, 594

panici, 594
Nitzschia, 2
Nobles, Miss, 473
Nocardia, 585

Nomenclature, rules for, 15
Nostoc, 11, 216, 217, 219
Nothofagus, 226, 236
Noivakowskiella, 63

macrospora, 63
Nuclear cap, 79, 81
Nucleus, diploid, 6

privileged, 341
Nurse cells, 312, 313
Nyctalis, 501

asterophora, 468
Nyland, 439

Obelidium, 57

rnucronatum, 57
Ochrosporae, 507

key to centrally stipitate genera, 523
Octojuga, 532, 543
Odomyxa, 34, 35
Octophore, 278
Odontia, 482
Oedocephalurn, 586, 594
Oedogonium, 49
Oehm, 468
Oenothera, 53
Oersted, 394

Oidiophore, 368, 370, 371
Oidium, 194, 335, 337, 368, 370, 371
453, 468, 488, 501

binucleate, 370

uninucleate, 370
Oidium, 589

aureum, 590
Olive, 26, 29, 30, 396
Olpidiaceae, 46, 55, 97, 99, 100
Olpidiopsidaceae, 32, 46, 49, 50, 96, 97,
98, 102, 633

key to genera, 120
OUpidiopsis, 2, 98, 100

luxurians, 98

schenkiawi, 49, 99

varians, 98

vexans, 98
Olpidium, 47, 49, 71, 95

brassicae, 47

radicale, 45, 49

trifolii, 47

778

INDEX

Olpidiuni — {Continued)

irifolii, sexual reproduction, 47

viciae, 47, 48

sexual reproduction, 47
Omphalia chrysophylla, 506
Onygena, 330, 444

caprina, 331

equina, 331
Omygenaceae, 325, 330, 331
Oogone, 106, 213

aplerotic, 129

plerotic, 129
Oomyceteae, 143
Oort, 377

Oosphere, 112, 136
Oospora, 194, 584, 585

ladis, 584, 585

nicotianae, 585
Oospore, 6, 95, 126, 136, 163

exogenous, 90

parthenogenetic, 133
Opegrapha atra, 220

subsiderella, 220
Operculatae, forms, 46, 224, 230
Operculate series, 45
Operculum, 45, 225, 226
Ophiostoma, 277, 278, 598

adiposum, 267

coeruleum, 267

fimbriaium, 267

paradoxum, 278, 576

piceae, 267

pluriannidatum, 267

ulmi, 277, 278, 598
Ophiostomataceae, 277, 323
Orange, 280
Orcein, 197, 219, 223
Orchidaceae, 2
Orobanche, 2
Orton, 284
Oryza sativa, 591
Osborn, 33

Ostiole, 203, 225, 262, 551
Ostropa, 230
Ostropaceae, 230, 254
Ostropales, 230
Oudemans, 414
van Overeem, 174
Oxalis, 390, 391

stricta, 395
Oxycoccus macrocarpus, 50, 53, tSO
Oxydotitia, 482
Ozonium, 602

omnivorum, 602

Pachynia, 602
cocos, 487, 602

Pachysterigma fugax, 456
Padus, 243
Pady, 383, 385, 393
Palm, 137

Palm and Burk, 33, 34
Palmella, 285
Palmeter, 270
Pampolysporium, 317
Panaeolus, 510
Panicum, 55
Papulospora, 5
mytilina, 240
Paragyrodon sphaerosporus, 496, 497
Paramyces, 468
Paraphysis, 203, 208, 210, 220, 273, 319,

466, 499
Paraphysoid, 273
Parasite, balanced, 8

destructive, 8, 132

facultative, 7

obligatory, 8
Parasitism, 7, 193

balanced, 194

destructive, 193
Paravicini, 410, 413
Parenchyma, true, 195, 190
Parmularia, 240, 294
Parodiopsis stevensii, 314
Parr, Miss, 166

Parthenogenesis, 6, 107, 108, 112, 129
Patella, 227

scutellata, 227
Patouillard, 321, 322, 327, 436, 470, 472,

476
Patouillardina, 455
Patterson, 128
Paxillaceae, 496
Paxillus, 496, 497, 498
Peck, 12
Pectose, 3, 151
Pedogamy, 341
Pellicularia, 5, 457, 467, 473

filamentosa, 465, 475, 602

isabellina, 474
Peltomyces, 35
Penicillin, 602, 603

Penicillium, 194, 323, 325, 326, 327, 573,
576, 586, 588, 591, 598, 602

chrysogenum, 587, 603

crustaceum, 328

expansum, 587

frequentans, 587

luteum, 328

notaium, 326, 587, 603

vermiculatuni, 328
Peniophora, 475, 482
Perfect stage, 18, 572
Perforations in septa, 195

INDEX

779

Periconia, 594
Pericystaceae, 337, 350, 352
Pericystis, 352
alvei, 352
apis, 352
Peridermium, 408
Peridiole, 548
Peridium, 531

Periphysis, 203, 207, 274, 319
Periplasm, 100, 101, 106, 114, 116, 117,

119, 126, 129, 132, 136, 137, 142
Perisporiaceae, 307, 317
Perisporiales, 292, 295, 307, 587
Perisporium, 307

granmieuni, 307
Perithecium, 196, 202, 203, 262

typical, 272
Peronoplasmopara, 141
Peronospora, 139, 141, 142
ficariae, 138
parasitica, 141
spinaciae, 141
Peronosporaceae, 18, 127, 138, 139, 140,
141, 142, 172
key to genera, 145
Peronosporales, 35, 49, 95, 108, 119, 126,
135, 142, 143, 151, 193, 631, 633,
634
keys to families and genera, 145
Persea, 280

Persoon, 12, 16, 17, 18, 227, 469, 554, 555
Pestalotia, 582

versicolor, 583
Pestalozzia, 582
Petch, 202, 333
Petersen, H. E., 55, 135
Petersen, N. F., 411
Petersenia, 100
Pethybridge, 132

Petrak, 273, 284, 291, 292, 299, 319, 354
Petrak and Sydow, 573, 577, 579
Peziza, 10, 12, 228
badia, 228
cacabus, 196
pustulata, 586
repanda, 228
vesiculosa, 228
Pezizaceae, 10, 202, 203, 227, 234
Pezizales, 197, 200, 210, 215, 223, 224,
237, 238, 243, 271, 295, 332, 334,
335, 573, 580, 582, 602, 639
key to families and genera of in-
operculatae, 251
of operculatae, 249
Pezizella oenotherae, 601
Phacidiaceae, 234, 239, 240, 582
Phacidium, 234
Phaeoradulum, 482

Phaeoseptoria, 580

Phallaceae, 9, 534, 543, 544, 545, 546

Phallales, 532, 535, 543

Phallogaster, 543, 544

Phallus, 10, 11, 12, 544

impudicus, 544

ravenelii, 544
Phaseolus, 334

limensis, 133

vulgaris, 395, 409, 475
Phellinus, 487

igniarius, 486, 493
Phellodon, 482

Phellorinia, 532, 553, 557, 558, 560
Phellorinieae, 558
Phialide, 5, 576, 586

definition, 575
Phialophora, 576
Phlebia, 482, 484
Phleogena, 439, 444, 556

decorticata, 444, 445

faginea, 444
Phleogenaceae, 440, 444, 445
Phlogiotis, 455
Phlyctidiaceae, 55
Phlyctochytrium, 55, 71

hallii, 56
Phlyctorhiza, 60

endogena, 60
Pholiota adiposa, 508

aurivella, 368, 468, 501
Phoma, 297, 577, 580

betae, 298, 578

stenobothri, 577
Phomopsis, 282

citri, 579
Phragmidium, 383, 394, 395, 402, 407, 408

rubi, 383, 400

rubi-idaei, 406

speciosum, 392

violaceum, 387
Phragmobasidiae, 437
Phycomyces, 160, 163

blakesleanus, 161

microsporus, 161, 164

nitens, 160, 161, 163
Phycomyceteae, 3, 4, 5, 7, 143, 182, 192,
193, 195, 334, 350, 572, 585

higher, phylogeny, 634

holocarpic bi flagellate, 95

nonflagellate, key to orders, 44
Phycopsis, 322
Phylacteria, 476, 481
Phylacteries, 481
Phyllachora, 284

graminis, 284
Phyllachoraceae, 282, 284, 290
Phyllachorineae, 284

780

INDEX

Phylladinia, 310, 314
eleagni, 315
guttata, 309
Phyllosticta, 577

tabifica, 577, 578
Phyrnatotrichum, 589

omnivorum, 589, 602
Physalacria, 478
Physalospora, 298, 579

cydoniae, 298
Physarella oblonga, 23
Physarum nutans, 28

polycephalum, 23, 24, 25
Physcia, 222, 223, 264
Physiologic forms, 8, 382

race, 389
Physocladia, 61

obscura, 61
Physoderma, 64, 66
maculare, 65, 66
zeae-maydis, 65, 66
Physodermataceae, 61, 64, 65, 67
Phytophthora, 127, 130, 133, 134, 135,
142, 634
arecae, 132
cactorum, 131, 133
hirnalayensis, 132
infestans, 133
phaseoli, 133
stellata, 133
Pierce, 242
Pierson, 390
Piersonia, 239
Pietra fungaia, 490
Pigment, 197
Pilacre, 444
Pilacrella, 444
Pilaira, 165, 166
Pilat, 539

Pileocystidium, 466
Pileolaria, 407
Piline, 317
Pilobolaceae, 155, 156, 165

key to genera, 184
Pilobolus, 49, 153, 156, 163, 165, 166,
169, 170, 175, 636
crystallinus, 155
kleinii, 155, 636
longipes, 636
Pinkerton, Miss, 586
Pmus, 387, 405, 497

s^robws, 383, 386, 395, 409
Piptocephalidaceae, 156, 159, 168, 178

key to genera, 185
Piptocephalis, 163, 168, 170
cruciata, 159
freseniana, 159
Piptoporus betulinus, 491

Piricularia, 590, 591

oryzae, 590
PiVms, 334

Pisolithus, 535, 547, 548
Pistillaria, 478
Pityriasis capitis, 347
Pityrosporum, 346, 347
Placenta, 291
Planocyte, 22, 23, 42
Planogametes, 6
Planont, 79, 82
Planospore, 5, 66
Plasmatoonkosis, 130
Plasmodiophora, 31, 34, 36, 94, 631

brassicae, 31, 32, 33, 35
Plasmodiophoraceae, 94
Plasmodiophorales, 23, 31, 35, 630

key to genera, 39
Plasmodium, 22, 24, 35
Plasynopara, 139, 140, 141
Platanus, 280
Platycarpa, 439, 441
Platygloea, 443
Plectania, 227

coccinea, 228
Plectascales, 205, 272, 275, 322
Plectobasidial structure, 533
Plectobasidiales, 546
Plectospira, 114
Pleospora, 296

herbarum, 597
Pleosporaceae, 266, 280, 296, 298
Pleotrachelus, 49, 100

fulgens, 49
Pleurage, 200, 275, 391

anserina, 264
Pleurocystidium, 466, 499
Pleurotus ostreatus, 502

ulmarius, 502
Plicatura, 484

Plowright, 379, 380, 394, 420
Plum pockets, 243
Pluteus cervinus, 502, 504
Poa, 316

pratensis, 418
Poaceae, 140, 316, 421
Podaxaceae, 535, 550, 558, 559
Podaxis, 532, 550, 553, 559, 560

pistillaris, 559, 560
Podetium, 218
Podosphaera, 312, 314
leucotricha, 316
oxyacanthae, 316
Podospora, 275
Pogonomyces hydnaides, 491
Poisson, 181
Poitras, 129, 168
Polycentric series, 42, 46

INDEX

781

Polychytrium, 63
Polymyxa, 35
Polyphagus, 57, 59

euglenae, 46, 57
Polyporaceae, 8, 9, 10, 11, 467, 469, 470,
473, 477, 484, 486, 490, 491, 492,
493, 498, 508, 511

key to genera, 516
Polyporales, 464, 469, 471, 473, 474, 481,
485, 490, 491, 492, 493, 495

key to families, 472
Polyporus, 5, 11, 489, 490, 491, 586, 602

caudicinus, 490

squamosus, 196, 486, 490

tuberaster, 490
Polysphondylium, 30

violaceuni, 29
Polysticius, 488, 489, 491

abietinus, 487
Polystigma, 287

rubrurn, 286
Polystomella, 294

Polystomellaceae, 290, 294, 319, 320
Polythrincium, 595

trifolii, 595
Pontisma, 100
Poole, 243
Populus, 405
Pore, 495

fungi, 2
Pores, 481
Porta, 488, 489, 494
Porothelium, 484
Portulaca oleracea, 137
Posteriorly uniflagellate fungi, key to

orders, 43
Prillieux, 578

Prillieux and Delacroix, 595
Primordiuin, basidial, 451
Primula, 227
Pringsheim, 130
Proabsidia, 184
Proactinomyces, 585
Probasidium, 458, 646
Proliferation, 109, 130, 153, 341
Promycelium, 379, 380, 381, 384, 394,
438, 453

internal, 384, 405
Propagula, 308, 321, 322
Prosenchyma, 196
Prosorus, 52
Prospodium, 399, 408

plagiopus, 401, 407
Prosporangium, 54
Proteomyxa, 36
Profoabsidia, 184
Protoachlya, 115
Proto-aecidium, 386

Protococcus, 217, 285
Protocoronospora, 481 ■
Protodontia, 455
Protogaster, 334, 535, 536, 539

rhizophilus, 535
Protogastraceae, 534, 535, 536
Protogastrales, 535, 536, 537
Protohydnurn, 455

gelatinosuvi, 454
Protomerulius, 455
Protomyces, 143, 144

inundatus, 143

macrosporus, 144
Protomycetaceae, 143, 144, 172
Protomycetales, 126, 143, 145
Protomycopsis, 143

leucantherni, 144
Protozoa, 22, 35, 116, 630
Protubera, 543
Pru7ius, 231, 243, 316, 582

avium, 243

cerasus, 243

domestica, 286
Psalliota, 502
Pseudocolus, 544

javanicus, 543
Pseudocoprinus, 509, 510
Pseudohypha, 348
Pseudolpidiopsis, 49, 99
Pseudolpidiurn, 99
Pseudoparenchyma, 195, 196
Pseudoperonospora, 141, 142

celtidis, 141

cubensis, 141

humuli, 141
Pseudopeziza, 224, 230

medicaginis, 230

n'bzs, 230, 573, 582

<ri/o/w, 224
Pseudoplasmodium, 29, 30
Pseudosepta, 4, 42, 78, 84, 86
Pseudosphaeriaceae, 296, 297
Pseudosphaeriales, 223, 240, 271, 272,
273, 274, 278, 280, 284, 291, 295,
331, 332, 334, 578, 596, 597

key to families, 302
Pseudosphaerita, 50, 100

euglenae, 99
Pseudotrichogyne, 266
Pteridophyta, 382
Pterula, 480
Ptychogaster, 487
Puccinia, 11, 17, 394, 402, 407, 408

arenariae, 396

asparagi, 395, 408

atropae, 393

caricis grossulariata, 388

coronata, 383, 388, 391, 395, 408

782

INDEX

Puccinia — {Continued)

glumarum, 408

graminis, 14, 18, 385, 388, 389, 390,
394, 395, 406, 408
agrostidis, 389
secalis, 389
tritici, 8, 382, 389

helianthi, 382, 388, 390, 394, 395

iridis, 382

malvacearum, 394, 396, 397

pringsheimiana, 388

prostii, 397

rubigo-vera, 408

tritici, 382, 388, 395, 396

sorghi, 17, 382, 390, 391, 395, 408

triticina, 388, 389, 390

tumidipes, 383
Pucciniaceae, 369, 398, 405, 406

key to genera, 425
Pucciniastrum, 399, 402

goeppertianum, 403
Puff ball, 2, 7
Pycnidium, 194, 219, 263

definition, 575
Pycniospore, 386
Pycnium, 386

Pycnoporus cinnabarinus, 491
Pycnothyriaceae, 581
Pycnothyriales, 581
Pyrenomycetes, 16, 262, 332

ostiolate, key to orders, 299
Pyrenophora, 202, 296

teres, 296, 297, 596

trichostoma, 296
Pyrenula, 285
Pyrenulales, 223, 271, 272, 285

key to families, 302
Pyronema, 193, 203, 210, 211, 220, 224,
225, 227, 243, 293, 351

omphalodes, 207, 209

omphalodes var. inigneum, 210
Pyrrhosorus, 97

Pythiaceae, 95, 97, 98, 126, 127, 128, 131,
133, 134, 138, 633

key to genera, 145
Pythiella, 100, 126
Pythiogeton, 135

transversum, 134
Pythiomorpha, 135
Pythiopsis, 105, 106, 109, 110

cyniosa, 110
Pythium, 8, 95, 100, 102, 114, 127, 130,
132, 133, 134, 135, 142, 634

aphanidermatum , 130

debaryanum, 130

dictyosporum, 128

monospermuni, 130

polysporum, 129

Pythium — (Continued)
proliferurn, 127, 128
torulosum, 128

Quadripolar sexual phases, 501
Queletia mirabilis, 556
Quercus, 239, 293, 497
Quintanilha, 377

Rabenhorst, 13
Races, biologic, 382

geographic, 378
Raciborski, 322, 334
Radicula armor acia, 591
Radiigera, 553
Radulaspore, 589

definition of, 575
Radulum, 482
Rafflesia, 2
Ramaria, 480
Ramlow, 201, 224
Ramularia, 573, 590, 598

armoraciae, 591

rosea, 590

tulasnei, 298, 591
Ramularisphaerella, 298
Ranunculaceae, 421
Raper, 29, 107, 156
Raper and Thorn, 587
Ravenelia, 394, 402, 408

acaciae-micranthae, 407
Rawitscher, 414
Rea, Carleton, 476, 477, 485, 486, 515,

557, 565
Rea, P. M., 557
Receptacle, 543
Receptive hypha, 389
Red Seaweeds, 2, 97
Reduction division, 112
Reed, 316
Reesia, 71

amoeboides, 71
Rehsteiner, 533, 537
Reijnders, 507, 509
Reuake, 218
Renisberg, Miss, 478
Renn, 30
Reproduction, asexual, 5

sexual, 5
Respiratory hyphae, 224
Resting sporangia, 65, 66, 67, 79, 83, 84,

85, 86, 143, 144
Resting spore, 79, 95, 96, 99, 102, 181,
182

INDEX

783

Reticularia, 24, 26
Beticidomyxa, 35
Rhamnus, 395

frangula, 383
Rheosporangiuyn, 130
Rhinotrichum, 589

Rhipidiaceae, 108, 117, 118, 119, 126,
127, 142

key to genera, 122
Rhipidium, 117

amertcanum, 118
Rhizidiaceae, 45, 54, 55, 56, 58, 59, 61,

78, 91
Rhizidiomyces, 71

apophysatus, 69, 70
Rhizidiomycetaceae, 70, 71
Rhizoclosmatmm, 55
Rhizoctonia, 5, 475, 602

solani, 602
Rhizoid, 7, 81
Rhizomorph, 4, 195
Rhizomorpha , 602

subcorWcalis, 602
Rhizomycelium, 61, 66, 67, 71
Rhizophlydis, 57

petersenii, 57

rosea, 57
Rhizophydium, 55, 57, 65, 71, 91, 96, 104

coronuni, 56

couchii, 57

graminis, 55

ovatxim, 57
Rhizoplast, 24
Rhizopoda, 179, 630
Rhizopogon, 539, 540
Rhizopogonaceae, 540
Rhizopus, 162, 165

nigricans, 11, 57, 160
Rhizothyriaceae, 581
Rhodochytrium, 630
Rhodophyllus, 468, 501, 504, 532
Rhodosporae, 507

key to centrally stipitate genera, 523
Rhodotorula, 348
Rhodotorulaceae, 348
Rhombiella cardamines, 410
Rhopalomyces, 586
Rhysotheca, 139, 140, 141

australis, 138

r^ziicoZa, 139, 140, 141
Rhytisma, 235

acerinum, 235
Bzftes, 230, 395, 405

nigrum, 396
Riccia, 98

Rice, Miss, 383, 389
Robak, 487
Roberts, J. M., 63

Roberts, Miss Catherine, 244

Rochlin, 31

Rogers, 438, 454, 455, 457, 458, 473, 474,

648
Rogers and Jackson, 473, 475
Rolland, 193
Rosa, 395
Rosellinia, 275, 284

aquila, 27 Q

necatrix, 275
Rostowzew, 141
Rotifers, 113
Routien, 537, 539, 541
RozeJla, 45, 47, 49
Rozellopsis, 97
Rubus, 334, 383, 386, 395, 396, 398, 408,

579
Rumex, 34

Russula, 193, 501, 508, 540
Russulaceae, 473
Rusts, 8, 14, 194, 197, 382, 424

autoecious, 394, 395

heteroecious, 394, 395
control, 395

macrocyclic, 396

microcyclic, 396

origin, 644

stem, of wheat, 8
Rutstroemia, 232

Saccardo, 13, 507, 577
Saccharomyces, 335, 344, 346, 637

cerevisiae, 342, 343, 344

paradoxus, 342, 343
Saccharomycetaceae, 244, 322, 341, 343,

344, 345, 346
Saccharomycetales, 192, 200, 202, 210,
211, 212, 272, 334, 350, 352, 585

key to families, 358

keys to genera, 358
Saccharomycodes, 344

ludwigii, 345
Saccharomycoideae, 345
Saccharum officinarum, 140
Saccoblastia,' 439, 442, 443
Saccobobis, 226
Sachs, 549, 633, 636, 638
Sagittaria, 411
Salix, 320, 405
Salmon, 310, 316, 317
Salvin, 110

Sammelzelle, 65, 66, 67
Sappin-Trouffy, 143, 380, 383, 400, 401,

405, 407
Saprolegnia, 100, 108, 110, 112, 114, 116,
130, 142, 153, 341, 634

784

INDEX

Saprolegnia — {Continued)

ferax, 98
monoica, 112

var. glomerata, 111
Saprolegniaceae, 71, 95, 97, 98, 99, 108,
109, 111, 113, 114, 115, 116, 119,
127, 142

key to genera, 121
Saprolegniales, 3, 35, 49, 94, 95, 104, 105,
107, 108, 126, 128, 130, 135, 142,
143, 150, 151, 630, 633, 634, 635

key to families and genera, 121
Sapromyces, 117
Saprophyte, 7, 8
Sarcodina, 630
Sartoris, 267, 409
Sartory, 326
Sass, 375

Satina and Blakeslee, 160
Savile, 390, 391
Sawyer, 177
Saxifragaceae, 405
Scheibe, 382
Scherffel, 45, 53, 99
Schikorra, 327
Schistodes, 314
Schizonella, 418

melanogramma, 418
Schizophijllum, 507

commune, 465
Schizosaccharomyces, 344, 345

odosporus, 335, 344

pombe, 344

versatilis, 335, 344
Schizosaccharomycetaceae, 341, 344, 345
Schizostoma montellicum, 279
Schizothecium, 200, 275, 375, 391

anserinum, 264, 266, 268, 269, 270
Schneepia, 294
Schneider, 218, 221, 345
Schostakowitsch, 153
Schroeter, 130, 135, 227, 494, 546, 573
Schultz, 141
Schwartz and Cook, 45
von Schweinitz, 12
Schweizer, 224
Schwendener, 216
Scirrhiineae, 284
Sclerenchymatous cells, 196
Sclerocystis, 171
Scleroderma, 531, 547, 549, 550, 555

aurantiacum, 547
Scleroderniataceae, 534, 535, 547, 550
Sclerodermatales, 535, 546, 553, 554, 558
Sclerospora, 140

graminicola, 140

macrosporn, 140
Sclerotia, 4, 5, 8, 195, 196, 203, 224

Sclerotia — (Continued)

subterranean, 487
Sclerotinia, 232, 602
fr-ucticola, 232
sclerotiorum, 194, 232
Sclerotiniaceae, 232, 347, 589
Sclerotiopsis concava, 601
Sclerotium, 602

rolfsii, 4
Scorias spongiosa, 318
Scott, 66

Scutellinia stercorea, 224
Scutiger,. 489

Seaver, 202, 225, 227, 249
Sebacina, 454
Secale cereale, 421

Secotiaceae, 534, 539, 540, 541, 550
Secotium, 532, 534, 540, 542
agaricoides, 540
coprinoides, 541
erythrocephalum, 540, 541
novae-zelandiae, 540
olbium, 541
Self-compatibility, 330
Self-incompatibility, 269, 375
Self-sterility, 226, 227
Senft, 193

Septobasidiaceae, 440, 445, 446
Septobasidiales, 440

Septobasidium, 436, 438, 439, 440, 443,
448, 449
burtii, 446
Septochytrium, 63

variabile, 64
Septocylindriuni, 590, 591
Septogloeum, 582

mori, 581
Septolpidium, 50
Septonema spilomcinn, 240
Septoria, 265, 680
aesculi, 579
apii, 580

apii-graveolentis, 580
lycopersici, 580
piricola, 298
Septorisphaerella, 298
Septotinia, 232

podophyllina, 201
Septum formation, 4
Serpula, 485
Sesamum indicum, 7

orientale, 197
Setae, 487
Seiaria, 140
Seuratia coffeicola, 321
Sexual phases, 241, 242, 370, 375, 388,
412, 453, 473, 501, 531
bipolar, 378

INDEX

785

Sexual phases — (Continued)

quadripolar, 378

reproduction, fundamental phenomena,
371

strains, 226, 267, 278

tendencies, 212
Sexuality, bipolar, 377, 487

quadripolar, 377, 487, 488
Seyfert, 409

Shanor, 90, 98, 132, 133, 168
Shear, 269, 391, 602
Shear and Dodge, 444, 445, 601
Shikorra, 323, 324
Sibelia, 382
Sideris, 130

Sigmoideomyces, 167, 586
Silver spoon test, 505
Simhlum, 544

Singer, 471, 473, 475, 481, 483, 485, 486,
488, 489, 490, 496, 507, 508, 509,
511, 518, 532, 652
Sinoto and Yuasa, 23, 24
Siphonales, 107, 634
Siphonaria, 55, 57

petersenii, 58
Sirobasidiaceae, 453, 454
Sirohasidium, 437, 451, 453

albidum, 454
Sirolpidiaceae, 96, 100

key to genera, 120
Sirolpidium, 100
Sistotrema, 482
Sjowall, 162

Skupienski, 24, 27, 29, 631
Sleumer, 409, 413
Slime molds, 1, 23
Smith, A. H., 520
Smith, A. H., and Miss Morse, 512
Smith, Miss A. L., 219, 223
Smut galls, 418
Smuts, 8, 15, 192, 194, 197, 409

hybridization, 416

manner of infection, 410

mutant forms, 417

origin, 644
Snell, 518
Solanuni tuberosum, 34, 50, 133, 475, 589,

600
Solenia, 476, 484

Candida, 476
Solidago, 405
Sommerstorffia, 113
Sordaria, 275

fimicola, 266
Sordariaceae, 274
Soredium, 219
Sorghum vulgare, 418
Sorodisciis, 34

Sorokin, 67, 69
Sorolpidium, 35
Sorosphaera, 34

radicalis, 34

veronicae, 33, 34
Sorosporium, 419

consanguineum, 413, 414

reilianum, 415, 416

saponariae, 418

syntherismae, 417
Sorus, 52, 135

uredial, 393
Southern beech, 226
Southworth, 583
Sparassis, 476, 479

ramosa, 476
Sparrow, 31, 35, 46, 49, 55, 57, 59, 60,
64, 65, 66, 70, 79, 89, 94, 95, 96,
100, 102, 104, 108, 113, 128, 630
Spathularia, 234
Speare, 177
Species, bipolar, 531

quadripolar, 531
Spegazzini, 13
Spegazzinia, 601
Sperm, 6, 386
Spermatia, 220, 222, 368
Spermatization, 391
Spermogonial nectar, 390
Spermogonium, 205, 206, 207, 216, 220,
222, 223, 368, 386

subcortical, 386, 387

subcuticular, 386, 387, 407

subepidermal, 386, 387, 407
Spermophthora, 352

gossypii, 345, 350, 351
Spermophthoraceae, 337, 345, 350, 351
Sphacelia, 289

Sphaceloma, 333, 334, 573, 583
Sphacelotheca, 418, 423

columellifera, 413

ischaemi, 415

schweinfurthiana, 413, 414

sorghi, 418
Sphaeria, 273

Sphaeriaceae, 273, 275, 276, 278, 284
Sphaeriales, 12, 210, 215, 271, 272, 273,
285, 286, 290, 291, 292, 295, 296,
299, 309, 319, 320, 322, 323, 458,
573, 578, 579, 580, 582, 591

key to families, 300
Sphaerioidaceae, 576
Sphaerita, 50, 100

dangeardii, 50, 99
Sphaerobolaceae, 535, 549, 554
Sphaerobolales, 535, 549
Sphaerobolus, 549, 550

grandis, 531

786

INDEX

Sphaerobolus — (Continued)

stellatus, 550, 555
Sphaerocladia, 82
Sphaerocysts, 501
Sphaeropsidaceae, 576, 578, 579, 601

key to genera, 604
Sphaeropsidales, 16, 240, 575, 576, 578
Sphaeropsis, 578, 579

malorum, 298, 578
Sphaerosporanginm, 130
Sphaerostilbe, 287

gracilipes, 289
Sphaerotheca, 314

castagnei, 312, 313

fuliginea, 314

humuli, 315, 316

mors-uvae, 310, 316

pannosa, 317
var. rosae, 311

phytoptophila, 310
Sphagnum, 419
Spicaria, 587
Spinacia oleracea, 141
Spinalia, 168
Spirogyra, 53, 99
Sponge mushrooms, 228
Spongospora, 34

subterranea, 33, 34
Sporangia, 5, 32

sympodially produced, 110
Sporangiole, 153, 156, 166, 170, 172

monosporous, 155
Spore, amyloid, 470, 484

aquatic, 575

forms, classification, 577

fruit, 7, 192, 197, 638
gymnocarpic, 496
ontogeny, 496
pseudoangiocarpic, 496
resupinate, 465

non-amyloid, 470

repeating, 381

secondary, 436

types, 508

wall, amyloid, 508
non-amyloid, 508
Sporendonenia, 576
Sporidiobolus, 349, 350
Sporidium, 366, 379

discharge, 369, 384

infection by, 385

quaternary, 385

secondary, 384

tertiary, 385
Sporobolomyces, 348, 349, 368, 379

roseus, 348
Sporobolomycetaceae, 348
Sporocarp, 170, 171

Sporocarp — (Continued)

consistency, 467
Sporocladia, 156, 161, 169
Sporodesmium, 240, 597
Sporodinia, 165

grandis, 157, 160, 161, 164
Sporodochium, 334, 599
Sporomyxa, 35
Sporonema, 280
Sporoschisma, 576
Spot anthracnose, 334
Stachylina, 178
Stahl, 220

Stakman, 14, 382, 383, 389, 409, 415
Staphylococcus, 602
Starbkck, 196
Steccherinuni, 482

Stelling-Dekker, Miss, 336, 345, 358
Stemonitis, 26, 28

fusca, 24
Stempell, 410, 421
Stemphylium, 596

botryosum, 597

sarcinaeforme, 597
StereocauloJi coralloides, 218
Stereum, 11, 467, 475
Sterigma, 168, 576, 586, 587

primary, 586

secondary, 586
Sterigmatocystis, 325, 586
Stevens, 136, 137, 317, 319, 354
Stichobasidial development, 437, 464,

477, 480
Stictis, 230
Stigeoclonium, 57
Stigmatea, 293

robertiani, 293
Stigmateaceae, 293
Siigmatomyces baeri, 214
Stigmatomycosis, 350
Stilbaceae, 584
Stilbella, 286, 287, 584
Stilbellaceae, 584, 598, 599

key to genera, 621
Stilbum, 444, 584
Stomatogene, 317
Stoneman, 279
Storage organs, 195
Streptomyces, 585,'603

scabies, 585
Streptomycetaceae, 585
Streptomycin, 602, 603
Strobilomyces floccopus, 497

strobilaceus, 497
Strobilomycetaceae, 496
Strobilophyta, 382

Stromatinia gladioli, 211, 224, 226, 233,
391

INDEX

787

Strong, 228, 229, 235, 290
Structure, types of, 532
Stylina, 423
Stylopage, 179
Stylospore, 170
Stypella, 454
Stysanus, 598

stemonites, 599
Subhymenium, 499
Subiculum, 476
Suborder Endosporeae, 25
Suchfaden, 417
Suillellus luridus, 497
Suillus, 11

luteus, 497
Suss, 504

Swarm cells, anteriorly or laterally bi-
flagellate, 94

posteriorly uniflagellate, 94
Swarm spore, 23
Swingle, 151
Swoboda, 552

Sydow, 273, 284, 290, 292, 293, 294, 296,
299, 303, 308, 316, 318, 319, 320,
326, 346, 354, 357, 399, 400
Sympodial development, 130
Synascus, 145, 353
Syncephalastrum, 168
Syncephalideae, 180
Syncephalis, 168

cornu, 159
Synchytriaceae, 50, 51, 52, 55

composition of cell wall, 54
Synchytrium, 45, 50, 53

decipiens, 52

endohioticwn, 50, 51, 53

fulgens, 52, 53

mercurialis, 45

papillatum, 50

vaccina, 50, 53
Synnema, 278, 598
Syringa vulgaris, 317
Syringospora, 347
Syzigites, 165
Syzygospora, 443

alba, 443
Szymanek, 132

Tabor and Bunting, 135
Taeniellaceae, 182
Taeniellopsis orchestiae, 182
Taphria, 243
Taphridium, 143, 144
Taphrina, 211, 212, 243, 637

cerasi, 243

communis, 243

Taphrina — (Continued)

deformans, 241, 242, 243, 244

epiphylla, 241

klebahni, 241

potentillae, 243

pruni, 243

robinsoniana, 243
Taphrinaceae, 241, 243, 639
Taphrinales, 215, 241, 368, 458, 510

key to genera, 256
Taraxacum, 17
Targioni-Tozzetti, 11
von Tavel, 556
Tear stain, 280
Tehon, 581, 607, 608
Tehon and Daniels, 595
Teleutospore, 449
Teliospore, 5, 379, 380, 399, 438

compound, 394
Teliosporeae, 349, 350, 366, 378, 379, 383,
436

and Heterobasidiae, summary, 457

key to orders and families, 424

relationships, 423

sexual reproduction, 382
TeUum, 399
Terfezia, 327

Terfeziaceae, 239, 325, 327, 331
Ternetz, 195
Terrostella, 554
Tetrachaetum, 575

elegans, 574
Tetrachytrium, 67, 69

triceps, 69
Tetracladium, bib

marchalianum, 574
Tetramyxa, 34
Thallus, crustose, 217

foliose, 217

fruticose, 217

pendent, 217
Thamnidiaceae, 157, 166

key to genera, 184
Thamnidium, 153, 166, 167

elegans, 157
Thamnocephalis, 167
Thaxter, 83, 86, 87, 88, 118, 159, 169,

174, 176, 214, 215, 244, 588
Theissen, 205, 271, 273, 284, 290, 292,
293, 294, 296, 299, 303, 308, 316,
318, 319, 320, 354, 357
Thelebolus stercoreus, 200, 201
Thelephora, 476, 481

terrestris, 476
Thelephoraceae, 457, 458, 469, 470, 473,
474, 476, 477, 480, 481, 482, 484,
486, 487, 489, 490, 494, 510, 536,
602

788

INDEX

Thelephoraceae — {Continued)

key to genera, 511
Thielaviopsis, 277, 278, 576

paradoxa, 576
Thirumalachar, 311, 396
Thorn, 325

Thorn and Raper, 587
Thomas, 134, 193, 634
Thraustochytriaceae, 94, 96, 104

key to family, 121
Thraustochytrium, 104
Thraustotheca, 109, 110, 115

clavata, 110

primoachlya, 109
Thrush, 337, 347
Tibicina septendecim, 177
van Tieghem, 159, 345
Tieghernella coerulea, 160

glauca, 160
Tilletia, 414, 419

caries, 410, 419

foeiida, 410, 419, 421

levis, 410, 419

sphagni, 419

tritici, 410, 419, 420
Tilletiaceae, 368, 381, 409, 410, 419, 420

key to genera, 427
Tilletiopsis, 348, 349
Tinsel type flagellum, 629
Tischler, 45, 468
Tisdale, 66
Tithymalus, 395
Tjibodasia, 443
Toadstool, 505
Tobler, 217, 218
Togashi and Oda, 508
Tolyposporium, 419

junci, 418
Tomentella, 467, 475, 482

flava, 474
Torula, 342, 346, 591
Torulopsidaceae, 346, 348

key to genera, 359
Torulopsis, 342, 346, 347

pulcherrima, 244
Torulopsoideae, 346
Tournefort, 10
Toxalbumins, 505
Trabutiineae, 284
Trachysphaera, 135
Trachyspora, 407
Tragopogon porrifolius, 137
Trama, 498, 531
bilateral, 499
inverse, 499
irregular, 499
mixed, 499
regular, 499

Trametes, 11, 491, 494

americana, 378, 487

serialis, 487
Tranzschelia pruni-spinosae, 408
Trematophlyctis, 35
Tremella, 11, 384, 438, 454, 455, 457

mesenterica, 452

reticulata, 452, 455
Tremellaceae, 452, 453, 454, 456

key to genera, 460
Tremellales, 197, 379, 437, 451, 452, 454,

456, 458
Tremellodendron, 455
Tremellodon, 454, 455
Trentepohlia, 217, 285
Trichaster, 554
Trichia, 28

varia, 24
Trichiaceae, 557
Trichocoma, 330
Trichocomaceae, 325, 330, 334
Trichoglossum, 234
Trichogyne, 210, 211, 213, 214, 215, 220,

222, 389
Tricholoma, 504

personatum, 502
Trichomonascus, 338
Trichopeltaceae, 295
Trichophore, 213
Trichoscyphella, 232

wilkommii, 232
Trichoihecium, 589
Trichothyriaceae, 308, 321

key to genera, 355
Trichothyrium, 322
Tricladiuni, 575
Trifolium, 595, 597

pratense, 224
Triticum, 55, 316, 395, 418

aestivum, 102, 382, 421

durum, 382
Trogia, 484
Trophic hyphae, 42
Trophogone, 208, 268, 323
Trow, 128
Truffle, 10, 239
Trypethelium, 285
Tsuga, 494

canadensis, 405
Tuber, 10, 11, 237, 239

aestivum, 237, 238, 239

brumale, 237

candidum, 201

lapideum, 237, 273

magnatum, 238

melanosporum, 237, 239

mesentericum, 237

panniferum, 237

INDEX

789

Tuber^(Continued)

riifum, 238
Tubera, 10

Tuberales, 203, 215, 236, 273, 291, 325,
327, 639

key to genera, 255
Tubercularia, 287, 600

vulgaris, 600
Tuberculariaceae, 584, 598, 599

key to genera, 622
Tuburcinia, 410, 421

trientalis, 420
Tucker, 132

Tulasne, 12, 13, 238, 289, 315, 538, 547
Tulasnella, 456, 457

violea, 456
Tulasnellaceae, 455, 456, 473

key to genera, 461
Tulasnellales, 379, 437, 455, 466
Tulipa, 397
Tulostoma, 436, 531, 553, 555, 557

campestre, 556

simulans, 555
Tulostomataceae, 535, 550, 555, 556, 557
Turbinate cell, 66

organs, 61
Tylopilus felleus, 497, 498
Type family, 16

genus, 16

species, 16

specimen, 16
Tijpha, 167
Typhella, 178
Tijphula, 478, 602
Typhulochaeta, 314

Ulmus americana, 598
Umbelliferae, 143
Uncinula, 314

necator, 310, 316

salicis, 315
Uncinulopsis, 314
Unipileate type of sporocarp, 533
Universal veil, 508

Uredinales, 5, 8, 14, 16, 368, 379, 380,
381, 382, 385, 392, 398, 400, 401,
403, 405, 406, 423, 436, 438, 449,
453, 458, 572, 639
Uredineae, 380
Uredinella, 438, 449
Uredinopsis, 393, 395, 402

struthiopteridis, 403
Urediospore, 381

primary, 399

secondary, 399
Uredium, 393, 399

Uredium — (Continued)
cupulate, 399
extrastomatal, 401
type, 401

with peridium, 401
Uredo, 408, 572
linearis, 18
mother cells, 449
sorghi, 17
Uredospore, 381, 449
Urobasidium, 481, 594
Urocystis, 421
anemones, 409, 421
gladioli, 409
occulta, 410, 421
tritici, 421
violae, 420, 421
Uromyces, 402, 407
aloes, 396
caladii, 383
caryophyllinus, 395
dianthi, 395
erythronii, 400
fabae, 406

phaseoli typica, 382, 389, 395, 408
phaseoli vignae, 389
Urophlyctis, 64, 66

alfalfae, 66, 67
Urtica, 141
Usnea barbata, 218

Ustilaginaceae, 6, 172, 379, 409, 412,
413, 415, 418, 423
key to genera, 426
teliospore longevity, 417
Ustilaginales, 8, 14, 16, 194, 337, 346, 379,
380, 381, 384, 409, 412, 413, 415,
418, 420, 422, 423, 424, 438, 453,
487, 572
Ustilago, 378, 409, 418, 419
avenae, 410, 414, 415, 416, 418
cardamines, 410
fischeri, 416
hordei, 415, 417, 418
ischaemi, 415
kuehneana, 418
levis, 409, 415, 416, 418
longissima, 378, 416, 417, 423
longissivia var. macrospora, 417
nuda, 411, 412, 418
perennans, 416
striiformis, 409, 416, 418
striiformis forma Hordei, 416
striiformis forma Poae-pratensis, 417
tritici, 411, 412, 418
violacea, 411, 414, 418
vuijckii, 413

zeae, 409, 410, 411, 412, 415, 416,
418

790

INDEX

Vaccinium, 403
Vaillant, 10
Valid publication, 16
Valkanov, 30
Valsa, 282, 578
Valsaceae, 280
Vulsella, 282

Vandendries, 14, 368, 370, 371, 377, 378,
488

and Brodie, 160

and Martens, 368, 468, 501
Vanterpool and Ledingham, 102, 103
Vararia, 475
Varicosporium, 575

elodeae, 574
Varitchak, 267, 341, 352, 353
Vaucheria, 71, 97, 107, 633, 634
Veins, 239, 330, 531
Venturia inaequalis, 8, 265, 266, 270, 298,

589
Veronica, 34
Verpa, 228, ^32

Verrucaria, 285 '

Verticillium, 589

albo-atrum, 588, 589
Vesicle, 95, 100, 102, 103, 117, 128, 129,
130, 134, 135

subsporangial, 153, 165, 166

substomatal, 393
Vetch, 47
Vihrissea, 230
Vicia, 47

Vigna sinensis, 600
Vincens, 274, 277
Viola, 421
Vitis, 141, 275, 310, 316, 334

vinifera, 297, 316
Volkartia, 143
Volutella, 600

dianthi, 600
Volva, 509
Volvaria, 501
Voss, 383
Vuillemin, 168, 575 •

Waterhouse, 389, 394

Webb, 34

Weber and Wolf, 158

Wehmeyer, 274, 277, 280, 281, 283

Weir, 384

Welsford, 286

Wernham, 409

Weston, 140

von Wettstein, 3, 26, 50, 79, 134, 151, 193

Whetzel, 201, 232

Whiffen, Miss, 46, 54, 59

Whiplash type flagellum, 629

White, 196, 493

White rust, 8, 137

Wieben, Miss, 211, 241, 242

Wilcox, 269

Willia, 336, 344

Williamson, 207, 210, 211

Wilson, M. and Cadman, 24

Wilson, O. T., 66

Wiltshire, 596, 597

Wingard, 345

Winge, 342

Winter, 587

van Wisselingh, 26, 31

Witches' broom, 241, 386

Withertip, 280

Wolf, F. A., 231, 265, 481, 495

Wolf, F. T., 85

Woronin, 14, 420

Woronina, 31, 35, 97

polycystis, 97
Woroninaceae, 96, 97

key to genera, 119

Xanthophyceae, 629, 632
Xenodochus, 407
Xenolachne, 454
Xylaria, 268, 284

hypoxylon, 16, 284

polyrnorpha, 284

subterranea, 283
Xylariaceae, 277, 280, 282, 283, 284, 287

Wager, 57

Waite, 243

Wakefield, 327

Waksman and Henrici, 585, 603

Walker, Miss, 170, 171, 340, 341, 507,

537, 550
Wang and Martens, 383, 391
Wartenweiler, 141
Water molds, 105

Yarwood, 316
Yeast-like growth, 151
Yeasts, 6, 200, 334, 335

asporogenous, 336, 342, 346, 585

bottom, 341

fission, 335

key to related families, 360

red, 348

sporogenous, key to genera, 358

top, 341

INDEX

791

Yeasts — (Continued)

true, 341
Yen, 413, 414
Young, E. L., 30, 31
Young, Miss, 323, 324, 327
Yu, 421
Yuasa, 24

Zaghouania, 384

Zahlbruckner, 223

Zea mays, 15, 66, 140, 395, 418, 594

Zeller, 225, 535, 540, 550, 553

Zeller and Dodge, 539, 542

Zeller and Walker, 537, 538

Zander, 336

Ziegenspeck, 202

Ziegler, 109

Zimmermann, 594

Zodiomyces vorticellarius, 213, 214

Zoopagaceae, 178, 179, 180, 585

Zoopagales, 150, 177, 634, 635

key to families and genera, 186
Zoopage, 179

phanera, 180
Zoosporangium, 5

filamentous, 129

formation by proliferation, 105

proliferated, 111, 112, 128, 142

resting, 119

smooth-walled, 119

Zoosporangium — (Continued)

spiny, 118, 119

sympodial formation, 105

toruloid, 128, 130
Zoospore, 5, 193

biflagellate, 78

dimorphic, 116

encysted, 113

primary, 100, 102, 104, 105, 109, 110,
112, 115, 116

secondary, 95, 100, 102, 105, 110, 112,
115, 116, 117, 126, 132
Zopf, 49, 99, 101, 155, 218
Zostera marina, 30
Zycha, 163
Zygnemataceae, 49
Zygochytrium, 67, 69

aurantiacum, 69
Zygomyceteae, 143, 380
Zygorhynchus, 163, 165

ynacrosporus, 157
Zygosporangium, 179
Zygospore, 6, 150, 152, 156, 163, 176, 179

anisogamous formation, 157
Zygosporium, 481, 594

echinosporum, 594

oschioides, 594
Zygote, 6, 7, 24

biflagellate, 82, 83, 85
Zythia, 580
Zythiaceae, 580

key to genera, 607

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