Soil Seed Fungi
Soil Seed Fungi
Soil and
Seed Fungi
Morphologies of Cultured Fungi
and Key to Species
Second Edition
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Pictorial Atlas of
Soil and
Seed Fungi
Morphologies of Cultured Fungi
and Key to Species
Second Edition
Tsuneo Watanabe
CRC PR E S S
Boca Raton London New York Washington, D.C.
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This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with
permission, and sources are indicated. A wide variety of references are listed. Reasonable efforts have been made to publish
reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials
or for the consequences of their use.
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identification and explanation, without intent to infringe.
Since the publication of the English edition of this book in 1994, the classification system of
organisms has changed dramatically, following the eight-kingdom concept based on the knowledge
obtained with molecular techniques. All soil fungi now belong to the kingdom of Fungi, Chromista,
or Protozoa; therefore, soil fungi described in this book are based on the traditional broad concept
of fungi belonging to one of these kingdoms. Although some fungal species have been identified
with the molecular techniques without observing the morphological characteristics, the fungi have
been originally named on morphology, and new fungi will be described and accepted scientifically
on the basis of morphological descriptions following the traditional classification system. Therefore,
compiled knowledge on these fungi is still essential and needed for any workers on this line.
In this revised second edition, the original text is partially revised by rearranging, correcting,
rewording, redrawing electronically and adding the recent knowledge, together with descriptions
of 45 additional fungal species, their pictures, illustrations, and references.
I would like to thank John Sulzycki, Senior Editor; Samar Haddad, Project Editor; Erika Dery,
Project Coordinator; and Barbara E. Norwitz, Publisher, at CRC Press, for their help, advice, and
considerations in publishing this edition.
Tsuneo Watanabe
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Preface
Biological elements related to higher plants and animals have been studied in relation to environ-
mental problems, but the significance of microorganisms, another biological element, has been
neglected. Microbial activities include nitrogen circulation and fixation, and disintegration of
organic matter. Most living entities may not exist on Earth without the activities of microorganisms.
Some edible mushrooms are cooked and eaten directly. Bread, cheese, syouyu (soy sauce),
miso, and various other foods, as well as alcoholic beverages, chemicals, medicines, including
acids and penicillin, and other agricultural chemicals such as gibberellin, are products of microbial
activities of yeasts, Aspergillus, Penicillium, and other fungi. Some fungi, however, attack directly
plants, animals, and human beings, causing diseases. Also, some fungi are often poisonous and
carcinogenic, excrete toxins contaminating foods. Dusts and fungi inhaled occasionally inside
houses and buildings, or in the fields, cause asthma and allergenic troubles. Houses, foods, industrial
products, books, and arts and craft materials are often damaged due to fungal activities. Some fungi
are useful, however, influencing our lives directly or indirectly, but most fungi live under soil
conditions separate from our lives and we do not understand their usability.
In this book, fungi detected or isolated from soil or plant seeds are provisionally termed as
“soil and seed fungi.” All these fungi influence one another, with antagonism or mutual symbiotic
or cooperative relationships. Some of them also influence plant growth, and may degrade ligno-
cellulose or deleterious atmospheric materials, including useless plastics and dioxin and its related
compounds. By studying the individual fungi mycologically and physiologically, and increasing
our knowledge of them, we may understand more about plant pathogenic and usable fungi, con-
comitantly from ecological views with plant protection and further improvement of human life.
Many books on mushrooms and fungi and the diseases they cause have been published. Drawings
and pictures of these fungi and diseases are helpful for identification and disease diagnosis; however,
only a few books are available on soil and seed fungi and their descriptions are very limited. In
the plant pathological fields, mycological studies are prerequisite to determining the causal agents
of the diseases, including isolation, preservation, classification, and identification.
The fungi in this text were detected, isolated, and stocked during such etiological and ecological
studies. Most of them are not pathogenic to plants, and have been neglected for now. However,
biocontrol practices have been emphasized recently to control plant diseases, and various
saprophytic fungi have been studied to find effective biocontrol agents among them. In addition,
physiologically active substances from fungal products may be screened and found by studying
soil fungi together with gene bank practices.
In this text, 308 species of fungi that have been studied pathologically and mycologically during
the course of soilborne plant disease studies, were selected among 10,000 living stock cultures of
the author’s own, and were described with pictures and illustrations.
Although purposes, materials, and methods of research are different, knowledge of soil fungi
itself may be helpful for those interested in mycological work. Furthermore, only a few references
and monographs are available on soil and seed fungi, and this book may therefore foster interest
in these fungi.
For the publication of this book, the author would like to thank Susumu Yoshida, President,
and Muneo Okazaki, Editor-in-Chief of Soft Science Publications, Tokyo, Japan, for their help
and advice.
Tsuneo Watanabe
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Soil fungal floras and fungi associated with plant seeds have been studied all over the world, and
some of them have been published. However, publications treating morphologies of these fungi
identified are rather limited, except those done by Gilman (1945), Barron (1968), Domsch and
Gams (1972), and Domsch et al. (1980a,b).
In this book, morphologies of 308 species of fungi, mainly collected in Japan, are shown based
on the author’s own photomicrographs and illustrations.
This book was originally written in Japanese (original title: Photomicrographs and Illustrations
of Soil Fungi), and published by Soft Science Publications, Tokyo, Japan, in May 1993. In the
English edition, the original Japanese edition was partially revised, with four newly described fungi,
together with 30 additional references.
The English edition was issued by invitation of Jon R. Lewis, Editor-in-Chief, Lewis Publishers,
Inc., who asked me to write a book on “ecology and fungi” or a related topic in May 1991,
when I was in the process of writing this book in Japanese, and by the acceptance of its translation
into English.
I hope the English edition may contribute more easily and widely to the understanding of the
ecology and distribution of fungi.
The author would like to thank Jon R. Lewis, Editor-in-Chief; Ed Norman and Jennifer Pate,
Project Editors, Lewis Publishers, Inc.; and the persons concerned with the publication of the
English edition for their help and advice; and S. Yoshida, President, Soft Science Publications,
Tokyo, Japan, who kindly agreed to publish the English edition of this book.
Tsuneo Watanabe
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More than 350 fungal species are described, including 46 Mastigomycetous species belonging
to seven genera, 33 Zygomycetous species (12 genera), 36 Ascomycetous species (16 genera),
9 Basidiomycetous species (more than 2 genera), and 240 mitosporic fungal (Deuteromycetous)
species (116 genera).
Latin binominals are adopted, mainly following publications by Farr et al. (1989), Hawksworth
et al. (1995), and Jong and Gantt (1987). Also, major synonyms that appeared in recent literature
are frequently consulted.
All of the fungi in this text are described alphabetically in order of Mastigomycotina, Zygo-
mycotina, Ascomycotina, Basidiomycotina, and Deuteromycotina. For each fungus, the literature,
morphology, and dimensions of each organ and material are described in order, occasionally with
the remarks. Colony characteristics are not generally included because of too much variation in
media used, and time of observation. Keys are prepared for all of the fungi to the genus level, and
to the species level for the genus with more than three species.
All illustrations and pictures are based on the author’s own work using the materials described.
The fungi examined are isolated from soil, plant roots, and seeds. Most of the samples were collected
in Japan, but some samples are from the Dominican Republic, Paraguay, Switzerland, and Taiwan,
the Republic of China (ROC). Details of isolation methods are described elsewhere in the text.
Scientific terms are mostly adopted following Hawksworth et al. (1995).
Living cultures of some fungi are deposited at the Gene bank, Ministry of Agriculture, Forestry
and Fishery (MAFF), National Institute of Agrobiological Sciences in Tsukuba, at Fermentation
Institute, Osaka (IFO), at American Type Culture Collection (ATCC), and at Centraalbureau Voor
Schimmelcultures (CBS), and most of them are listed in the appendix of this book.
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Acknowledgments
We wish to acknowledge the following journals for granting us copyright permissions: Ann.
Phytopath. Soc. Jpn. (the Phytopathological Society of Japan), Mycologia (the Mycological Society
of America, the New York Botanical Garden) Mycologia Helvetica (the Swiss Mycological Society),
Phytopathology (the American Phytopathological Society), Trans. Br. Mycol. Soc. (the British
Mycological Society), and Trans. Mycol. Soc. Jpn. and Mycoscience (the Mycological Society
of Japan).
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The Author
Tsuneo Watanabe earned his Ph.D. degree in plant pathology from the University of California,
Berkeley in 1967. He has been active as a plant pathologist and mycologist in the field of soilborne
diseases, their pathogens, and soil fungi for more than 30 years at the National Institute of
Agroenvironmental Sciences, Forestry and Forest Products Research Institute, and National Institute
of Advanced Industrial Science and Technology of Japan.
He is the author of more than 150 scientific papers on the subject, and the book titled Dictionary
of Soilborne Plant Diseases (Asakura Publishing Company, Ltd., Tokyo, 1998).
Dr. Watanabe is currently studying lignocellulose and dioxin-decomposing fungi as a part of
the national Bioconsortia Project of Japan.
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Table of Contents
3 Identification of Fungi............................................................................................................17
3.1 Basal Knowledge for Identification ...........................................................................17
3.2 Necessity of Experimentation ....................................................................................18
3.3 Selection of Appropriate Binomials...........................................................................18
3.4 Morphologies to be Observed for Identification .......................................................18
3.5 References for Fungal Taxonomy and Identification ................................................20
References ......................................................................................................................................457
Index ..............................................................................................................................................479
Afterword .......................................................................................................................................486
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1
Study on Soil and Seed Fungi
1
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In Japan, Takahashi (1919) isolated some fungi from field soils at Komaba in Tokyo, identified
25 species belonging to 13 genera, and reported in the first volume of Ann. Phytopathol. Soc. Jpn.
It must be the pioneer soil fungal study in Japan, although some other early works on this line may
have been conducted (note the early issues of J. Agrochem., Jpn.).
Waksman is famous for his discovery of streptomycin, and published a series of papers on soil
fungi since 1916 and advocated a theory that common fungi live in any soil. To demonstrate this,
he isolated more than 200 fungal species belonging to 42 genera from 25 locations in the U.S.
Among these fungi, four genera, viz. Aspergillus, Mucor, Penicillium, and Trichoderma, live com-
monly in soil of any location. Especially, he noted that Mucor and Penicillium commonly occur in
soil of temperate or cool-climate areas, Aspergillus in tropical soil, and Trichoderma occurs fre-
quently in wet or acidic soils. In addition to these four genera, the following eight genera were
pointed out to be dominant: Acrostalagmus (syn. Verticillium), Alternaria, Cephalosporium (syn.
Acremonium), Cladosporium, Fusarium, Rhizopus, Verticillium, and Zygorhynchus. He further
found that soil fungi tend to colonize on moribund plant residues, many kinds and numerous
populations of bacteria and Actinomycetes live in soil, and the more fertile the soils, the more
numerous and massive are the fungi.
Although there are differences in fungal species among soil isolates by different isolation
methods, 17 common soil fungi were nominated including 12, listed by Waksman, and the other
five, namely Absidia, Botrytis, Chaetomium, Cylindrocarpon, and Stemphylium, by Burges (1965).
Some fungi, such as Pythium, Mortierella, Rhizoctonia, and Basidiomycetous fungi, had not
been listed or were very rarely listed as members of soil fungi before 1949 (Chesters, 1949), but
by the hyphal isolation method devised by Warcup (1959), these fungi became common as soil
fungus members. Especially, about 40 species of Basidiomycetous fungi have been isolated from
soil and were identified in his work.
According to Burges (1965), over 600 fungus species, including 200 Phycomycetous species
(Mastigomycotina and Zygomycotina), 32 Ascomycetous species, and 385 Deuteromycetous (Mito-
sporic fungal) species are recorded as soil fungi in the first edition of “A Manual of Soil Fungi”
written by Gilman in 1945. Since then, soil fungus study has been serious and active, but nearly
1800 to 2000 species may be considered reasonable as the total number of soil fungi. This is
calculated on the basis of Gilman’s figure (600) in 1945 plus 1200, the figure increased since 1945.
Its nearly 800 species of fungi are now being catalogued in the Index of Fungi every year (Hawk-
sworth et al., 1995); if 20 of them (2.5%) are assumed to be soilborne, more than 1120 species
must be added to the number of soil fungi for the past 56 years.
These figures are less than 2.7% of 72,065 species of total fungi recorded (Hawksworth et al.,
1995). These small figures must be due to the lack of attention to individual organisms, because
soil microorganisms have been treated previously en masse in the categories of bacteria, actino-
mycetes, fungi, algae, and so on.
Researchers want to identify soil fungi and know their genus and species names, their numbers,
and the common and dominant species in soil surrounding us, although research purposes may
vary considerably. Concomitantly, it is helpful to compare fungi or fungus floras in different soils,
paying attention to the characteristics of the floras, and their similarity rates. To get this information,
various isolation methods and media have been used, including soil dilution and soil plate methods,
often with rose bengal streptomycin agar medium.
For particular genera or species, we like to compare their occurrence or detection frequency
(isolation frequency) and coefficient of similarity. Values of coefficient of similarity are obtained,
following the equation S = 2W/(a + b), where “w” is the number of common species in two fungal
populations and “a” and “b” are the numbers of species in both populations; namely, as the similarity
between two populations increases, the value of the coefficient of similarity approaches one.
By using these approaches, interrelationships among habitats of higher plants, soil fungus floras,
many different soils including forest, grassland, uncultivated and cultivated soils, and comparison
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of soil fungus floras in diseased vs. healthy soils, various soil atmospheres including soil pH, soil
type, organic contents, soil depth in different seasons, and factors influencing soil fungus floras
become subjects of study; through the knowledge of soil fungus floras we are able to understand
the various habitats of higher plants and their activities, soil fertility, and disease occurrence areas
and some environmental problems.
Sphaerostilbe repens, the root rot pathogen of tea and various plants in the tropics, is induced
to form hyphal bundles and synnema by the influence of Aspergillus spp., Penicillium spp., and
Verticillium lamelicola (Botton and El-Khouri, 1978). All these are examples of the morphogenesis
of certain fungi under the influence of activities of other organisms, and these activities are related
to alcohols including hexanal, ethyl and methyl alcohol; fatty acids including linoleic acid; oils
and fats; and unidentified substances excreted by microorganisms.
Generally speaking, with the increase of research on soil fungus floras, soil fungi may be
observed more individually, rather than en masse, and new fungi may be discovered and knowledge
on their classification will be more and more increased.
For Mastigomycetous fungi, Pythium spp. are generally isolated from old or declining roots,
occasionally with Aphanomyces, and Phytophthora, and aqueous fungi such as Dictyuchus,
Saprolegnia, and Pythiogeton. Among Zygomycetous fungi, the genus Mortierella is the most
common, followed by Mucor and Rhizopus. In addition, Absidia, Cunninghamella, Gongronella,
and Syncephalastrum may often be isolated. Saksenaea is not so often recorded as a member of
soil fungus floras, but it was isolated from sugarcane roots in Okinawa. Helicocephalum, one of
the Zygomycetous fungi which are not cultured singly and purely in vitro, was detected on agar
cultures together with various organisms, and lived for more than 6 months by the mixed culture.
These are rather rare fungi, because for over 30 years, the author has only once isolated them.
For the ascomyceteous fungi, Chaetomium is most commonly isolated, and the number of its
species isolated from soil is numerous. In addition, Thielavia and seven others are often isolated
from soil. Basidiomycetous fungi have not been commonly isolated from soil, but Coprinus and
Thanatephorus are examples of fruiting in vitro, but most of the Basidiomycetous isolates are
not successful for fruiting in vitro. They are just judged to be Basidiomycetous because of their
having clamp connections.
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Mitosporic (Deuteromycetous) fungi are most frequently isolated, and they are rich in species.
Alternaria and Penicillium are always listed as the members of fungus floras. Generally speaking,
a total of 9 to 74 mitosporic fungus genera have been isolated from the soil of each location
(see the Supplement at the end of this chapter).
Unsporulated, sterile, and unidentified fungi are different in their treatment in the literature,
and they are classified as independent items. It is not clear if these fungi do not sporulate because
of their innate nature, or if we could not induce their sporulation because of lack of technique. In
addition, there are a few unidentified fungi, although they sporulated.
In the fungal taxonomy, fungi are classified, identified, and described mainly based on morph-
ology observed in nature. For example, fungi belonging to the genus Pestalotia (syn., Pestalotiopsis)
form sporodochia with morphologically characteristic conidia, but they do not usually form acervuli
on agar cultures, the morphology of which has often been used as criteria for taxonomy. It appears
to be difficult to induce morphogenesis by inoculation in nature, because of the excessive labor
and time required. However, with the increase of knowledge on agar cultures of various fungi,
unknown fungi may be identified more readily.
Isolation Fungi
Sample (method and media) (no. of genera, species, and remark)a Ref.
North America
Canada
Manitoba, Dilution plate Total: 64 g, 178 spp. Bisby et al.
75 samples with two media (Zygomycota 7 g, 22 spp.; (1933, 1935)
Ascomycota 9 g, 13 spp.;
Basidiomycota 2 g, 2 spp.;
Deuteromycotina 46 g, 141 spp.)
Canada
Ottawa, Dilution plate Total: 17 g, 38 spp. Timonin (1940)
Rhizosphere soil of wheat, (Zygomycota 3 g, 3 spp.;
oats, alfalfa, and pea Deuteromycotina 14 g, 35 spp.)
Floras compared between rhizo- and
nonrhizosphere
Canada
10 forest soils Dilution with soil Total: 22 g, over 56 spp. Morral and
extract agar (Zygomycota 2 g, 12 spp.; Vanterpool (1968)
Ascomycota 1 g, 1 spp.;
Deuteromycotina 19 g, 43 spp.)
Dominant fungi: Mortierella, Pullularia,
Trichoderma, and Penicillium
Canada
Ontario, Soil washing Total: 41 g, 75 spp. Widden and
four conifer (Zygomycota 5 g, 10 spp.; Parkinson (1973)
forest soils Ascomycota 5 g, 5 spp.;
Deuteromycotina 31 g, 60 spp.)
Dominant fungi: Mortierella, Penicillium,
and Trichoderma
Canada
Tundra at the North pole Soil washing and plate Total: 26 g, 46 spp. Widden and
(Zygomycota 1 g, 1 sp.; Parkinson (1979)
Ascomycota 2 g, 2 spp.;
Deuteromycotina 23 g, 43 spp.)
Dominant fungi: Sterile fungi, Penicillium,
Chrysosporium, and Cylindrocarpon;
Trichoderma not detected. 27 species from
leaves also studied
U.S.
New Jersey, grassland Dilution and direct Total: over 29 g, 106 spp. Waksman (1916)
and others, 8 samples inoculation with four (Zygomycota 4 g, 19 spp.,
media Ascomycota 2 g, 5 spp.
Deuteromycotina 23 g, 82 spp.)
Dominant genera: Penicillium, Mucor,
Aspergillus, Trichoderma, and
Cladosporium
U.S.
New Jersey, grassland Dilution and direct Total: 42 g, 137 spp. Waksman (1917)
and others, 25 samples inoculation with four (Zygomycota 4 g, 18 spp.;
media Ascomycota 5 g, 7 spp.;
Deuteromycotina 33 g, 112 spp.)
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Isolation Fungi
Sample (method and media) (no. of genera, species, and remark)a Ref.
U.S.
Texas, forest soils, Dilution with Total: 13 g, 32 spp. Morrow (1940)
4 positions Waksman’s medium (Zygomycota 3 g, 5 spp.;
Ascomycota 1 g, 1 sp.;
Deuteromycotina 9 g, 26 spp.)
Dominant genera: Penicillium, Aspergillus
U.S.
Wisconsin southern Dilution with soil Total: 20 g, 50 spp. Dominant fungi: Tresner et al. (1954)
hardwood forest soils, extract agar (Zygomycota 4 g, 9 spp.;
13 locations Ascomycota 1 g, 1 sp.;
Deuteromycotina 15 g, 40 spp.)
Dominant genera: Absidia, Mucor,
Mortierella, and Zygorhynchus
Fungus floras reflected in higher plant
vegetation
U.S.
Georgia, forest and Dilution, direct Total: 63 g, 165 spp. Miller et al. (1957)
cultivated soils, inoculation and (Zygomycota 11 g, 22 spp.
45 samples others with rose Ascomycota 8 g, 10 spp.
bengal streptomycin Basidiomycota 2 g, 2 spp;
agar and various Deuteromycotina 42 g, 131 spp.)
media Dominant genera: Penicillium, Aspergillus,
Cunninghamella, Trichoderma, and
Rhizopus
Fungus floras compared between summer
and winter
The floras became poorer with soil depth
U.S.
Southern Wisconsin forest Dilution Total: >36 g, 199 spp. Christensen et al.
soils, five locations (Zygomycota >4 g, >9 spp.; (1962)
Ascomycota >2 g, >2 spp.;
Deuteromycotina >30 g, >83 spp.)
Dominant genera: Penicillium, Paecilomyces,
Gliocladium, and Humicola
Similarity rates among the respective
samples, 18.9–40.7
U.S.
Southern 10 states, Dilution and soil plate Total: 45g, 121 spp. Hodges (1962)
30 forest nurseries with rose bengal (Zygomycota 8 g, 16 spp.;
streptomycin agar Ascomycota 4 g, 11 spp.;
Deuteromycotina 33 g, 94 spp.)
Dominant fungi: Aspergillus, Penicillium,
Trichoderma, and Fusarium; Pythium and
Rhizoctonia not isolated
U.S.
Northern Wisconsin bogs Dilution Total: 57 g, 130 spp. Christensen and
and swamps, (Zygomycota 2 g, 11 spp.; Whitlingham
15 locations Ascomycota 8 g, 14 spp.; (1965)
Deuteromycotina 47 g, 105 spp.)
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Isolation Fungi
Sample (method and media) (no. of genera, species, and remark)a Ref.
U.S.
Alaska, uncultivated soils, Dilution with ether Total: 14 g, 23 spp. Yokoyama et al.
19 samples treatment (Zygomycota 3 g, 3 spp.; (1979)
Ascomycota 2 g, 3 spp.;
Deuteromycotina 9 g, 17 spp.)
Dominant fungi: Mortierella, Penicillium
U.S.
South Dakota, grassland Dilution Total: 13 g, 62 spp. Clarke and
10 samples (Zygomycota 1 g, 1 sp.; Christensen (1981)
Deuteromycotina 12 g, 40 spp.)
Dominant fungi: Penicillium, Acremonium,
Aspergillus, Chrysosporium, and Fusarium
Fungus floras compared domestically and
internationally
U.S. and Mexico
Arizona-Mexico Soil dilution and soil Total: 104 g, 230 spp. Ranzoni (1968)
Sonoran desert, plate with 5 media (Mastigomycota 1 g, 1 sp.;
24 locations, 30 samples Zygomycota 7 g, 15 spp.;
Ascomycota 21 g, 77 spp.;
Basidiomycota 1 g, 4 spp.;
Deuteromycotina 74 g, 133 spp.)
No specific fungus flora in desert. Curvularia,
colored fungi, and pycnidium-forming fungi
frequently isolated
Isolation Fungi
Sample (method and media) (no. of genera, species, and remark)a Ref.
Australia
Australia
Southern Australia, wheat Dilution, soil plate, Total: 57 g, 94 spp. Warcup (1957)
field soils and hyphal isolation (Mastigomycota 3 g, 4 spp.;
Zygomycota 9 g, 19 spp.;
Ascomycota 11 g, 13 spp.;
Basidiomycota 3 g, 3 spp.;
Deuteromycotina 31 g, 53 spp.)
Isolation methods compared
Basidiomycetous fungi isolated only by
hyphal isolation method
Australia
Wheat field soils Dilution, soil plate, Total: 54 g, 74 spp. Warcup (1976)
and plant residue (Mastigomycota 1 g, 8 spp.;
Zygomycota 10 g, 11 spp.;
Ascomycota 6 g, 8 spp.;
Basidiomycota 5 g, 5 spp.;
Deuteromycotina 32 g, 42 spp.)
Fumigation effect on fungus floras studied
Asia
India
Tea rhizosphere soils Dilution with rose Total: 26 g, 50 spp. Agnihothrudu
bengal streptomycin (Zygomycota 4 g, 5 spp.; (1960)
agar Ascomycota 4 g, 5 spp.;
Deuteromycotina 18 g, 40 spp.)
More numerous species isolated from roots
infected by Usulina zonata than from
healthy roots
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Isolation Fungi
Sample (method and media) (no. of genera, species, and remark)a Ref.
Iraq
Liwaas, Central Iraq, Not recorded Total: 41 g, 150 spp. Al-Doory et al.
5 locations (Mastigomycota 1 g, 8 spp.; (1959)
Zygomycota 3 g, 18 spp.;
Ascomycota 3 g, 5 spp.;
Deuteromycotina 34 g, 119 spp.)
Dominant fungi: Cladosporium, Aspergillus,
Fusarium, Alternaria, Penicillium,
Humicola, Mucor, and Pythium
Israel
Peanut fields, 12 locations Dilution with two Total: 42 g, 95 spp. Joffe and Borut
media (Zygomycota 8 g, 14 spp.; (1966)
Ascomycota 6 g, 30 spp.;
Deuteromycotina 28 g, 51 spp.)
Dominant fungi: Mucor, Rhizopus,
Aspergillus, Trichoderma,
Cephalosporium, and Fusarium
Israel
Mikve, cultivated soils Dilution and direct Total: 46 g, 147 spp. Joffe (1963)
inoculation with (Zygomycota 7 g, 13 spp.;
13 media Ascomycota 7 g, 60 spp.;
Deuteromycotina 32 g, 74 spp.)
Dominant fungi: Rhizopus, Aspergillus,
Penicillium, Alternaria, and Fusarium
Effect of fertilization and vegetation on
fungal floras studied
Japan
Osaka, paddy field soils, Dilution plate, Total: 21 g, 37 spp.: Ito et al. (1981)
4 locations isolated at 42°C (Zygomycota 2 g, 3 spp.;
Ascomycota 9 g, 16 spp.;
Deuteromycotina 10 g, 18 spp.)
Fungus floras in both meso- and
thermophilic fungi
Japan
Ogasawara (the Bonin Direct inoculation, Total: 47 g, 81 spp.: Watanabe et al.
Islands), forest and baiting with (Mastigomycota 1g, 10 spp. (2001a)
uncultivated soil, cucumber seeds and Zygomycota 3 g, 9 spp.
11 locations toothpicks Ascomycota 1 g, 2 spp.
Basidiomycota 1g, 1 sp.
Deuteromycotina 41g. 59 spp.)
the Bonin Islands: known as the Ocean
Islands
Malaysia
Forest and cultivated soils Dilution plate Total: 26 g, 54 spp. Varghese (1972)
in the west (Zygomycota 3 g, 4 spp.;
Ascomycota 4 g, 5 spp.;
Basidiomycota 1 g, 7 spp,;
Deuteromycotina 18 g, 38 spp.)
Pythium and Mortierella not isolated
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Isolation Fungi
Sample (method and media) (no. of genera, species, and remark)a Ref.
Europe
England
Brown forest and podozol Sieve soaking plate Total: 22 g, 50 spp. Thornton (1956)
soils and hyphal isolation (Zygomycota 3 g, 16 spp.;
Deuteromycotina 19 g, 33 spp.)
Dominant fungi: Trichoderma, Mucor,
Penicillium, and Mortierella
England
Suffolk grassland, five Soil plate Total: 48 g, 148 spp. Warcup (1951)
locations (Mastigomycota 1 g, 3 spp.;
Zygomycota 10 g, 30 spp.;
Ascomycota 9 g, 24 spp.;
Deuteromycotina 28 g, 91 spp.)
Dominant fungi: Penicillium, Mortierella,
Absidia, Cephalosporium, and Fusarium
England
Kidney bean root and its Soil plate Total: 17 g, 52 spp. Dix (1964)
rhizosphere soils (Mastigomycota 1 g, 1 sp,;
Zygomycota 3 g, 5 spp.;
Ascomycota 1 g, 1 sp.;
Deuteromycotina 12 g, 19 spp.)
Sweden
Forest soils in the south Soil washing Total: 21 g, 90 spp. Söderström (1975)
(Zygomycota 3 g, 18 spp.;
Deuteromycotina 18 g, 36 spp.)
Mortierella, Penicillium, and Trichoderma
occupied more than 71% of the total
isolates
Africa
Nyasaland
Coffee field Soil plate Total: 39 g, 81 spp. Siddiqi (1964)
(Zygomycota 7 g, 8 spp.:
Ascomycota 3 g, 4 spp.;
Basidiomycota 3 g, 3 spp.;
Deuteromycotina 26 g, 34 spp.)
Dominant fungi: Aspergillus, Trichoderma,
Cephalosporium, and Fusarium
a Aspergillus, Penicillium, and Rhizoctonia are variously treated for their ana- or teleomorph in the original literature. The
total number of genera and species of the respective classes in each work do not always coincide because figures of unknown
and sterile fungi may be included in one work, but not in another.
1118-ch02-Frame Page 13 Sunday, February 24, 2002 9:06 AM
2
Materials and Methodology
All fungi in the text are isolated, identified, and described on the basis of the following experimental
methods and samples.
13
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Using glass needles with capillary tips, single spores may be separated from a spore mass on
water agar under a dissecting microscope. In this technique, thick plates (5 mm thick) of 3 to 4%
plain water agar may be prepared. Isolations may be practiced with aseptic glass needles sterilized
by soaking in boiled water for each isolation procedure.
1. Wash plant materials under running tap water for at least 30 min.
2. For isolation from diseased plants, freshly infected parts may be selected, and they are cut into tissue
segments of less than 5 mm. From the aged, infected tissues, the more numerous saprophytic fungi
may be isolated.
3. Sterilized plant tissues may be prepared with antiformin (a strongly alkaline solution of sodium
hypochlorite) or ethanol, together with unsterilized ones. Concentrations of chemicals may be
different, according to the samples used, but generally they may be soaked in 1 to 5% antiformin
or 70% ethanol for 30 sec to 5 min. Unsterilized samples may be included to reduce failure of
isolation of the fungi susceptible to such chemicals. Part of the samples used for microscopic
examination may also be used for samples for fungal isolation.
4. Plant tissue segments placed on isolation media are incubated at the appropriate temperature for
1 to 7 days. The isolation media, specific for individual fungi, may be used, but for general purposes,
plain water agar may be one of the best media because bacteria and some contaminated fungi may
be suppressed for lack of nutrition, and individual hyphae can be observed readily during isolation
procedures. The treated plates may be incubated, under lower temperatures below 15°C, at 20 to
25°C, the optimum for most fungi, and at higher temperatures above 34°C. On the plates incubated
under variable temperatures, morphogenesis may be stimulated, resulting in swift sporulation.
Identification practices may become easy.
5. Hyphae may be elongated from the tissue segments plated within a few days, and single hyphal
tippings may be practiced as soon as possible to get rid of extra contamination. The plates used may
be further incubated for continuous observations and finally, ascocarps and other fruiting structures
may be formed on such plates.
6. For isolation of Pythiaceous fungi, including Aphanomyces spp., the fungi may be trapped initially
by susceptible plants or some other trapping substrates, and these materials may be soaked further
under water for observation and isolation.
1. Use some rich agar media, including cornmeal agar, Czapek (Dox) agar, malt agar, oatmeal agar,
PDA, and V-8 juice agar. Some Ascomycetous fungi may often form fruiting structures in agar after
a long incubation period, and therefore the quantity of media in the plates should be increased.
2. Alter the cultural environments, including light conditions and temperatures, for successful sporulation.
3. Change the balance of nutrition in agar. The cultures may be drastically changed from nutrient-rich to
poor nutrient cultures, including plain water agar culture.
4. Use natural media prepared by mixing dried and propylene oxide-treated plant tissues into agar
(Hansen and Snyder, 1947).
5. Soak a bit of culture, infected plant tissue, or substrate in pond water, well water, or petri salts
solution to induce zoospore discharge for some Mastigomycetous fungi.
6. Case (cover) cultures with soils, or grow in wood chip medium, including rice bran, to obtain
successful fruiting for some Basidiomycetous fungi.
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3
Identification of Fungi
Fungus species, or the taxa (singular: taxon) are usually named on the basis of morphological
characteristics. The names of fungi may be determined by comparing the already known species
in morphologies. Generally speaking, any fungi may be identified if they are known species, and
their morphologies are observed clearly.
Observations may be conducted at various levels from the naked eye through stereomicroscope
or compound microscope to electron microscope. However, on the basis of satisfactory observations
by compound microscope, identification may be possible, and the fungi may be correctly named
due to individual abilities of observation of morphologies, or technical abilities for inducing
sporulation in agar cultures.
17
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Keys are based on the standard and general characteristics, but some keys are too artificial by
nature, including some exceptions. Therefore, we just refer to such selections after keying out.
Some fungi may be named without detailed studies and, therefore, with further studies, other
significant characteristics may be found and added; old literature may be consulted later, resulting
in reclassification or the production of synonyms or new combinations.
All these things may occur routinely, and we understand this readily because there are numerous
synonyms. Therefore, without completing identification just by following the keys, we have to
check and consult fungi on the basis of the original descriptions, often comparing them with the
type of specimen. At any rate, overall judgment is essential for identification.
Identification of Fungi 19
The following points should be paid attention for observation, although there are some differ-
ences in individual fungi. In addition, physiological characteristics such as temperature responses
and host ranges are included in the keys, and should be similarly studied, together with morpho-
logical characteristics.
A. Cultural Characteristics
1. Color and tint in colony surface and reverse
2. Smell or fragrance
3. Quantity of aerial hyphae
4. Colony surface texture: cottony, shrunken, sloppy, resupinate, velvety, powdery (floury),
crustaceous, water soaked, embedded, yeast-like, sticky, homogeneous or heterogeneous,
presence or absence of elevation
5. Colony margin: smooth, irregular, restricted, spreading
6. Pattern: zonate, radiate, flowery, arachnoid
7. Pigment exuded: color, watery
8. Organs formed: fruiting structures, sclerotia, rhizomorphs, synnema, sporodochia, stroma, setae
B. Morphology
1. Size: length, width, thickness, etc.
2. Color: refer to standard color charts (i.e., Rayner (1970), Ridgway (1912))
3. Shape
a. General characteristics for all fungi
Hyphae (septate, aseptate, septum location, clamp connection, hyphopodia), appressoria,
chlamydospores, rhizomorphs, synnema (pl. -ta), and others
b. Differences in the respective classes
Mastigomycetous fungi: oogonium (pl.-a), antheridium (pl. -a), oospores (plerotic, aplerotic),
sporangium (pl. -a), or hyphal swellings (hypha-like sporangia, sphaerosporangia,
lobate sporangia)
Zygomycetous fungi: sporangium (columella, apophysis), sporangiola, sporangiophores,
sporangiospores, zygospores, rhizoid, creeping hyphae, zygospores
Ascomycetous fungi: ascocarp (nakid ascocarp, perithecium, apothecium, etc.), appendages
(hairs, setae), ascoma wall [peridium, tissue (textura) type], stroma, ascus (pl. asci)
(disposition, apical structure, evanescent or nonevanescent), paraphysis (pl. paraphyses),
ascospores
Basidiomycetous fungi: basidiocarp (volva, stipe, annulus, umbrella, hymenium (location,
formation pattern, shape, gill, pore, needle shape, cystidia), basidia, spore print,
basidiospores
Mitosporic fungi (Deuteromycotina): apothecium (pl. -a), pycnidium (pl. -a), sporodochium
(pl. -a), conidiophore (erect, creeping, resupinate, simple, or branched; branching pattern),
conidial types (aleuriosporae, annellosporae, arthrosporae, blastosporae, phialosporae,
porosporae, radulasporae, sympodulosporae), papulaspore
4. Number: number of spore septum, zoospore flagellum, oospores per oogonium, antheridium per
oogonium, oil globules per spore, ascospores per ascus, and basidiospores per basidium, etc.
5. External and internal structures of tissues: smooth, echinulate, warty, presence or absence of
hair, texture of peridium, component tissues of ascocarp or pycnidia (conidiocarp), component
hyphae in Basidiomycetes (presence or absence of primary hyphae, skeleton hyphae, and/or
uniting hyphae)
6. Presence or absence of protuberances: number and shape
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C. Physiological Characteristics
1. Temperatures: growth temperatures, optimum temperature for growth, cardinal temperatures
for growth, growth rate
2. Growth media and nutritional requirements: suitable media for sporulation and growth
3. Reaction for reagent and staining: lactophenol, cotton blue, acid fuchsin, KOH, FeSO4, Melzer
reagent
4. Resistance against chemicals
5. Hyphal anastomosis
6. Co-culture reaction
7. Parasitic nature, pathogenicity
4
Key to Classes of Soil Fungi
Although soil fungi now belong to three kingdoms, viz. Kingdom of Protozoa including Plasmo-
diophora, Kingdom of Chromista including Phytophthora and Pythium, and Kingdom of Fungi
including the rest of fungi according to the recent classification system (Hawksworth et al., 1995),
they are treated as they were without the special separation.
Key words: ascospore(s), basidiospore(s), clamp connection, conidium (pl. -a), hypha (pl. -e), oospore(s),
rhizomorph, sclerotium (pl. -a), sporangiospore(s), zoospore(s), zygospore(s)
1. Hyphae aseptate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
septate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
7. Spores formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
none . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
21
1118-ch04-Frame Page 22 Sunday, February 24, 2002 9:08 AM
KEY TO MASTIGOMYCETES,
KINGDOM OF CHROMISTA
Key words: antheridium (pl. -a), chlamydospore(s), oogonium (pl. -a), oospore(s), sporangiospore(s), spo-
rangium (pl. -a), vesicle, zoospore(s), zoospore differentiation, zoosporangium (pl. -a)
3. Sporangia globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
not so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
6. Zoosporangia hypha-like. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
not so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
4. Sporangia globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
flask-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Saksenaea
7. Sporangia columellate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
without columella or poorly columellate . . . . . . . . . . . . . . . . . . . . Mortierella
3. Ascospores 1-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
over 2-celled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
5. Ascocarps hairy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
setose, and cylindrical asci and pigmented ascospores . . . Phaeotrichosphaeria
Key: none
1118-ch04-Frame Page 25 Sunday, February 24, 2002 9:08 AM
1. Conidiomata formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
not formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Conidia formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
not formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sterile fungi
A. Pycnidium-Forming Fungi
Key words: aggregate, appendage, conidia, discrete, filiform, ostiole, pycnidia, setae
1. Pycnidia globose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
occasionally cup-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hainesia
2. Conidia 1-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
over 2-celled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
3. Conidia hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
8. Conidia 2-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
over 3-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stagonospora
9. Conidia hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
B. Sporodochium-Forming Fungi
Key words: appendage, conidia, conidiophores, filiform, setae, sporodochia, vesicle
1. Setae formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
not formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3. Conidia 1-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
over 2-celled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wiesneriomyces
6. Conidia simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
complicated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
C. Synnema-Forming Fungi
D. Aleuriosporae
Key words: clamp connection, conidia, conidiophore, septa, setae, sterigma (pl. -ta)
1. Conidia 1-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
over 2-celled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Conidia hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
E. Arthrosporae
F. Blastosporae
9. Conidia 1-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
over 2-celled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephaliophora
11. Conidia formed at the apex of conidiogenous cell, 1–16 per cell . . . . Oedocephalum
formed apically and laterally, numerous . . . . . . . . . . . . . . . .Chromelosporium
G. Phialosporae
Key words: appendage, catenulation, conidia, conidiophores, fertile (conidium-forming) area, foot cell,
penicillate, phialide(s), spore mass
6. Conidia pigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
hyaline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
9. Conidia branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
almost unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stachybotrys
H. Porosporae
Key words: beak, conidia, conidiophore(s), longitudinally or transversely septate, sympodulate
4. Conidia catenulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
not so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
I. Sympodulosporae
Key words: appendages, biconical, conidia, conidiophore(s), filiform, hilum, sporogeneous (fertile) area
2. Conidia 1-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
over 2-celled. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
5
List of Fungus Genera
MASTIGOMYCOTINA
1. Aphanomyces 5. Pythiogeton
2. Dictyuchus 6. Pythium
3. Phytophthora 7. Saprolegnia
4. Plectospira
ZYGOMYCOTINA
1. Absidia 7. Mucor
2. Circinella 8. Rhizopus
3. Cunninghamella 9. Saksenaea
4. Gongronella 10. Syncephalastrum
5. Helicocephalum 11. Umbelopsis
6. Mortierella 12. Zygorhynchus
ASCOMYCOTINA
1. Achaetomium 9. Microascus
2. Anixiella 10. Monosporascus
3. Apiosordaria 11. Nectria
4. Chaetomium 12. Phaeotrichosphaeria
5. Didymella 13. Preussia
6. Eudarluca 14. Sordaria
7. Glomerella 15. Thielavia
8. Massarina 16. Zopfiella
BASIDIOMYCOTINA
Armillaria, Coprinus, and six taxa
35
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DEUTEROMYCOTINA (continued)
85. Scolecobasidium 101. Thielaviopsis
86. Selenophoma 102. Thysanophora
87. Sepedonium 103. Torula
88. Septonema 104. Torulomyces
89. Spegazzinia 105. Trichocladium
90. Sporidesmium 106. Trichoderma
91. Sporobolomyces 107. Trichothecium
92. Sporoschisma 108. Trichurus
93. Sporotrichum 109. Trinacrium
94. Stachybotrys 110. Tripospermum
95. Stagonospora 111. Tritirachium
96. Staphylotrichum 112. Ulocladium
97. Stemphylium 113. Vermispora
98. Taeniolella 114. Verticillium
99. Tetracladium 115. Volutella
100. Tetraploa 116. Wiesneriomyces
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6
MORPHOLOGY OF
SOIL FUNGI
Mastigomycotina
39
1118-ch06A-Frame Page 40 Sunday, February 24, 2002 4:26 PM
Aphanomyces de Bary
Jahrb. Wiss. Bot. 2:178, 1860.
Type species: A. stellatus de Bary
A B
Aphanomyces cladogamus Drechsler
Aphanomyces cladogamus. A,B: Hypha-like sporangia containing encysted zoospores in a row (A) and
bearing a mass of encysted zoospores at the tip (B). C–E: Oogonia, antheridia, and oospores.
1118-ch06A-Frame Page 41 Sunday, February 24, 2002 4:26 PM
Mastigomycotina 41
Dictyuchus Leitg.
Bot. Ztg. 26:503, 1868.
Type species: D. monosporus Leitg.
Dictyuchus sp.
References: Johnson 1951; Nagai 1931, 1933; Padgett and Seymour 1974;
Rattan et al., 1979.
C
Morphology: Zoosporangia inflated, terminal, cylindrical with the primary A
zoospores developed, and readily encysted inside the sporangia, thus showing
a net-like appearance, each zoospore releasing a secondary zoospore. Immature
sexual organs with irregularly distributed spines are formed. Chlamydospores
(gemmae) globose or subglobose.
Dimensions: Zoosporangia 125–250 × 20-30 µm. Oogonia 27–28 µm in
diameter. Chlamydospores 22–23 µm in diameter. B
Dictyuchus sp. A: Zoosporangium with encysted zoospores. B: Sexual organs, chlamydospores, and vacant
and intact zoosporangia.
1118-ch06A-Frame Page 42 Sunday, February 24, 2002 4:26 PM
Phytophthora de Bary
J. R. Agric. Soc. 2, 12:240, 1876.
Type species: P. infestans (Mont.) de Bary
Key to Species
5. Chlamydospores single . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
catenulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. cyptogea
Mastigomycotina 43
Phytophthora capsici. A,B: Sporangia. C: Differentiated zoospores inside the sporangium and their release.
1118-ch06A-Frame Page 44 Sunday, February 24, 2002 4:26 PM
F G
Phytophthora cryptogea. A–E: Sporangia with external and internal proliferation (E). F,G: Clustered (F) or
simple hyphal swellings.
1118-ch06A-Frame Page 45 Sunday, February 24, 2002 4:26 PM
Mastigomycotina 45
Remarks: This fungus was trapped from soil with potato cubes as the substrate.
Phytophthora megasperma. A–D: Sporangia and external (B,C) and internal proliferation (D). E,F: Oogonia,
antheridia, and oospores (E: paragynous; F: amphigynous). G: Twisted and knobby hypha.
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Mastigomycotina 47
Phytophthora melonis. A,B: Sporangia. C: Oogonium with amphigynous antheridium, and oospore.
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Mastigomycotina 49
Plectospira Drechsler
J. Agric. Res. 34:288, 1927.
Type species: P. myriandra Drechsler
Mastigomycotina 51
Plectospira myriandra. A: Sporangia. B,C: Sporangia and encysted zoospores at the exit (C). D,E: Oogonia
and antheridia. F: Oospores formed parthenogenetically (left) and sexually (right). (From Watanabe, T. 1987.
Mycologia, 79:77–81. With permission.)
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Pythiogeton Minden
Mycol. Unters. 2(2):241, 1916.
Type species: P. ramosum Minden
Pythiogeton ramosum. A–C: Cylindrical sporangia on slender sporogenous hyphae. D: Germinated sporangium
with a single germ tube. E: Vacant sporangium with an emission tube. F: Zoospores encysted. (From Watanabe,
T. 1974a. Trans. Mycol. Soc. Jpn., 15:343–357. With permission.)
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Mastigomycotina 53
Pythium Pringsheim
Jahrb. Wiss. Bot. 1:304, 1858.
Type species: P. monospermum Pringsheim
Key to Species
6. Sporangia single . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
catenulate, deciduous . . . . . . . . . . . . . . . . . . . . . . . . . . P. intermedium
Mastigomycotina 55
Pythium acanthicum. A–C: Sporangia (A), oogonia, antheridia, and oospores. (From Watanabe, T. 1974a.
Trans. Mycol. Soc. Jpn., 15:343–357. With permission.)
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Mastigomycotina 57
Pythium acanthophoron. A–D: Oogonia, antheridia, and oospores. (From Watanabe, T. 1990b. Ann. Phyto-
pathol. Soc. Jpn., 56:549–556. With permission.)
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Pythium afertile. (A,B) and H-Zs (C–E). A,C,D: Vesicle formed from hypha-like sporangia. B: Chlamydospore-
like structures. E: Somewhat thick-walled dendroid hyphae. (From Watanabe, T. et al. 1977. Phytopathology,
67:1324–1332. With permission.)
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Mastigomycotina 59
Pythium angustatum. A: Zoospore differentiation inside vesicle developed from hypha-like sporangium.
B–D: Oogonia and antheridia (B,D: monoclinous; C: diclinous). (From Watanabe, T. et al. 1977. Phytopathology,
67:1324–1332. With permission.)
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Mastigomycotina 61
Pythium aphanidermatum. A–D,F: Sporangia and vesicle formation. E: Encysted zoospores and germination.
G: Sporangia and sexual organs (diclinous). H–J: Oogonia, antheridia, and oospores. (From Watanabe, T. 1974a.
Trans. Mycol. Soc. Jpn., 15:343–357. With permission.)
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B E
Pythium apleroticum. A–C: Oogonia, antheridia, and oospores (B). D: Conidia. E: Vesicle formation. (From
Watanabe, T. 1974a. Trans. Mycol. Soc. Jpn., 15:343–357. With permission.)
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Mastigomycotina 63
Pythium carolinianum. A–C: Sporangia and vesicle formation (B,C). D: Internally proliferated sporangium.
(From Watanabe, T. et al. 1977. Phytopathology, 67:1324–1332. With permission.)
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Pythium catenulatum. A,B: Lobate sporangia (A) and conidia (B). C: Direct germination of conidium.
D,E: Zoospore differentiation in vesicle (D) and zoospore discharge (E). F: Encysted zoospores. G,H: Sexual
organs and oospores (H). (From Watanabe, T. 1974a. Trans. Mycol. Soc. Jpn., 15:343–357. With permission.)
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Mastigomycotina 65
Mastigomycotina 67
D E
Pythium dissimile. A,B: Sporangia and oospores formed parthenogenetically (B: close-up of A).
C–D: Oospores formed parthenogenetically. E: Catenulate appressorium-like hyphal swellings. (From
Watanabe, T. 1990b. Ann. Phytopathol. Soc. Jpn., 56:549–556. With permission.)
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Pythium dissotocum. A: Vesicle formation from hypha-like sporangium. B,C: Oogonium, sessile antheridium,
and oospore.
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Mastigomycotina 69
Pythium echinulatum. A: Immature oogonium. B: Oogonia and antheridium. C,D: Oogonia containing
plerotic oospores. E: Direct germination of sporangium. F,G: Oogonia, aplerotic oospores, and a part of the
sporangium (G). (From Watanabe, T. et al. 1977. Phytopathology, 67:1324–1332. With permission.)
1118-ch06A-Frame Page 70 Sunday, February 24, 2002 4:26 PM
Pythium elongatum. A,B: Lobate sporangia and vesicle formation. C: Inflated hyphae. (From Watanabe, T.
et al. 1977. Phytopathology, 67:1324–1332. With permission.)
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Mastigomycotina 71
Pythium graminicola. A–C,E,H: Oogonia and antheridia. D: Appressoria. F: Oospore. G: Lobate sporangia.
I,J: Immature oogonium-like structures formed aerially and on agar. K: Vesicle formation. L: Germination of
encysted zoospores. (From Watanabe, T. 1974a. Trans. Mycol. Soc. Jpn., 15:343–357. With permission.)
1118-ch06A-Frame Page 72 Sunday, February 24, 2002 4:26 PM
Pythium cf. indigoferae. A–D: Lobate hyphal swellings and sexual organs with aplerotic oospores. Note
parthenogenetically developed oospores (C,D). (From Watanabe, T. et al. 1998. Mycologia Helvetica, 10:3–13.
With permission.)
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Mastigomycotina 73
Pythium inflatum. A,B: Lobate sporangia. C–E: Oogonia and antheridia. F: Oospore. G–I: Vesicle formation.
J: Germination of encysted zoospores. (From Watanabe, T. 1974a. Trans. Mycol. Soc. Jpn., 15:343–357. With
permission.)
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Pythium intermedium. A: Globose sporangia in a chain. B: Young sporangia basipetally developed. C–F: Detached
sporangia. G,H: Direct germination of sporangia. I: Chlamydospore. J: Inflated hyphae. (From Watanabe, T.
et al. 1977. Phytopathology, 67:1324–1332. With permission.)
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Mastigomycotina 75
Pythium myriotylum. A,B: Lobate sporangia and vesicle formation (B). C: Appressorium. D–F: Oogonia,
antheridia, and oospores (F). (From Watanabe, T. 1977a. Ann. Phytopathol. Soc. Jpn., 43:306–309. With permission.)
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Mastigomycotina 77
Pythium nayoroense. A,C,D: Sphaerosporangia and vesicle formation. B: Sporangia and subsporangial
hyphal swellings.
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Pythium oedochilum. A–C: Sphaerosporangia with vesicle formation (B) and internally proliferated (C).
D,E: Oogonia, antheridia, and oospores (E). F: Chlamydospore. (From Watanabe, T. et al. 1977. Phytopathology,
67:1324–1332. With permission.)
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Mastigomycotina 79
Pythium paroecandrum. A,B: Sphaerosporangia and sexual organs. C,D: Sphaerosporangia and vesicle for-
mation (D). E–G: Oogonia, antheridia, and oospores (E,G). (From Watanabe, T. 1985. Trans. Mycol. Soc. Jpn.,
26:41–45. With permission.)
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Mastigomycotina 81
Pythium periplocum. A: Culture on PDA. B,C: Oogonia, antheridia, and oospores (C). D: Lobate sporangia.
(From Watanabe, T. et al. 1987b. Trans. Mycol. Soc. Jpn., 28:475–481. With permission.)
1118-ch06A-Frame Page 82 Sunday, February 24, 2002 4:26 PM
Pythium rostratum. A,B: Sporangia (A), oogonia, antheridia, and oospores (A,B). (From Watanabe, T. 1988b.
Ann. Phytopathol. Soc. Jpn., 54:523–528. With permission.)
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Mastigomycotina 83
Pythium salpingophorum. A,B: Intercalary intact and vacant globose hyphal swellings. C-E: Oogonia with
monoclinous (C,D) and diclinous antheridia (E).
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Pythium spinosum. A,B: Oogonia, antheridia, and oospores (B). C: Sporangia. D: Immature oogonia with
digitate protuberances. (From Watanabe, T. et al. 1977. Phytopathology, 67:1324–1332. With permission.)
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Mastigomycotina 85
Pythium splendens. A: Sporangia. B,C: Aborted oogonia and antheridia. (From Watanabe, T. et al. 1977.
Phytopathology, 67:1324–1332. With permission.)
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D
C
Pythium sulcatum. A: Sporangia. B,C: Oogonia and antheridia. D: Oospores. (From Watanabe, T. et al. 1986b.
Ann. Phytopathol. Soc. Jpn., 52:287–291. With permission.)
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Mastigomycotina 87
Pythium sylvaticum. A,B: Sporangia. C–E: Oogonia and antheridia. F: Sporangium surrounded by hypha-like,
appressorium-like structures. (From Watanabe, T. et al. 1977. Phytopathology, 67:1324–1332. With permission.)
1118-ch06A-Frame Page 88 Sunday, February 24, 2002 4:26 PM
Pythium torulosum. A,B: Vesicle formation from lobate sporangia. C–F: Oogonia, antheridia, and oospores (E–G).
(From Watanabe, T. et al. 1977. Phytopathology, 67:1324–1332. With permission.)
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Mastigomycotina 89
Pythium ultimum. A–C: Oogonia, antheridia, and oospores. D: Sporangia and sexual organs. (From
Watanabe, T. et al. 1977. Phytopathology, 67:1324–1332. With permission.)
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Pythium vexans. A: Vesicle formation from sphaerosporangium. B,C: Oogonia, antheridia, and oospores (C).
(From Watanabe, T. 1986. Trans. Mycol. Soc. Jpn., 26:41–45. With permission.)
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Mastigomycotina 91
Saprolegnia Nees
Nova Acta Leop.-Carol. 11:513, 1823.
Type species: S. ferax (Gruith.) Thuret
Saprolegnia anisospora. A,B: Oogonia, diclinous antheridia, and eccentric oospores. C–F: Sexual organs
bearing some oospores per oogonium. G: Zoosporangia. H: Gemmae.
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MORPHOLOGY OF
SOIL FUNGI
Zygomycotina
93
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Absidia repens. A–C: Sporangiophores and sporangia. D,E: Sporangia after deliquescence showing apophyses,
columellae, and sporangiospores.
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Zygomycotina 95
Circinella muscae. A,B: Sporangiophores, sporangia, and columella (B) and sporangiospores. C: Sporangio-
spores. D. Chlamydospore.
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Cunninghamella Matr.
Cunninghamella echinulata. A–F: Sporangiophores, vesicles (C–F), and sporangiospores. G: Hyphal swellings.
(From Watanabe, T. 1975a. Trans. Mycol. Soc. Jpn., 16:18–27. With permission.)
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Zygomycotina 97
Cunninghamella elegans. A–D: Sporangia, and sporangiophores showing vesicles (B–D), and sporangio-
spores (B,C).
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Gongronella Ribaldi
Riv. Biol. Gen., N. S. 44:164, 1952.
Type species: G. urceolifera Ribaldi = G. butleri (Lendn.) Peyronel &
Dal Vesco
Gongronella butleri. A,C,D: Sporangiophores and sporangia. B,E: Columellae and apophyses. F: Sporangio-
spores. G: Rhizoid. H: Chlamydospores. (From Watanabe, T. 1975a. Trans. Mycol. Soc. Jpn., 16:18–27. With
permission.)
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Zygomycotina 99
Helicocephalum Thaxter
Bot. Gaz. 16:201, 1891.
Type species: H. sacophilum Thaxter
Helicocephalum oligosporum. A–D: Sporangiophores (A,B), sporangia (B,C), and sporangiospores (C,D).
Note apexes of sporangiophores after deliquescence of sporangia (A,D). C,E: Rhizoids. (From Watanabe, T.
and Koizumi, S. 1976. Trans. Mycol. Soc. Jpn., 17:1–3. With permission.)
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Mortierella Coemans
Bull. Acad. R. Sci. Belg. 2, 15:288, 1863.
Type species: M. polycephala Coemans
Key to Species
1. Sporangiospores formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
not formed, chlamydospores echinulate . . . . . . . . . . . . . . . M. chlamydospora
2. Sporangiophores simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
branched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Zygomycotina 101
Mortierella alpina. A,B: Sporangiophores and sporangia. C,D: Sporangiospores on deliquescens of sporangia.
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Zygomycotina 103
Zygomycotina 105
Zygomycotina 107
Mortierella epicladia var. chlamydospora. A,B: Sporangiophores and sporangia. C–E: Sporangiophore and
sporangiospores. F: Sporangiophore and chlamydospore. (From Watanabe, T. et al. 2001e. Three New Mortier-
ella from Soil in the Bonin Islands, Japan. In preparation. With permission.)
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A, B
Zygomycotina 109
A C
Mortierella humilis. A,B: Sporangiophore and sporangiospores. C: Apical part of sporangiophore and
sporangiospores.
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Zygomycotina 111
Zygomycotina 113
Zygomycotina 115
Zygomycotina 117
Mortierella zychae. A: Sporangiophores and sporangium. B,C: Apical parts of sporangiophores and sporangio-
spores. D: Sporangiospores. E: Apex of sporangiophore. F,G: Rhizoids. H–J: Chlamydospores. (From Watanabe, T.
1975a. Trans. Mycol. Soc. Jpn., 16:18–27. With permission.)
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Zygomycotina 119
Key to Species
Zygomycotina 121
Mucor hachijoensis. A: Sporangium and zygospore. B: Columellae. C: Zygospore with opposite unequal
suspensors. D: Sporangiospores. E: Chlamydospores and hyphae. (From Watanabe, T. 1994b. Mycologia,
86:691–695. With permission.)
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Mucor hiemalis f. luteus. A: Sporangiophores and sporangia. B: Columella on the apical part of sporangiophore.
C: Sporangiospores. (From Watanabe, T. 1971. Trans. Mycol. Soc. Jpn., 12:35–47. With permission.)
1118-ch06A-Frame Page 123 Sunday, February 24, 2002 4:26 PM
Zygomycotina 123
Mucor microsporus. A: Sporangiophores and sporangia. B: Collapsed sporangia and apical parts of sporangio-
phores. C: Columella on apical sporangiophore and sporangiospores. D: Rhizoids.
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Mucor plumbeus. A,B: Sporangiophores and columellae on apical parts of sporangiophores and sporangio-
spores (A). C: Chlamydospore.
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Zygomycotina 125
Rhizopus Ehrenb.
Icon. Fung. 2:20, 1838.
Type species: R. stolonifer (Ehrenb. : Fr.) Vuill.
Rhizopus oryzae. A: Sporangiophore with columella on dehiscence of sporangium and rhizoid. B: Columella
and sporangiospores. C: Sporangiospores. D: Chlamydospore-like structure at the basal part of sporangiophore.
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Saksenaea Saksena
Mycologia 45:434, 1953.
Type species: S. vasiformis Saksena.
Saksenaea Saksena
Saksenaea vasiformis. A,B: Sporangiophores, sporangia, and sporangiospores. C: Rhizoids. (From Watanabe, T.
1971. Trans. Mycol. Soc. Jpn., 12:35–47. With permission.)
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Zygomycotina 127
Syncephalastrum Schroeter
Krypt. Fl. Schles. 3(2):217, 1886.
Type species: S. racemosum (Cohn) Schroeter
References: Amos and Barnett 1966; Arx 1982; Kendrick et al. 1994;
Watanabe et al. 2001.
C Morphology: Sporangiophores hyaline, erect, short, forming vesicles
(globose inflations) at the middle, elongating 2 to more than 12 branches
with the terminal single sporangia, often proliferated from the sporangia.
Sporangiospores hyaline, 1-celled, deciduous.
Dimensions: Sporangiophores 1–50 µm long from base to primary
vesicle, 1.6–4 µm wide, 10–54 µm long from primary vesicles to
sporangia. Sporangia (3-) 4–7 (-10) µm in diameter. Sporangiospores
2.5–3 (-4.5) µm in diameter.
D
Material: 99-454 (= MAFF 238015, Forest soil, Takabotchi, Shiojiri,
Nagano, Japan), 99-481 (= MAFF 238018, Forest soil, Kataoka,
Shiojiri, Nagano, Japan); 99-485 (= MAFF 238019, Forest soil, Ina,
Nagano, Japan).
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Zygomycotina 129
Umbelopsis nana. A: Six-day-old colonies of two isolates on PDA. B–E: Sporangiophores and sporangia.
F: Sporangiophores after detachment of sporangiospores, and sporangiospores. G,H: Proliferated and aggregated
sporangiophores and sporangia with intercalary chlamydospores and vacant hyphal swelling (H). (From
Watanabe, T. et al. 2001b. Mycoscience. In Press. With permission.)
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Umbelopsis vinacea. A: Sporangiophores and sporangia. B,C: Vesicles and sporangia. D: Sporangiophore
and sporangiospores. E: Sporangiospores. F: Chlamydospores. (From Watanabe, T. 1977b. Trans. Mycol. Soc.
Jpn., 18:242–244. With permission.)
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Zygomycotina 131
Zygorhynchus Vuill.
Bull. Soc. Mycol. Fr. 19:116, 1903.
Type species: Z. heterogamus Vuill.
MORPHOLOGY OF
SOIL FUNGI
Ascomycotina
133
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Ascomycotina 135
Achaetomium sp. A,B: Erumpent (A) and embedded perithecia (B). C: Asci, paraphyses and ascospores.
D: Aureobasidium anamorph.
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Ascomycotina 137
References: Arx and Gams 1966; Mirza and Cain 1969; Stchigel et al. 2000;
Udagawa and Furuya 1972; Watanabe 1971.
D B
Morphology: Perithecia mostly embedded, black, flask-shaped, ovate, soft, api-
cally papillate: peridium with soft skin, pseudoparenchymatous. Asci hyaline, clavate, 4-ascosporous.
Ascospores 2-celled, composed of black, ovate, granulate, thick-walled cell, and hyaline, triangle cell. Para-
physes present.
Dimensions: Perithecia 210–340 × 220–290 µm: ostioles 30–35 µm in diameter. Asci 125–150 × 12.5–15 µm.
Paraphyses 110–120 × 2.7–2.8 µm. Ascospores 31–35 × 13–14.8 µm: pigmented cells 20.5–22.5 × 13–14.8 µm:
hyaline cells 10.5–12.5 × 7.2–7.8 µm.
Material: 69-300 (= ATCC 24150, Pineapple field soil, Okinawa, Japan).
Remarks: A key to the 16 soilborne species was provided by Stchigel et al. (2000).
Apiosordaria verruculosa var. maritima. A: Perithecium. B,C: Asci and ascospores. D: Ascospores.
E: Cladorrhinum st. (From Watanabe, T. 1971. Trans. Mycol. Soc. Jpn., 12:35–47. With permission.)
1118-ch06A-Frame Page 138 Sunday, February 24, 2002 4:26 PM
Chaetomium Kunze
Mykol. Hefte 1:16, 1817.
Type species: C. globosum Kunze
Key to Species
3. Perithecia ellipsoidal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
vase-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Ascomycotina 139
Chaetomium aureum. A: Part of perithecium. B: Part of terminal hairs. C: Asci and ascospores.
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Ascomycotina 141
Ascomycotina 143
Chaetomium erectum. A,B: Perithecia and terminal hairs. C: Asci and ascospores. D: Ascospores.
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Chaetomium funicola. A,B: Perithecia and terminal hairs. C: Asci and ascospores.
1118-ch06A-Frame Page 145 Sunday, February 24, 2002 4:26 PM
Ascomycotina 145
Chaetomium fusiforme. A: Perithecia and terminal hairs. B: Asci and ascospores. C: Ascospores. D: Part of
the terminal hairs.
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Ascomycotina 147
Chaetomium reflexum. A: Terminal hairs on perithecium. B: Terminal hair, asci, and ascospores.
1118-ch06A-Frame Page 149 Sunday, February 24, 2002 4:26 PM
Ascomycotina 149
Chaetomium spirale. A: Perithecium and terminal hairs. B: Ascospores. C: Asci and ascospores.
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Chaetomium torulosum. A: Perithecium. B: Asci and ascospores from crushed perithecium. C: Ascospores.
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Ascomycotina 151
Chaetomium virescens. A,C: Perithecia. B: Terminal hairs and ascospores. D,E: Asci and ascospores.
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Didymella Sacc.
Syll. Fung. 1:545, 1882.
Type species: D. exigua (Niessl) Sacc.
A
Didymella effusa (Niessl) Sacc.
Ascomycotina 153
Eudarluca Speg.
Rev. Mus. La Plata 15:22, 1908.
Type species: E. australis Speg.
A
Eudarluca biconica Katumoto
Material: 85-99, 85-100 (= IFO 32539, Flowering cherry seed, Hachioji, Tokyo, Japan).
Eudarluca biconica. A: Perithecia. B: Ascospores in ascus. C: Ascospores. (From Watanabe, T. 1989d. Trans.
Mycol. Soc. Jpn., 30:395–400. With permission.)
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Glomerella glycines. A: Perithecia. B: Asci. C: Ascospores. D: Asci extruded from perithecium. E: Sporodochia.
F: Setae and conidia.
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Ascomycotina 155
Massarina Sacc.
Syll. Fung. 2:153, 1883.
Type species: M. eburnea (Tul.) Sacc.
Massarina sp. 1. A
Massarina sp. 1. A: Perithecia embedded in agar culture. B: Asci and ascospores. C: Ascospores.
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Massarina sp. 2.
References: Bose 1961; Srinivasulu and Sathe 1972.
A Morphology: Cleistothecia black, globose, subglobose or disc-shaped: peridium brown,
opaque, with soft skin. Asci cylindrical, bitunicate, 8-ascosporus. Paraphyses lacking.
Ascospores hyaline, fusiform, often curved, 4–7-celled.
Dimensions: Cleistothecia 225–300 µm in diameter. Asci 80–100 × 7.5–8.8 µm.
Ascospores 27.5–32 × 3.8–5 µm.
Material: 73-463 (Strawberry field soil, Shizuoka, Japan).
Remarks: This fungus was identified as Massarina species on the basis of ascospore
morphology and its dimensions, but differs from Keissleriella gloeospora (Berk. et Curt)
Bose, closely related to this fungus which bears setae around ostioles.
B C
Massarina sp. 2. A: Cleistothecia embedded in agar culture. B: Part of ascus and ascospores. C: Ascospores.
D: Ascospores in an ascus.
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Ascomycotina 157
Microascus Zukal
Verh. Zool.-Bot. Ges. Wien 35:333, 1885.
Type species: M. longirostris Zukal
Microascus longirostris. A,D: Perithecia. B: Catenulate ascigerous hyphae. D: Asci and ascospores. E: Anamorph.
(From Watanabe, T. and Sato, Y. 1988. Trans. Mycol. Soc. Jpn., 29:143–150. With permission.)
1118-ch06A-Frame Page 158 Sunday, February 24, 2002 4:26 PM
Monosporascus cannonballus. A: Crushed perithecium and ascospores. B: Immature ascus and ascospore,
and paraphyses. C: Cultured perithecia. D–E: Asci and ascospores. (From Watanabe, T. 1979. Trans. Mycol.
Soc. Jpn., 20:312–316. With permission.)
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Ascomycotina 159
Nectria Fr.
Summa Veg. Scand. 287, 1847.
Type species: N. cinnabarina (Tode : Fr.) Fr.
Key to Species
2. Anamorph formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
not formed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .N. asakawaensis
Nectria asakawaensis. A: Perithecium. B: Asci and ascospores. C: Peridium and ascospores. D: Asci and
ascospores. (From Watanabe, T. 1990c. Trans. Mycol. Soc. Jpn., 31:227–236. With permission.)
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Ascomycotina 161
Nectria fragariae. A: Perithecia. B: Peridium, part of ascus and ascospores. C: Ascospores in ascus.
D: Ascospore and conidia. E–H: Anamorph: catenulate conidia (E), part of conidiophores and conidia (F–K),
and phialides (L), M: Chlamydospores. (From Watanabe, T. 1990c. Trans. Mycol. Soc. Jpn., 31:227–236.
With permission.)
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Ascomycotina 163
Nectria gliocladioides. A: Perithecia. B,C: Asci and ascospores. D: Ascospores. E: Anamorph; conidiophores
and conidia. F: Chlamydospore.
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Ascomycotina 165
Nectria hachijoensis. A: Perithecium. B: Peridium. C: Ascospores. D,E: Asci and ascospores. F–K:
Anamorph: Seta-like structures (F,G), conidiophores and conidia (H,I), phialides and conidia (J,K). L: Conidia and
ascospore. M: Chlamydospore. (From Watanabe, T. 1990c. Trans. Mycol. Soc. Jpn., 31:227–236. With permission.)
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Phaeotrichosphaeria Sivanesan
Trans. Br. Mycol. Soc. 81:313, 1983.
Type species: P. indica Sivan. & N. D. Sharma
Phaeotrichosphaeria sp.
A B
References: Sivanesan 1983.
Morphology: Perithecia black or dark brown, flask-shaped or
subglobose, ostiolate, covered with stiff hairs or setae. Asci
hyaline, cylindrical or clavate, 8-ascosporous in 1 row. Para-
physes hyaline, cylindrical, curved, thread-like. Ascospores
ellipsoidal, brown, biguttulate.
C Anamorph: Aureobasidium-like. Conidiophores lacking or
very short. Conidia hyaline, cylindrical. Setal bodies with stiff
setae: setae dark brown or black, septate.
Dimensions: Perithecia 275–300 × 175–200 µm: ostiolar
D
regions 50 µm long, 60–80 µm wide. Asci 60–74 × 4–6 µm.
Ascospores 4.4–7.6 × 3–3.4 µm. Paraphyses nearly 68–70 µm long, 1–2 µm wide. Conidia 5–6 × 1.2–2.4 µm.
Setal bodies 90–180 µm including setae: setae 24–240 × 2.8–4 µm.
Material: 99-467 (Forest soil, Agematsu, Shiojiri, Nagano, Japan).
Remarks: This fungus characteristically forms setal perithecia with brown ascospores, and Aureobasidium
anamorph.
Phaeotrichosphaeria sp. A,B: Tomentose (A) and setose perithecia (B). C: Asci, paraphyses, and ascospores.
D: Dark ascospores, and hyaline conidia in Aureobasidium anamorph.
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Ascomycotina 167
Preussia Fuckel
Fungi Rhenan. Suppl., Fasc. 3:1750, 1866.
Type species: P. funiculata (Preuss) Fuckel
E
Preussia terricola Cain
Preussia terricola. A–C: Asci and ascospores. D: Separated component cells of conidia. E: Cleistothecium.
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Key to Species
2. Perithecia over 200 µm in diameter, with asci over 125 µm long . . . . . . . . . S. fimicola
under 150 µm in diameter, with asci under 117.5 µm . . . . . . . . . S. tamaensis
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Ascomycotina 169
Sordaria fimicola. A: Perithecia. B: Ascospores in ascus. (From Watanabe, T. 1989d. Trans. Mycol. Soc. Jpn.,
30:395–400. With permission.)
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Sordaria nodulifera. A: Perithecia. B: Part of asci and ascospores. C: Ascospore. (From Watanabe, T. 1989d.
Trans. Mycol. Soc. Jpn., 30:395–400. With permission.)
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Ascomycotina 171
Sordaria tamaensis. A: Asci and ascospores extruded from crushed perithecium. B: Part of ascus and
ascospores. C: Ascospore. (From Watanabe, T. 1989d. Trans. Mycol. Soc. Jpn., 30:395–400. With permission.)
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Thielavia Zopf
Verh. Bot. Ver. Brandenb. 18:101, 1876.
Type species: T. basicola Zopf
C
Thielavia terricola (Gilman et Abbott) Emmons
References: Booth 1961; Booth and Shipton 1966; Malloch and Cain 1973.
A Morphology: Cleistothecia superficial, black, globose, covered with white hairs:
D peridium brown, double-membranous, pseudoparenchymatous. Asci ovate, clavate,
narrowed basally with foot cells, 8-ascosporus, evanescent. Ascospores, olive-colored,
dark green, ellipsoidal, with germ pores at one end, truncate at another end, blue lines recognizable.
Anamorph lacking.
Dimensions: Cleistothecia (50-) 75–170 (-190) µm in diameter. Asci (20-) 27.5–35 (-40) × 13.7–17.5 (-19) µm.
Ascospores 9–15.5 × (5-) 6.2–8 µm.
Material: 73-30 (Strawberry root, Shizuoka, Japan); 73-451 (Strawberry field soil, Shizuoka, Japan); 84-532
(Japanese red pine seed, Higashichikuma, Nagano, Japan); 86-64 (Japanese cedar seed, Kasama, Ibaraki, Japan).
Thielavia terricola. A: Cleistothecium. B: Crushed cleistothecium, asci, and ascospores. C: Asci and
ascospores. D: Ascospores.
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Ascomycotina 173
Zopfiella Winter
Krypt. Fl. 1:56, 1887.
Type species: Z. tabulata (Zopf) Wint.
Key to Species
Zopfiella curvata. A,B: Cleistothecia on agar. C: Part of peridium and setae. D,E: Asci and ascospores (E).
F: Ascospores.
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Ascomycotina 175
Zopfiella pilifera. A,B: Asci and ascospores. C: Cleistothecium. (From Watanabe, T. 1971. Trans. Mycol. Soc.
Jpn., 12:35–47. With permission.)
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MORPHOLOGY OF
SOIL FUNGI
Basidiomycotina
177
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Basidiomycetous fungi
References: Warcup 1959; Warcup and Talbot 1962, 1963; Watanabe 1986.
A-1
Morphology: Basidiomycetous fungi are generally not fruitful in agar cultures,
except Coprinus spp. which often fruited in vitro (Figures A,B). Many Basidio-
mycetous fungi have hyphae with clamp-connections (Figures D–I), but others
A-2 do not. Rhizomorphs are often formed in vitro by Armillaria mellea (Vahl:Fr.)
A
Kumm. (Figure C). However, some fungi form arthroconidia (Figure H) or
A-3
chlamydospores. Because of nonfruiting in vitro, identification is very difficult,
although their colonies are clearly differentiated to one another.
H Dimensions: Coprinus sp. (Isolate 85-17): hyphae 2.5–7 µm broad: pileus 8 mm
long: stipe 35–60 mm tall, ca. 1 mm wide basally: basidiospores 5.7–9.5 × 5–6.3 µm:
sclerotia 105–155 µm in diameter.
Material: 73-50, 73-83, 73-145 (Strawberry root, Shizuoka, Japan); 82-735 (Bell
pepper field soil, Wakayama, Japan); 84-481 (Japanese red pine seed, Saihaku,
D-G Tottori, Japan); 84-595 (Japanese red pine seed, Kamiina, Nagano, Japan); 85-P101
(Paulownia root, Itapua, Paraguay); 85-17 (Flowering cherry seed, Hachioji, Tokyo,
Japan); 85-153 (Flowering cherry root, Hachioji, Tokyo, Japan).
Remarks: To identify unknown Basidiomycetous fungi, it is essential to compare them with already known
isolates culturally or cultures derived from natural fruiting structures. Trials for inducing fruiting in agar cultures
with soils under various atmospheres are recommended (Warcup, 1959; Warcup and Talbot, 1962; 1963).
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Basidiomycotina 179
Basidiomycetous fungi. A,B: Coprinus spp. sporulated on agar (A: Isolate 85-17, B: 82-735). C: Rhizomorph
formation of Armillaria mellea (Isolate 85-153) on agar culture. D–G,I: Hyphae with clamp connections on
agar (D: Isolate 84-481; E: 84-595, F: P101, G: 73-50, I: 84-145). H: Hyphae and arthrospores (Isolate 73-83).
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Coprinus Pers.
Basidiomycotina 181
Coprinus sp. A: Colony on PDA with the residues of two fruit bodies. B,C: Part of cap component cells
and basidiospores. D: Basidiospores. E: Part of stipe tissues. F: Unclumped hyphae. F: The fungus on the
sectioned pieces of chopsticks recovered from soil. G,H: Eight-day-old colony on PDA.
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MORPHOLOGY OF
SOIL FUNGI
Deuteromycotina (Motosporic Fungi)
183
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Acremonium Link
Acremonium macroclavatum. A: Conidiophores and spore masses. B: Conidia and crustaceous hyphae.
(From Watanabe, T. et al. 2001b. Mycoscience. In press. With permission.)
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Acremonium sp.
Synonym: Cephalosporium sp.
References: Gams 1971; Obayashi and Watanabe 1987.
Morphology: Conidiophores (phialides) hyaline, erect, simple or branched,
gradually tapering from base toward apex, bearing spherical spore masses termi-
B
nally. Conidia phialosporous, hyaline, 1-celled, cylindrical or long-ellipsoidal,
pointed at one or both ends, with 3–7 oil globules.
Dimensions: Conidiophores (phialides) 17.5–37.5 (-50.4) × 3.2–4 µm. Spore
masses 27.5–45 µm in diameter. Conidia 6–8.5 (-9.3) × 2.1–2.8 (-4) µm.
Material: 87-3 (Radish roots, Miura, Kanagawa, Japan).
Remarks: This fungus causes round, brown lesions on radish roots. For a few
plant pathogens such as C. gramineum Nisikado et Ikata and C. gregatum Allington
et Chamberlain, the genus name Cephalosporium is more commonly used. C
Acremonium sp. A,B: Conidiophores and spore masses. C: Conidiophores and conidia.
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Alternaria alternata. A,B: Conidiophores and conidia. (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn.,
16:264–267. With permission.)
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Arthrinium st.
Arthrobotrys Corda
Prachtflora p. 43, 1839.
Type species: A. superba Corda
Key to Species
Aspergillus brevipes. A: Conidiophore and spore mass. B: Vesicle, phialides, and conidia. C: Conidia.
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Aspergillus fumigatus. A: Conidiophores and spore mass. B: Vesicle and phialides. C: Conidia. (From
Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 193 Sunday, February 24, 2002 7:32 PM
Aspergillus niger. A: Conidiophore and spore mass. B,C: Vesicles, phialides, and conidia (B).
1118-ch06B-Frame Page 194 Sunday, February 24, 2002 7:32 PM
Aspergillus parasiticus. A: Conidiophore and catenulate conidia. B,C: Vesicles, phialides, and conidia. (From
Watanabe, T. 1971. Trans. Mycol. Soc. Jpn., 12:35–47. With permission.)
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Aspergillus sp., Sect. Clavati. A: Conidiophore and spore mass. B: Conidiophore with terminal vesicle.
C: Phialides. D: Conidia. (From Watanabe, T. and Sato, T. 1988. Trans. Mycol. Soc. Jpn., 29:143–150. With permission.)
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Aureobasidium sp.
References: Cooke 1959, 1962; Ellis 1971; Hermanides-Nijhof 1977; Watanabe
A et al. 1986a.
Morphology: Conidiophores lacking. Conidia blastosporous, lateral directly on
hyphae, forming spore masses on hyphae, hyaline, cylindrical, 1-celled.
Dimensions: Conidia 32–54 × 2–5 µm.
Material: 99-5 (Cucumber seed, Tateyama, Chiba, Japan).
B C
Remarks: Cultures on PDA are resupinate, spreading, often black yeast-like.
Beltrania Penzig
Nuovo G. Bot. Ital. 14:72, 1882.
Type species: B. rhombica Penzig
Beltrania rhombica. A,B: Conidiophores and conidia. C: Apex of conidiophore and conidia. (From
Watanabe, T. 1971. Trans. Mycol. Soc. Jpn., 12:35–47. With permission.)
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Biporalis Shoemaker
Can. J. Bot. 37:879, 1959.
Type species: B. maydis (Nisikado) Shoemaker
Key to Species
Biporalis holmii. A,B: Conidiophores and conidia. C: Part of germ tube and end cell of conidium. (From
Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Biporalis sacchari. A–D: Conidiophores and conidia. E: Germ tubes from both end cells of germinated conidium.
(From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Bispora Corda
Icon. Fung. 1:9, 1837.
Type species: B. antennata (Pers. : Fr.) Mason.
Botryodiplodia Sacc.
Syll. Fung. 3:377, 1884.
Type species: B. juglandicola (Schw.) Sacc.
Botryodiplodia sp.
Camposporium Harkn.
Bull. Calif. Acad. Sci. 1:37–38, 1884.
Type species: C. antennatum Harkn.
Camposporium laundonii. A,C: Sporulation on root segment of Aralia elata (substrate). B: Conidium on
substrate. D,E: Sporulation on agar. F: Conidiophore on substrate. G: Conidia formed on agar culture.
H: Catenulate chlamydospores. (From Watanabe, T. 1983b. Trans. Mycol. Soc. Jpn., 24:25–33. With permission.)
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Candelabrella sp.
References: Rifai and Cooke 1966.
Morphology: Conidiophores hyaline, erect, tapering from base toward
apex, bearing 2–4 conidia at sterigmata developed sympodially on the apical
D
fertile part. Conidia sympodulosporous, hyaline, cylindrical, 2-celled with
narrowed basal end. Chlamydospores dark brown, globose.
Dimensions: Conidiophores 45–135 × 2.5 µm. Conidia 22.5–42.5 ×
3.5–6.3 µm. Chlamydospores 12.5–13.8 µm in diameter.
B
E
Material: 74-525 (Strawberry root, Shizuoka, Japan).
A
Remarks: Conidiophores of this fungus are shorter, and conidia are narrower
than those of C. musiformis.
Candida Berkhout
De shimmelgeslachaten Monilia, Oidium, Oospora en Tolura, Thesis
Utrecht (1923).
Type species: C. albicans (Robin) Berkhout
Candida sp.
Cephaliophora Thaxter
Bot. Gaz. 35:153, 1903.
Type species: C. tropica Thaxter
A-D
Chaetomella Fuckel
Symb. Mycol. P. 401, 1870.
Type species: C. oblonga Fuckel
Chaetomella sp. A B
References: Ramchandra Rao & Baheker 1964; Stolk 1963; Sutton & Sarbhoy 1976.
Morphology: Pycnidia black, superficial, ellipsoidal, setose: setae dark, bristle,
gradually tapering toward coiled apex. Conidiophores branched, septate. Conidia
hyaline, 1-celled, cylindrical or fusiform, occasionally slightly curved.
Dimensions: Pycnidia mostly 200–250 × 170 µm: setae mostly 70 µm long.
C D
Conidiophores 20–25 × 1–1.5 µm. Conidia 6–8.5 × 2–2.5 µm.
Material: 75-P-110 (Paulownia root, Pirapo, Paraguay).
Remarks: Although this fungus is close to Amerosporium Speg. on the basis of setose pycnidia and aseptate
conidia, its conidiophores were branched and no raphe was observed on its pycnidia. The conidiophores of
Amerosporium are simple, and no raphe is present.
Chaetomella sp. A,B: Setose pycnidia. C: Crushed pycnidium and extruded conidia. D: Conidiophore.
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Chloridium Link
Syst. Mycol. 1:46, 1821, Figure 76a.
Type species: C. viride (Link) Hughes
Chloridium virescens var. chlamydosporum. A,D: Conidiophores and spore masses in simple (A,D) or pro-
liferated conidiophore (A). B: Conidiophores on dehiscence of spore masses. C: Apexes of conidiophores,
conidia, and chlamydospores. E: Conidiophore apexes bearing two conidia at each apex, and chlamydospores.
F: Conidiophore after dehiscence of spore mass.
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Chromelosporium Corda
in Sturm, Deutschl. Fl., Pilze 3, 3:81, 1833.
Type species: C. ochraceum Corda
C
Chromelosporium fulvum (Link) McGinty, Hennebert & Korf
Chrysosporium Corda
in Sturm, Deutschl. Fl., Pilze 3, 3:13:85, 1833.
Type species: C. merdarium (Link ex Grev.)
Carmichael
Chrysosporium keratinophilum. A,B: Conidiophores and conidia. (From Watanabe, T. 1975c. Trans. Mycol.
Soc. Jpn., 16:149–182. With permission.)
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Cladorrhinum bulbillosum. A: Conidiophores and spore masses. B: Hypha. C,D: Phialides. E: Sclerotia.
F: Component cells of sclerotium. G: Phialides and conidia. (From Watanabe, T. 1975d. Trans. Mycol. Soc.
Jpn., 16:264–267. With permission.)
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Cladorrhinum samala. A,B: Conidiophores and spore masses. C: Phialides and conidia. D: Chlamydospore.
E: Conidiophores. F: Conidia. (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 222 Sunday, February 24, 2002 7:32 PM
Cladosporioum Link
Syst. Mycol. 3:368, 1832.
Type species: C. herbarum Link ex Fr.
Codinaea Maire
Publ. Inst. Bot. Barcelona 3:15, 1937.
Type species: C. aristata Maire
Codinaea parva. A: Conidiophores and spore masses. B–E: Conidiophores and conidia.
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Codinea state of Chaetosphaeria talbotii. A: Conidiophore and spore mass. B: Conidiophore. C: Phialide
and conidia. D: Conidiophores and conidia. E: Conidia.
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Colletotrichum Corda
in Sturm, Deutschl. Fl., Pilze 3:41, 1831.
Type species: C. dematium (Fr.) Grove
Key to Species
2. Conidia lunar-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
cylindrical. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Colletotrichum coccodes. A: Sclerotia with setae. B: Conidiophores, conidia and hyphae. C: Appressoria.
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Colletotrichum truncatum. A. Conidia and setae. B: Sclerotia. C: Sporodochia with setae and sclerotia. (From
Watanabe, T. 1972a. Ann. Phytopathol. Soc. Jpn., 38:106–110. With permission.)
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Coniothyrium Corda
Icon. Fung. 4:38, 1840.
Type species: C. palmarum Corda
Corynespora Güssow
Z. Pflanzenkr. 16:13, 1906.
Type species: C. cassiicola (Berk. & Curt.) Wei
References: Ellis 1963, 1971; Sato and Kitazawa 1980; Wei 1950.
Morphology: Conidiophores hyaline or pale brown, simple, erect, 3- to
7-celled. Conidia porosporous, terminal, solitary or 2–3 conidia in a chain A
acropetally developed, brown, cylindrical, long-ellipsoidal, often curved, occa-
sionally branched, usually 9- to 18-celled, with conspicuous hilum basally.
C
Dimensions: Conidiophores 35 over 250 × 7.5–10 µm. Conidia 165–180 ×
17.5–22.5 µm.
B
Material: 83-191 (Soybean root, Shiojiri, Nagano, Japan).
Remarks: Colonies on PDA are yellowish gray green, zonate. This fungus
is a root and stem rot pathogen of soybean.
C
Remarks: Conidia of this fungus are narrower than those of
A
C. cassiicola.
Corynespora citricola. A,B: Conidiophores and conidia. C: Conidia. D: Apical part of conidiophore and
conidia. E,F: Apex of conidiophore.
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Curvularia Boedijn
Bull. Jard. Bot. Buitenzorg 3, 13(1):127, 1933.
Type species: C. lunata (Wakker) Boedijn
Key to Species
8. Conidia almost uncurved, large and dark in central cells . . . . . . . . . . . . . . . .C. affinis
curved totally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. senegalensis
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B C
Curvularia clavata. A,B: Conidiophores and conidia. C: Conidia and chlamydospore. D,E: A part of
conidiophores and conidia.
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A-C D
B
Teleomorph: Cochliobolus tuberculatus Sivan.
References: Ellis 1971; Jain 1962.
Morphology: Conidiophores erect, (pale) brown, simple, conspicuously
curved, often zig zag-shaped, rarely branched, bearing conidia apically and
laterally. Conidia porosporous, cylindrical or long-ellipsoidal, dark brown,
warted all over, mainly 4-celled, more pigmented and larger in central
2 cells, with or without hilum.
Dimensions: Conidiophores 75–145 × 2.8–5 µm. Conidia 37.5–47.5 ×
10–20 µm.
Material: 79-137 (Taro root, Naha, Okinawa, Japan).
A
Cylindrocarpon Wollenw.
Phytopathology 3:225, 1913.
Type species: C. cylindroides Wollenw.
Key to Species
Cylindrocarpon janthothele. A: Habit. B: Conidiophore and conidia. C,D: Germination of conidia with germ
tubes from both ends.
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Cylindrocladium Morgan
Bot. Gaz. 17:191, 1892.
Type species: C. scoparium Morgan
Key to Species
1. Conidia 2-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
4-celled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Cylindrocladium camelliae. A: Conidiophore with stipe and vesicle (arrow) and chlamydospores. B,F: Part of
the conidiophores and branches. C: Chlamydospores. D:E: Vesicles. G: Part of the conidiophore and conidia.
(From Watanabe, T. 1994a. Mycologia, 86:151–156. With permission.)
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Cylindrocladium colhounii. A,B: Conidiophores, spore masses, stipes, and vesicles. C,D: Part of the
conidiophores, vesicle (arrow), and conidia. E: Catenulate chlamydospores. (From Watanabe, T. 1994a.
Mycologia, 86:151–156. With permission.)
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Cylindrocladium floridanum. A,B: Conidiophores, stipes, vesicles, conidia, and sclerotium (A). B: Conidia
and stipe. C: Sclerotia.
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Cylindrocladium parvum. A,B: Conidiophores, stipes, vesicles, and conidia. C: Conidia. D: Branches and
phialides. E,F: Chlamydospores. (From Watanabe, T. 1992f. Trans. Mycol. Soc. Jpn., 33:231–236. With permission.)
1118-ch06B-Frame Page 256 Sunday, February 24, 2002 7:32 PM
Cylindrocladium scoparium. A: Conidiophore with spore mass, stipe, and vesicle. B: Conidiophore, branches,
and phialides. C: Overview of spore mass. D: Conidia. E,F: Chlamydospores and sclerotium-like structure.
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Cylindrocladium tenue. A–D: Conidiophores and spore masses. E,F: Conidiophores and conidia. G: Conidia.
H–J: Branches, phialides, and conidia. K: Catenulate chlamydospores. Note conidia borne on phialides (H,I).
(From Watanabe, T. 1994a. Mycologia, 86:151–156. With permission.)
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Cylindrocladium sp.
References: Crous and Wingfield 1994; Peerally 1991; Watanabe 2001c.
Morphology: Conidiophores hyaline, erect, branched 1–3 times, mainly
D
penicillate at the fertile portions, bearing spore masses at phialides on the
respective branches, with stipes and terminal vesicles characteristic of most
of Cylindrocladium species. Vesicles globose to subglobose, often constricted
or furrowed. Conidia phialosporous, hyaline, cylindrical, 1 to 4-celled (mainly
4-celled). Chlamydospores globose, reddish brown, thick-walled, catenulate.
Sclerotia dark reddish brown, spherical or irregular, composed of catenulate
and aggregated chlamydospores.
Dimensions: Conidiophores mostly 140–330 µm tall, nearly 6–8 µm wide
basally, 3–4 µm wide apically: primary branches 15 µm long: secondary
B
branches 20 µm long: tertiary branches 12.5 µm long: phialides 7–19 × 2.4–4
µm: vesicles 14–20 × 6–14 µm. Conidia 32–48 × 3–4 µm. Chlamydospores
12.5–26 µm in diameter. Microsclerotia 65–115 µm or more in diameter.
A C
Material: 00-52 (= MAFF 238171, Forest soil, Chibusa-yama, Hahajima,
the Bonin Islands, Tokyo, Japan).
Remarks: This fungus resembles C. citri (Fawcett & Klotz) Boedijin & Reitsma morphologically except for
formation of chlamydospores and microsclerotia, although these structures were described and illustrated by
Crous and Wingfield (1994).
Cylindrocladium sp. A: Conidiophores with penicillate fertile portions, stipes and vesicles, and conidia. B:
Conidia germinated or not germinated. C: Catenulate chlamydospores forming microsclerotia. D,E: Part of
stipes and vesicles (E,F), penicillate fertile portion, and detached conidia (E). (From Watanabe, T. et al. 2001c.
Mycoscience. In press. With permission.)
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Cytospora sacchari. A–D: Pycnidia and spore mass on apical neck (arrow in D). E: Spore mass on apical
pycnidium. F,G: Conidiophores and conidia. H: Conidia. (From Watanabe, T. 1975b. Trans. Mycol. Soc. Jpn.,
16:28–35. With permission.)
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Dactylaria Sacc.
Michelia 2:20, 1880.
Type species: D. purpurella Sacc.
A
Dactylaria candidula (Hohn.) Bhatt & Kendrick
References: Bhatt and Kendrick 1968; Hoog 1985; Timms and Hepden 1965.
Morphology: Conidiophores hyaline, simple, erect, bearing 2–8 conidia sympo-
dially at sterigmata on apical fertile parts. Conidia sympodulosporous, hyaline, B
cylindrical, 2-celled, rarely 3- to 4-celled, with septum at the median.
Dimensions: Conidiophores up to 28 µm tall. Conidia 22.5–28.8 × 0.7–1.3 µm.
Material: 73-139, 73-268 (Strawberry root, Shizuoka, Japan).
Remarks: The conidia of this fungus are slightly narrower than those in the
original description. C-E
C B
Dactylella Grove
J. Bot. Lond. 22:195, 1884.
Type species: D. minuta Grove
Dactylella chichisimensis. A: Habit. B–D: Conidiophores and aleurioconidia. E: Chlamydospore and aleurio-
conidia. F: Sclerotium. (From Watanabe, T. et al. 2001c. Mycoscience. In press. With permission.)
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Dematophora Hartig
Unters. Forstbot. Inst. München 3:95, 1883.
A Type species: D. necatrix Hartig
B Teleomorph: Rosellinia necatrix (Hartig) Berl. : Prill.
Dematophora necatrix. A: Hyphae with pear-shaped cells. B: Sclerotia. C–E: Conidiophores and conidia.
F: Fertile portions of conidiophores and conidia. G: Germinated conidia (arrow) on conidiophores. H: Ger-
minated conidia. (From Watanabe, T. 1992b. Ann. Phytopathol. Soc. Jpn., 58:65–71. With permission.)
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Didymostilbe P. Henn.
Hedwigia 41:148, 902.
Type species: D. coffeae P. Henn.
Didymostilbe sp.
Diplodia Fr.
Summa Veg. Scand. 416, 1849.
Type species: D. mutila Fr.
C
Diplodia frumenti. A,D: Crushed pycnidia. B,E: Conidiophores and conidia. C,F: Conidia (A–C: Isolate
69-438; D–F: Isolate P9).
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B
Morphology: Conidiophores yellowish brown, short, not well
differentiated from ordinary hyphae. Conidia solitary or aggregated
forming sporodochia, brown, globose, rough on the surface, occa-
sionally with cylindrical hilum, muriform composed of transverse
and longitudinal septa.
C
Dimensions: Conidiophores up to 20 µm long. Conidia 15–20 µm
in diameter.
Material: 84-526 (Japanese red pine seed, Hiba, Hiroshima, Japan).
Fumago Pers.
Myc. Eur. 1:9, 1822.
Lectotype: F. vagans Pers.
Fumago sp.
A, B
References: Friend 1965; Hughes 1976, 1983; Sahni 1964.
Morphology: Papulospore-like spores borne intercalarly on
ordinary hyphae or terminally on short conidiophores, reddish
brown or dark brown, subglobose or irregular, muriform when
immature, composed of several component cells. Hyphae often
rich in oil globules.
Dimensions: Muriform spores mostly 14–28 µm in diameter.
Sporephore 4–32 × 2–3.6 µm. Hyphae 3–4.4 µm wide.
Material: 00-76 (Uncultivated soil, Chichijima, the Bonin
C D
Islands, Tokyo, Japan).
Remarks: Colonies on PDA are dark gray, velvety, homogeneous. The genus Fumago is selected to accom-
modate a member of soil fungi only with hyphae and dark brown muriform spores simply or in chains, although
its validity is often controversial. This fungus may resemble Sarcinella Sacc. with hyphopodiate hyphae.
However, Fumago and Sarcinella are commonly known as members of sooty molds with a few other
hyphopodiate fungi. Fumago is also listed as a member of soil fungi (Ranzoni, 1968).
Fumago sp. A,B: Muriform spores and hyphae. C,D: Intercalary muriform spores. E,F: Both apical and
intercalary muriform spores.
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Key to Species
Material: 00-54 (Forest soil, Mt. Chibusa, Hahajima, the Bonin Islands, Tokyo, Japan).
Remarks: Colonies on PDA are homogeneous, yellowish brown centrally, white marginally, resupinate.
Fusarium ventricosum. A: Conidiophore and microconidia. B: Conidiophore with elongated branches and
microconidia. C: Macroconidium and microconidia. Note a microconidium bearing chlamydospore. D: Macro-
conidia and chlamydospores.
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Geotrichum candidum. A: Hypha and chlamydospores. B–E: Conidia and sporulation (B,D,E). (From
Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Gliocladium Corda
Icones Fung. 4:30, 1840.
Type species: G. penicillioides Corda
Key to Species
1. Conidiophores penicillate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
penicillate and verticillate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
C D
Gliocladium catenulatum. A: Conidiophores with verticillate branches and spore masses. B: Conidiophore
with penicilliate branches, and catenulate conidia. C: Spore mass formed on verticillate phialides. D: Conidia.
(From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Gliocladium roseum. A: Conidiophores and spore masses. B: Conidiophores and conidia. C: Part of the
conidiophores and conidia. (From Watanabe, T. 1975c. Trans. Mycol. Soc. Jpn., 16:149–182. With permission.)
1118-ch06B-Frame Page 282 Sunday, February 24, 2002 7:32 PM
Gliocladium virens. A: Conidiophores and spore masses. B: Conidia and chlamydospores. C,D: Conidio-
phores with phialides and conidia. (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267.
With permission.)
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Gliocladium viride. A,B: Conidiophores and spore masses. C,D: Conidiophores with fertile portions and conidia.
(From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 285 Sunday, February 24, 2002 7:32 PM
Gloeocercospora sorghi. A: Sporodochia and sclerotia formed on natural medium containing sorghum straw.
B: Sporodochium. C: Conidia. D: Sclerotia. E: Germinated conidia. F: Sporodochium on sclerotium. (From
Watanabe, T. and Hashimoto, K. 1978. Ann. Phytopathol. Soc. Jpn., 44:633–640. With permission.)
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Gonatobotrys Corda
Prachtflora 9, 1839.
Type species: G. simplex Corda
Gonatobotrys sp.
Gonatobotrys sp. A–C: Conidiophores and spore masses. D: A portion of the apical conidiophore and
macroconidia. E: Conidiophores and microconidia.
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Gonytrichum macrocladum. A: Apical sterile branches formed on natural substrate. B,D: Conidiophores and
spore masses. C: Overview of phialides and conidia. E: Conidiophore with phialides and conidia. F: Rhizoids.
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Hansfordia Hughes
Mycol. Pap. 43:15–24, 1951.
Type species: H. ovalispora Hughes
Dimensions: Conidiophores 52.5–250 × 1.2–3 µm. Conidia (3.9-) 5–8 × 1.9–3 µm.
Material: 69-424 (Pineapple leaf near soil, Nakijin, Okinawa, Japan).
Remarks: This fungus was isolated from leaf partially embedded in soil, and treated as a member of soil
fungi. Cultures on PDA are pale brown with blackish fragments distributed all over. Dicyma Boulanger may
be a synonym of this genus (Arx, 1981).
Helicomyces Link
Linn. Spec. Pl. 1:131, 1824.
Type species: H. roseus Link
A-D
References: Goos 1985; Moore 1955, 1957.
Morphology: Conidiophores not differentiated from ordinary
hyphae (micronematous), short, simple, hyaline to subhyaline,
bearing single conidia laterally on hyphae. Conidia sympodulo-
sporous, hyaline to subhyaline, white in mass, tightly coiled in
1 plane mostly 3 times, conidial filaments elongated when wet
(hygroscopic), numerously septate, often with over 28 septations.
Dimensions: Conidia 30–35 µm in diameter: conidial filament
3–3.6 µm wide.
Helicomyces roseus. A–D: Conidia and hyphae with short conidiophores. E: Elongated conidium after
mounting in water.
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Humicola Traaen
Nyt Mag. Naturvid. 32:20, 1914.
Type species: H. fuscoatra Traaen
Key to Species
Humicola fuscoatra. A,B: Hyphae and aleurioconidia. C–E: Sporulation of phialoconidia and aleurio-
conidia. F: Aleurioconidia and phialoconidia. (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267.
With permission.)
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Remarks: This fungus is morphologically close to H. fuscoatra, but different in forming more pigmented
darker brown, and larger aleurioconidia than the latter. Phialoconidia are rarely formed, and often neglected
as a morphological criterion, but they are broadly ovate.
Humicola sp.
References: Barron 1968; Domsch et al. 1980a,b.
Morphology: Conidia of two kinds; aleurio-
conidia single or in twos, or rarely catenulate, in
A a short chain, globose or spindle-shaped, verru-
cose, pale brown, terminal or intercalary; phialo-
conidia aggregated in a spore mass, borne directly
on hyphae or on aleurioconidia, hyaline, ovate,
ellipsoidal, cylindrical or barrel-shaped, occasion-
ally apiculate at one end.
Dimensions: Aleurioconidia globose 5–6.5 (-7.5) µm
in diameter; spindle-shaped, 6.5–8.8 × 4.5–6.8 µm.
Phialoconidia 2–2.3 (-3.5) × (1.2-) 2.3–2.5 µm.
Phialide, mostly 3.8–12.5 µm long.
Materials: 76-55 (Gentian root, Chino, Nagano,
Japan).
C-D E Remarks: This fungus is characterized in form-
ing small aleurioconidia and phialoconidia in
spore masses.
Humicola sp. A: Aleurioconidia and hyphae. B: Aleurioconidia and a mass of phialoconidia formed on
hypha. C,D: Phialides and phialoconidia formed on aleurioconidia, and aleurioconidia. E: Phialoconidia.
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G-I
Hyalodendron Diddens
Zentbl. Bakt. Parasitkde., Abt. 2, 90:315, 1934.
Type species: H. lignicola Diddens
Hyalodendron sp.
Hyphodiscosia radicicola. A: PDA colony. B,C: Sporodochia and conidia. D: Conidia. E–G: Conidiophores
and conidia. (From Watanabe, T. 1992a. Mycologia, 84:113–116. With permission.)
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Macrophomina Petrak
Annl. Mycol. 21:314, 1923.
Type species: M. philippinensis Petrak = Tiarosporella phaseoli
(Maubl.) van der Aa = Macrophoma phaseolina Tassi B, C
Macrophomoina phaseolina. A: Hypha. B,C: Sclerotia on agar culture. D: Sclerotia on bean straw in natural
media. E: Pycnidia. F: Crushed pycnidia and conidia. G: Conidiophores and conidia. H: Conidia. (From
Watanabe, T. 1972a. Ann. Phytopathol. Soc. Jpn., 38:106–110. With permission.)
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Mammaria Ces.
Flora 12:207, 1854.
Type species: M. echinobotryoides Ces.
Mammaria sp. A, E
Mammaria sp. A: Spore masses of phialoconidia and aleurioconidia. B: Phialoconidia and aleurioconidium.
C: Aleurioconidia. D,E: Phialoconidia and conidiophores.
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Metarhizium Sorokin
Les Maladies des Plantes, etc. 2:268, 1883.
C
Type species: M. anisopliae (Metschn.) Sorokin
Metarhizium anisopliae. A: Sporodochia. B,C: Conidiophores, phialides, and conidia. (From Watanabe, T.
1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 307 Sunday, February 24, 2002 7:32 PM
Microsphaeropsis Höhn.
Hedwigia 59:267, 1917.
Type species: M. olivacea (Bonord.) Höhn.
Synonym: Coniothyrium olivaceum Bonord.
Microsphaeropsis sp.
A B, C
References: Sutton 1980.
Morphology: Pycnidia globose, white when immature, pig-
mented gradually with age, covered with pigmented hairs, brown
when mature, with black spore masses extruded from indistinct
apical ostioles: peridium yellowish brown, pseudoparenchyma-
tous. Conidia globose, smooth, hyaline or slightly pigmented D
initially, brown, 6- to 7-angled, subglobose, rough at the margin,
1-celled, thick-walled when mature.
Dimensions: Pycnidia ca. 250 µm. Conidia 7.5–8 µm in diameter.
Material: 69-313 (Pineapple field soil, Ishigaki, Okinawa, Japan).
Remarks: Cultures on PDA are pale brown with black spots distributed, reverse pale reddish brown.
Sporogenous cells not observed. However, its angular globose conidia are characteristic morphologically.
This fungus is different from any of the 14 genera of Sphaeropsidales with globose conidia recorded by
Sutton (1980). The genus Microsphaeropsis is introduced conveniently by Sutton (1977), but it is close to
the genus Coniothyrium morphologically.
Microsphaeropsis sp. A: Pycnidium. B: Spore masses extruded from pycnidia. C,D: Peridium and conidia.
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Monacrosporium Oudem.
Ned. Kruidk. Arch. 2, 4:250, 1885.
Type species: M. elegans Oudem.
Key to Species
Monilia pruinosa. A–D: Hyphae and monilioid cells. E: Detached monilioid cells. (From Watanabe, T.
1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Monilia sp.
References: Domsch and Gams 1972; Gilman 1957.
Morphology: Conidiophores lacking or indistinct, not well differentiated
from hyphae. Hyphae hyaline, nonaerial, creeping, bearing catenulate
conidia developed by budding directly from hyphae: conidial chains
simple or branched. Conidia hyaline, blastosporous, 1-celled, ellipsoidal.
Dimensions: Conidia 8.7–12.5 × 5.5–8 µm.
Material: 77-302 (Uncultivated field soil, Nishigahara, Tokyo, Japan).
Remarks: This fungus is abundant in autoclaved field soil.
Mycocentrospora Deighton
Taxon 21:716, 1972.
Type species: M. acerina (Hartig) Deighton
Mycocentrospora acerina. A,B: Conidiophores and conidia. C: Chlamydospores. D,E: Conidiophores and
part of conidia.
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Mycoleptodiscus Ostazeski
Mycologia 59:970, 1967.
Type species: M. terrestris (Gerd.) Ostazeski
Synonym: Leptodiscus terrestris Gerdemann
F
C, D
Mycoleptodiscus terrestris (Gerd.) Ostazeski
Mycoleptodiscus terrestris. A,B: Young (A) and mature (B) sporodochia. C: Stroma with extruded phialide.
D: Young conidia attached to the sporogenous cells by short stalks. E,F: Conidia. Note 1-septate (E,F) and
aseptate conidia (F). G,H: Germination of conidia on wet slides 5 days after treatment at 25°C. Note formation
of appressoria on elongated germ tubes (H).
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Myrioconium Syd.
Ann. Mycol. 10:449, 1912.
Type species: M. scirpicola (Ferd. & Winge) Ferd. & Winge
Myrioconium sp.
Myrioconium sp. A–C: Conidiophores and conidia. D: Conidia and phialides. E: Sclerotium.
1118-ch06B-Frame Page 318 Sunday, February 24, 2002 7:32 PM
Key to Species
Myrothecium cinctum. A: Sporodochia. B: Conidiophores and conidia. (From Watanabe, T. 1975d. Trans.
Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 320 Sunday, February 24, 2002 7:32 PM
F G
Myrothecium dimorphum. A: Sporodochia with erect seta-like conidiophores. B,C: Apical fertile portions
of seta-like conidiophores bearing microconidia, macroconidia, and phialides (C). D: Seta-like conidiophore,
setae, and conidia. E: Phialides and macroconidia detached. (From Watanabe, T. 1975d. Trans. Mycol. Soc.
Jpn., 16:264–267. With permission.)
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Naranus T. Watanabe
Mycol. Res. Trans. 99:806, 1995.
Naranus cryptomeriae. A,B: Initiation of hyphal swellings. C,D: Catenulate and aggregated conidia.
E: Detached conidia. (From Watanabe, T. 1995. Mycol. Res., 99:806–808. With permission.)
1118-ch06B-Frame Page 323 Sunday, February 24, 2002 7:32 PM
Neta quadriguttata. A: Habit showing aleurioconidia. B: Aleurioconidia and one allantoid hyaline conidium.
C–E: Aleurioconidia and allantoid conidia formed on the identical hyphae. F: Mass of allantoid conidia.
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Nigrospora Zimmerm.
Zentbl. Bakt. Parasitkde. Abt. 2, 8:220, 1902.
Type species: N. panici Zimmerm.
Nigrospora oryzae. Conidiophore and conidium. (From Watanabe, T. and Tsudome, K. 1970. Trans. Mycol.
Soc. Jpn., 11:64–71. With permission.)
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Nigrospora sacchari. A,B: Conidiophores and conidia (A: Isolate 84-509; B: 70-1841).
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Nodulisporium Preuss
Herb. Viv. Mycol. no. 1272, 1849.
Type species: N. ochraceum Preuss.
Nodulisporium melonis. A–F: Conidiophores and conidia. G: Conidia. H–K: Apical fertile portions
of conidiophores. (From Watanabe, T. and Sato, M. 1995. Ann. Phytopathol. Soc. Jpn., 61:330–333.
With permission.)
1118-ch06B-Frame Page 328 Sunday, February 24, 2002 7:32 PM
Oedocephalum Preuss
Linnaea 24:131, 1851.
Type species: O. elegans Preuss
E
Oedocephalum nayoroense T. Watanabe
B
References: Baksi 1950, 1951; Dodge 1937; Hughes 1953a; Stalpers
1974; Thaxter 1891; Watanabe 1991.
Morphology: Conidiophores erect, simple or rarely branched, clavate,
bearing 1–16 conidia at sterigmata borne on apical fertile portions, on
which appear basidia and basidiospores. Conidia yellowish brown or
brown, ellipsoidal, rough or minutely echinulate on the surface, 1-celled.
No clamp connection present on hyphae.
Dimensions: Conidiophores 92.5–195 × 12.5–20 µm. Conidia 11.5–15
× 6–7.5 µm.
Material: 81-498 (= IFO 32546, Potato field soil, Nayoro, Hokkaido,
Japan).
C D
Remarks: This fungus is also close to the genus Spiniger morphologically.
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Oedocephalum nayoroense. A–G: Conidiophores and conidia. (From Watanabe, T. 1991. Mycologia, 83:524–529.
With permission.)
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Oidiodendron Robak
Nyt. Mag. Naturvid. 71:243, 1932.
Type species: O. fuscum Robak = O. tenuissimum (Peck) Hughes
Key to Species
A, B
Paecilomyces Bain.
Bull. Soc. Mycol. Fr. 23:26, 1907.
Type species: P. variotii Bain.
Key to Species
5. Conidia of two kinds: subglobose and ovate, over 3.2 µm long . . . . . . . . P. variabilis
subglobose, under 3 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. victoriae
C E
Paecilomyces inflatus. A–C,F: Conidiophores and conidia. D: Conidia. E: Conidiophore. (From Watanabe, T.
1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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A-C
A C
Paecilomyces puntoni. A,B: Conidiophores and conidia. C: Basal parts of conidiophores. D: Conidia.
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B, C
Paecilomyces variabilis. A–D: Conidiophores and conidia. E: Conidia. (From Watanabe, T. 1975d. Trans.
Mycol. Soc. Jpn., 16:264–267. With permission.)
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Papulaspora Preuss
Linnaea 24:113, 1851.
Type species: P. sepedonioides Preuss
Key to Species
Papulaspora irregularis. A: Hyphae. B,C: Papulospores and component cells. (From Watanabe, T. 1975d.
Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Papulaspora nishigaharanus. A,B: Papulospores and component cells (B-1). C: Papulospore and catenulate
conidia. D: Conidiophores and conidia. E–K: Process of papulospore formation. (From Watanabe, T. 1991.
Mycologia, 83:524–529. With permission.)
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Papulaspora pallidula. A–C: Process of papulospore formation. D: Papulospore. (From Watanabe, T. 1975d.
Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Papulaspora pannosa. A–C: Papulospores formed on aerial hyphae (A), and on agar medium (B,C). (From
Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Papulaspora sp. 1.
References: Hotson 1912, 1917, 1942; Weresub and LeClair 1971.
Morphology: Spores dimorphic, i.e., papulosporous, and
phialosporous. Papulospores borne intercalarly on ordinary
hyphae, or terminally on short conidiophores undifferentiated A
from hyphae, pale brown or brown, subglobose composed of
globose component cells. Conidiophores phialidic, hyaline,
erect, gradually tapering toward apex or acutely narrowed in
the middle, 1-septate. Conidia phialosporus, hyaline, ovate, api-
culate at one end.
B
Dimensions: Papulospores 15–25 (-30.5) µm in diameter.
Phialides (7.5-) 10–17.5 (-22.5) × 2–2.5 µm. Conidia 2.2–2.5
× 0.7–1 µm.
Material: 73-1071 (Uncultivated soil, Nishigahara, Tokyo,
Japan), 74-535; -615; -675 (Strawberry root, Shizuoka, Japan).
Remarks: Colonies on PDA are yellowish brown, with C, D
irregular concentric zonation. Papulospores of this fungus were
always formed on agar cultures, but conidia not always formed.
Papulaspora sp. 1. A,B: Papulospores and hyphae. C: Phialides and conidia. D: Conidia.
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Papulaspora sp. 2.
References: Hotson 1912, 1917, 1942; Weresub
and LeClair 1971.
Morphology: Spores dimorphic, i.e., papu-
losporous, and arthrosporous. Papulospores
borne directly on ordinary hyphae, or terminally
on short conidiophores undifferentiated from
A, B, D
hyphae, embedded, yellowish brown or brown,
fleshy, subglobose composed of globose com-
ponent cells, smooth marginally. Arthrospores
hyaline, erect, borne directly on hyphae, aerial.
Conidia arthrosporous, hyaline, cylindrical.
Dimensions: Papulospores 100–225 µm in
diameter: component cells 11.2–17.5 µm in
diameter. Conidia 11.2–27.5 × 5–7 µm.
C Material: 73-467 (Uncultivated soil, Nishiga-
hara, Tokyo, Japan); 83-415 (Strawberry root,
Shizuoka, Japan).
Remarks: Colonies on PDA are dark yellow-
ish green.
Papulaspora sp. 2. A,B,D: Arthrospores in chains (A,B) and detached (B). C: Papulaspore.
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Penicillium Link
Mag. Ges. Natur. Freunde, Berlin, 3:16, 1809.
Type species: P. expansum Link
Key to Species
Penicillium corylophilum. A–C: Conidiophores, phialides, and conidia. (From Watanabe, T. 1975d. Trans.
Mycol. Soc. Jpn., 16:264–267. With permission.)
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Penicillium lanosum. A: Conidiophores and conidia. B: Conidia. C,D: Phialides and conidia. (From
Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 355 Sunday, February 24, 2002 7:32 PM
Penicillium nigricans. A: Conidiophores and conidia. B: Phialides and conidia. (From Watanabe, T. 1975d.
Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Penicillium resticulosum. A: Conidiophores and conidia. B,C: Phialides and conidia. (From Watanabe, T.
1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Periconia Tode
Syn. Fung. Carol. Sup. 125, 1822.
Type species: P. lichenoides Tode : Mérat
Key to Species
A B
Periconia byssoides. A,B: Conidiophores and conidia. (From Watanabe, T. and Sato, Y. 1988. Trans. Mycol.
Soc. Jpn., 29:143–150. With permission.)
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Periconia macrospinosa. A,B: Conidiophores and conidia. C,D: Apical fertile part of conidiophore and conidia.
(From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 360 Sunday, February 24, 2002 7:32 PM
A B
Pestalotia de Not.
Mem. R. Acad. Sci. Torino 2, 3:80, 1839.
Type species: P. pezizoides de Not.
Pestalotia spp.
H D
Synonym: Pestalotiopsis spp.
References: Dube and Bilgrami 1966; Guba 1932, 1961; Steyaert 1949,
1955, 1956; Sutton 1969; Watanabe et al. 1986a.
Morphology: Spodochia (spore masses) on agar cultures hemispherical,
mucilagenous, black, occasionally leaked around sporodochia. Conidio-
phores short, simple. Conidia spindle-shaped or ellipsoidal, 4- to 5-celled,
with 2–3 central, pigmented cells (especially darker in 2 cells), with B F
2–4 appendages (setulae) in apical cells and 1 short appendage (pedicel)
in basal cells.
Dimensions: Conidia (excluding apical and basal appendages) 19.7–50
× 5.0–10 µm: pigmented cells (7.5-) 10–20 µm long: apical appendages
(4.7-) 10–30 µm long: basal appendages 1.2–12.5 µm long.
Material: 69-319 (Pineapple field soil, Okinawa, Japan); 70-1752 A E
(Pineapple field soil, Hachijo, Tokyo, Japan); 83-1, 83-2, 83-5 (Japanese
red pine seed, Hiba, Hiroshima, Japan); 83-3 (Japanese black pine seed, Kumano, Mie, Japan); 83-4 (Japanese
cedar seed, Yoshino, Nara, Japan); 85-47, 85-48 (Flowering cherry seed, Hachioji, Tokyo, Japan).
Remarks: On the basis of the number of component cells of conidia, the genus Pestalotia may be further
split into Pestalotia (6 cells), Pestalotiopsis Steyaert. (5 cells), and Truncatella Steyaert (4 cells).
In agar cultures, no acervuli are generally formed and thus, it is rather difficult to identify unknowns by
comparing the morphology of known species. Among Pestalotia isolates with almost similar conidial dimen-
sions obtained from Japanese pine seeds, conidia are formed with central larger and more darkly pigmented
cells in the isolate 83-2 (A), with short setulae (B) in the 83-1, with thick and black central septum (C) in
the 85-5, and with constriction at the septum in the 83-3 (D). Isolate 69-319 formed conidia with predominantly
2 setulae (G).
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Pestalotia spp. Conidia (A: 83-2, B: 83-1, C: 83-5, D: 83-3, E: 85-88, F: 85-49, G: 69-319).
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Pestalotia sp.
Synonym: Pestalotiopsis sp.
References: Dube and Bilgrami 1966; Guba 1932, 1961;
Steyaert 1949, 1955, 1956; Sutton 1969.
Morphology: Sporodochia (spore masses) on agar cultures
hemispherical, mucilagenous, black. Conidiophores indis-
tinct. Conidia ellipsoidal, 4- to 5-celled, with 3 central,
pigmented cells, with 3–6 (mainly 4) appendages (setulae)
in apical cells and 1 short appendage (pedicel) in basal cells
that are often lacking. Chlamydospores globose, thick-
walled, borne in one or two cells of conidia with age. A
Pestalotia sp. A,B: Conidia. C,D: Chlamydospores and conidia. E–G: Germination of conidia.
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Phialophora Medlar
Mycologia 7:200, 1915.
Type species: P. verrucosa Medlar
Key to Species
8. Phialides aggregated, with indistinct collarette, conidia mainly cylindrical. . . .P. cinerescens
not aggregated, with conspicuous collarette, conidia mainly ovate . . . P. fastigiata
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A, B
Remarks: Short and thick phialides are characteristic of this fungus.
Phialophora cyclaminis. A: Conidiophores (phialides) and spore masses. B: Conidia. C,D,F: Phialides.
E,G: Chlamydospores (A–E: Isolate 72-X142; F,G: 85-P32). (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn.,
16:264–267. With permission.)
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Phialophora malorum. A: Conidiophores and spore masses. B: Conidiophores. C: Phialides and conidia.
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Remarks: Colonies on agar cultures are dark gray to black. This fungus is morphologically close to
Gaeumannomyces graminis (Sacc.) v. Arx & Olivier. Comparative studies of both fungi were conducted by
Decon (1973, 1974).
Phialophora richardsiae. A,B: Conidiophores, phialides, and spore masses. C: Ovate and globose conidia.
(From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
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Phialophora sp. 1.
References: Cole and Kendrick 1973; Schol-Schwarz 1970.
Morphology: Conidiophores (phialides) pale brown, erect,
simple or branched, developed often on aerial hyahae, bearing
catenulate conidia (often over 165 µm long) or spore masses:
phialides ampulliform (flask-shaped) or cylindrical with conspic-
uous collarette. Conidia hyaline, subglobose, truncate at one end.
A, C E
Dimensions: Conidiophores 7–11.3 × 3–4.8 µm: collarettes
1.2–2.3 µm wide. Conidia 1.2–2.0 µm in diameter.
Material: 73-410 (Strawberry root, Tsu, Mie, Japan).
Remarks: This fungus is morphologically close to P. verrucosa
Medlar, in forming flask-shaped phialides with conspicuous col-
larettes, but their conidia are globose with truncate base, and B D
ellipsoidal, respectively.
Phialophora sp. 1. A–C: Conidiophores and conidia in chains or spore masses. D,E: Phialides and spore masses.
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Phialophora sp. 2.
References: Cole and Kendrick 1973; Schol-Schwarz 1970.
Morphology: Conidiophores (phialides) pale brown, erect, simple
or branched, bearing spore masses apically: phialides tapering from
A base toward apex with well-developed collarette. Conidia hyaline,
B subglobose, apiculate at one end. Sclerotia brown.
Dimensions: Phialides 12.5–30 × 2–3.8 µm: collarettes ca. 2 µm
wide, 0.7 µm deep. Conidia 1.7–2.5 µm in diameter. Sclerotia
30–75 µm in diameter.
Material: 74-750 (Strawberry root, Mie, Japan).
Remarks: This fungus is morphologically close to P. cyclaminis
except for sclerotium formation.
C D
Phoma Sacc.
Michelia 2:4, 1880.
Type species: P. herbarum Westend
Key to Species
Remarks: Among over 2000 species were described in Phoma, nearly 40 are valid (Hawksworth et al., 1995).
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A
Morphology: Pycnidia globose or subglobose, dark brown, conspicuously
ostiolate: peridium dark brown, pseudoparenchymatous. Conidia hyaline,
ellipsoidal, 1-celled. Chlamydospores dark brown, subglobose, muriform with
transverse and longitudinal septa rather irregularly.
Dimensions: Pycnidia 50–150 µm: ostioles 7–8 µm wide. Conidia 4.5–5.3 ×
1.7–2.5 µm. Chlamydospores 15–32.5 (-42.5) × (10-) 13.8–16.8 (-20) µm.
Materials: 77-160 (Strawberry root, Shizuoka, Japan); 83-17 (Japanese red pine
seed, Hiba, Hiroshima, Japan).
C, D
Remarks: Conidiophores were indistinct. This fungus is well characterized by
pycnidia, and chlamydospores that resemble conidia of genus Alternaria.
Phoma glomerata. A: Part of the pycnidium and chlamydospore. B: Chlamydospores. C,D: Conidia and
chlamydospores.
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Phoma sp.
References: Dennis 1946; Dorenbosch 1970; Sutton 1964, 1980;
Watanabe 1975b.
Morphology: Pycnidia globose, subglobose, or disc-shaped,
A, B conspicuously ostiolate: peridium dark brown, pseudoparenchy-
matous. Conidia hyaline, ellipsoidal, 1-celled. Chlamydospores
solitary, dark brown, granulate, thick-walled.
Dimensions: Pycnidia 107–146 × 97.3–107 µm: ostioles 4.8–5 µm
in diameter. Conidia 4.8–6.1 × 2.1–2.7 µm. Chlamydospores
10.2–18.3 µm in diameter.
Materials: 72-X24 (Sugarcane root, Taiwan, ROC).
D E
Remarks: Conidiophores were indistinct.
Phoma sp. A,B: Pycnidia. C: Pycnidial ostiole. D: Conidia. E,F: Chlamydospores. (From Watanabe, T.
1975b. Trans. Mycol. Soc. Jpn., 16:28–35. With permission.)
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References: Hahn 1930; Hobbs et al. 1985; Sutton 1980; Watanabe et al. 1986a.
Morphology: Pycnidia half-embedded in agar media, solitary or aggregated,
globose, irregular with indistinct ostioles: peridium dark brown, pseudo-
parenchymatous. Conidia hyaline, 1-celled, dimorphic: the alpha type, often B C
borne in pale pink spore masses, spindle-shaped or ellipsoidal; and the beta
type, borne in white spore masses, filiform, curved.
Dimensions: Pycnidia ca. 450 × 800 µm. Conidiophores ca. 25 × 1.2 µm. Conidia: alpha type (5.7-) 8.4–12.5 ×
1.8–2.5 µm, and beta type 16.5–28 × 0.5–1.3 (-1.6) µm.
Materials: 83-9 (Japanese black pine seed, Kumano, Mie, Japan); 83-33 (Japanese cedar seed, Yoshino, Nara,
Japan); 84-517, 84-518 (Japanese red pine seed, Saihaku, Tottori, Japan).
Remarks: Nearly 100 species are distributed widely (Hawksworth et al., 1996). The two conidial types are
variously produced in frequency by the isolates studied.
Pyrenochaeta De Not.
Micromycetes Ital. 5:15, 1845.
Type species: P. nobilis De Not.
Key to Species
Pyrenochaeta gentianicola. A,B: Pycnidia. C,D: Pycnidia and setae. E: Hyphae with rich oil globules. F:
Conidiophores and conidia. G: Sclerotia. H: Conidia.
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Pyrenochaeta globosa. A–F: Pycnidia. E: Part of peridium and setae. F: Conidiophores (phialides) on
peridium and conidia. G–I: Part of pycnidia and setae. J: Conidia. (From Watanabe, T. 1992c. Trans. Mycol.
Soc. Jpn., 33:21–24. With permission.)
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Pyrenochaeta terrestris. A: Pycnidium crushed. B: Setae around ostiolar region. C: Conidiophores. D: Conidia.
(From Watanabe, T. 1975b. Trans. Mycol. Soc. Jpn., 16:28–35. With permission.)
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Ramichloridium anceps. A,B: Conidiophores and conidia. (From Watanabe, T. et al. 1987a. Trans. Mycol.
Soc. Jpn., 28:453–469. With permission.)
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Ramichloridium subulatum. A–C: Conidiophores and conidia. (From Watanabe, T. and Sato, Y. 1988. Trans.
Mycol. Soc. Jpn., 29:143–150. With permission.)
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Rhizoctonia spp.
References: Parmeter (Ed.) 1970; Sneh et al. 1991.
Morphology: Conidia not formed. Hyphae pale brown,
branched angularly with side branches septated closely near the
main hyphae, and constricted basally. Monilioid cells usually
formed. Sclerotia discrete or aggregated, pale brown to brown,
various in shape and size.
B
Dimensions: Hyphae 6–10 µm wide. Sclerotia 1 mm long.
Material: 85-60, 85-61, 85-62, 85-63, 85-64, 85-65 (Flowering
cherry seed, Hachioji, Tokyo, Japan).
Remarks: Cultures of Rhizoctonia spp. on PDA are separated
into at least 9 colony types for the cherry seed isolates, showing
6 types among them in Figure A. The sclerotia of Isolate 85-62
are characteristically fragment-like. There are over 20 anasto-
C D mosis groups in binucleate Rhizoctonia.
Rhizoctonia spp. obtained from flowering cherry seed. A: Colonies (from upper left to lower right: Isolates
85-60, -61, -62, -63, -64, -65). B: Hypha (Isolate 85-65). C: Monilioid cells. D. Sclerotia (Isolate 85–62).
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Robillarda Sacc.
Michelia 2:8, 1880.
Type species: R. sessilis Sacc.
Sarcopodium Ehrenb.
Sylv. Myc. Berol. pp. 12, 23, 1818.
Type species: S. circinatum Ehrenb.
Sarcopodium araliae. A,B: Fruiting structures. C: Conidia. D: Fruiting structure and Gliocladium-like sporula-
tion on agar culture. E: Fruiting structure after removal of spore mass. Note setae and rhizoid. F: Palisade
layer of conidiophores. G: Conidiophores with phialides. (From Watanabe, T. 1993b. Mycologia, 85:520–526.
With permission.)
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Sclerotium spp. A,B,G: Hyphae with (G) and without clamp connections (A,B). C,D,F: Sclerotia formed in
agar (C,D) and on aerial hyphae (F). E: Colony (A,B: 72-X110; C,D: 73-226; E–G: 85-15). (From Watanabe, T.
1987b. Trans. Mycol. Soc. Jpn., 28:475–481. With permission.)
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Scolecobasidium Abbott
Mycologia 19:29, 1927.
Type species: S. terreum Abbott.
Scopulariopsis Bainier
Bull. Soc. Mycol. Fr. 23:98, 1907.
Type species: S. brevicaulis (Sacc.) Bainier.
Key to Species
D E
Scopulariopsis asperula. A,B,D: Conidiophores and conidia. C,E: Phialides and conidia.
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Selenophoma Maire
Bull. Soc. Bot. Fr. 53:87, 1906.
Type species: S. catananches Maire
D
Selenophoma obtusa Sprague et Johnson
Selenophoma obtusa. A: Pycnidium with extruded spore mass. B,C: Conidiophores and conidia. D: Conidia.
(From Watanabe, T. 1975b. Trans. Mycol. Soc. Jpn., 16:28–35. With permission.)
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Sepedonium chrysospermum. A–C: Conidiophores and conidia formed on agar (A,C) and on aerial hyphae
(B). D,E: Conidia.
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Sepedonium sp.
References: Damon 1952; Watanabe et al. 1987a.
Morphology: Spores dimorphic, i.e., aleuriosporous and sympodu-
A losporous. Conidiophores hyaline, erect, simple or branched, mainly
simple for aleuriospores, but branched oppositely or verticillately for
sympodulospores. Aleurioconidia pale brown, globose, double mem-
branaceous, occasionally deciduous, often pedicellate, occasionally
with apically pointed setae. Sympoduloconidia hyaline, ellipsoidal,
ovate, 1-celled.
Dimensions: Branches for sympodulospores 17.5–37.5 µm tall. Aleurio-
conidia 42.5–37.5 µm in diameter. Sympoduloconidia 5–9 × 3.7–5.5 µm.
B C Material: 85-P106 (Paulownia root, Misiones, Argentina).
Remarks: None of at least 10 described species in this genus fits this fungus morphologically.
Septonema Corda
Icon. Fung. 1:9, 1837.
Type species: S. secedens Corda
Septonema chaetospira. Hypha and conidia. (From Watanabe, T. and Sato, Y. 1988. Trans. Mycol. Soc. Jpn.,
29:143–150. With permission.)
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Spegazzinia Sacc.
Michelia 2:37, 1880.
Type species: S. tessarthra (Berk. et Curt.) Sacc.
References: Damon 1953; Ellis 1971, 1976; Hughes 1953; Watanabe and
Sato 1988.
Morphology: Conidiophores brown, erect, various in height, bearing 1 to
several conidia apically. Conidia dark brown, at least of two kinds:
smooth, square- and disc-shaped, ellipsoidal, 2- to 4-celled; and muriform,
composed of cross- and longitudinal septa with setae, constricted at or
A C near septa.
Dimensions: Conidiophores 75–142.5 × 2.2–2.8 µm. Conidia (excluding protuberances) 12.5–16.3 µm in
diameter, ca. 7.5 µm thick: protuberances 5–8.8 µm long.
Material: 86-59 (Japanese cypress seed, Kasama, Ibaraki, Japan).
Remarks: Process of sporulation may be interesting to study.
Spegazzinia tessarthra. A: Conidia and sporulation. B: Conidiophore and conidia. C: Conidia. (From
Watanabe, T. and Sato, Y. 1988. Trans. Mycol. Soc. Jpn., 29:143–150. With permission.)
1118-ch06B-Frame Page 407 Sunday, February 24, 2002 7:32 PM
Sporidesmium filiferum. A–I: Conidiophores bearing conidia. J: Detached conidia. Note the truncate bases
and apical appendages of conidia. (From Watanabe, T. 1996. Mycoscience, 37:367–369. With permission.)
1118-ch06B-Frame Page 410 Sunday, February 24, 2002 7:32 PM
Sporobolomyces sp.
A
References: Barnett and Hunter 1987; Watanabe et al. 1986a.
Morphology: Conidiophores indistinct. Conidia borne singly on sterigmata
developed from hyphae or detached conidia, reproduced by budding, hyaline,
cylindrical, lunar- or sickle-shaped, 1-celled.
Dimensions: Hyphae 1.8–2.7 µm wide. Conidia 7.5–12.6 × 1.5–2.5 µm:
B sterigmata on detached conidia 2.1–2.2 µm long.
Material: 84-578 (Japanese black pine seed, Okawa, Kagawa, Japan).
Remarks: Cultures on PDA are yeast-like, yellowish white in color.
Sporoschisma saccardoi. A: Conidia in a long chain. B,D: Conidiophores, conidia, and capitate hyphae with
mucilaginous envelopes. C: Apical portion of conidiophore and conidia. E: Germination of conidium. F: Rhizoid.
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Sporotrichum aureum. A,B: Hyphae, conidiophores, and conidia. Note hyphae with clamp connections (arrow).
C: Conidia. (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 414 Sunday, February 24, 2002 7:32 PM
Sporotrichum sp.
References: Barron 1968.
Morphology: Conidiophores erect, simple or branched, bearing
single aleurioconidia apically on alternate or verticillate branches,
and catenulate arthroconidia on simple conidiophores. Conidia of
two kinds: aleuriosporous or sympodulosporous, hyaline, pale
brown, ellipsoidal, spindle-shaped, ovate or subglobose, truncate
or pedicellate basally; and arthrosporous, cylindrical, 1-celled.
Dimensions: Conidiophores 3–29 × 0.3–1 µm. Conidia: aleurio-
A, B
conidia (ellipsoidal) (3.5-) 5–8.5 × (1.5-) 3–5.5 µm, and arthro-
conidia 3–5 × up to 0.5 µm.
Material: 86–76 (Japanese black pine seed, Kyoto, Japan).
Remarks: Colonies on PDA are brownish gray, homogeneous,
C and floury, the reverse dark gray with slight radiation.
Stachybotrys Corda
Icon. Fung. 1:21, 1837.
Type species: S. chartarum (Ehrenb. : Link) Hughes
Stachybotrys bisbyi. A–C: Conidiophores and spore masses. D: Conidia. E: Conidiophore. (From Watanabe, T.
1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 416 Sunday, February 24, 2002 7:32 PM
Stagnospora subseriata. A–C: Pycnidia. D: Conidiophores. E,F: Conidia. (From Watanabe, T. 1975d. Trans.
Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 418 Sunday, February 24, 2002 7:32 PM
Stemphylium Wallr.
FI. Crypt. Germ. 2:300, 1833.
Type species: S. botryosum Wallr.
Taeniolella Hughes
Can. J. Bot. 36:816, 1958.
Type species: T. exilis (Karst.) Hughes
A, B
Taeniolella phialosperma T. Watanabe
Taeniolella phialosperma. A,B: Conidia and conidiophores in Taeniolella state. C: Taeniolella and Phialophora
state on the identical hypha. D,E: Phialophora state. F: Catenulate and branched conidia in Taeniolella state.
G: Aleurioconidia in Taeniolella state. (From Watanabe, T. 1992e. Mycologia, 84:794–798. With permission.)
1118-ch06B-Frame Page 421 Sunday, February 24, 2002 7:32 PM
Tetracladium de Wild.
Ann. Soc. Belg. Microsc. 17:35, 1899.
Type species: T. marchalianum de Wild.
Tetracladium setigerum. A: Conidiophore and spore mass. B,C: Conidiophores and conidia. D: Conidia.
(From Watanabe, T. 1975e. Trans. Mycol. Soc. Jpn., 16:348–350. With permission.)
1118-ch06B-Frame Page 422 Sunday, February 24, 2002 7:32 PM
Thielaviopsis Went
Meded. Proefst. West Java 7:4, 1893.
Type species: T. ethacetica Went = T. paradoxa (de Seynes) Hoehnel
Thielaviopsis paradoxa. A,B: Conidiophores and conidia. C,E: Conidia and chlamydospores. D: Catenulate
chlamydospores. (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:149–182. With permission.)
1118-ch06B-Frame Page 425 Sunday, February 24, 2002 7:32 PM
Thysanophora Kendrick
Can. J. Bot. 39:817, 1961.
Type species: T. penicillioides (Roum.) Kendrick
Torula herbarum. Conidiophores and conidia. (From Watanabe, T. and Sato, Y. 1988. Trans. Mycol. Soc.
Jpn., 29:143–150. With permission.)
1118-ch06B-Frame Page 427 Sunday, February 24, 2002 7:32 PM
Torula sp.
References: Schoknecht and Crane 1977; Watanabe et al. 1987.
Morphology: Conidiophores lacking. Conidia blastosporous, apically devel-
oped, forming simple or branched chains borne apically and laterally on hyphae,
often resulting in spore masses, hyaline, pale brown or brown, globose cylin-
drical, 1-celled. Chlamydospores dark brown, globose, double-membranous.
Dimensions: Conidia 6.2–8.3 × 2.5–3.8 µm. Chlamydospores 8.7–10 µm in A, B
diameter.
Material: 85-PP70 (Paulownia root, Itapua, Paraguay); 86-67 (Japanese cypress seed, Higashichikuma,
Nagano, Japan).
Torula sp. A–C: Conidiophores, conidia, and chlamydospores. (From Watanabe, T. et al. 1987a. Trans.
Mycol. Soc. Jpn., 28:453–469. With permission.)
1118-ch06B-Frame Page 428 Sunday, February 24, 2002 7:32 PM
Torulomyces Delitsch
Systematik der Schimmelpilze p. 91, 1943.
Type species: T. lagena Delitsch
A
Torulomyces lagena Delitsch
Torulomyces lagena. A: Habit. B: Conidiophores and conidia. C: Conidia. (From Watanabe, T. and Sato, Y.
1988. Trans. Mycol. Soc. Jpn., 29:143–150. With permission.)
1118-ch06B-Frame Page 429 Sunday, February 24, 2002 7:32 PM
Trichocladium Harz
Bull. Soc. Imp. Moscou 44:125, 1871.
Type species: T. asperum Harz
Trichocladium pyriformis. A,B: Hyphae, conidiophores, and conidia. (From Watanabe, T. 1991. Mycologia,
83:524–529. With permission.)
1118-ch06B-Frame Page 431 Sunday, February 24, 2002 7:32 PM
Key to Species
Trichoderma harzianum. A,B: Conidiophores and conidia. C,D: Conidiophores, phialides, and chlamydospores
(C) or conidia (D). (From Watanabe, T. 1975c. Trans. Mycol. Soc. Jpn., 16:149–182. With permission.)
1118-ch06B-Frame Page 435 Sunday, February 24, 2002 7:32 PM
Trichoderma koningi. A: Conidiophores and spore masses. B,C: Conidiophores and phialides. D: Conidia.
E: Chlamydospores. (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 436 Sunday, February 24, 2002 7:32 PM
Trichoderma pseudokoningi. A,B: Conidiophores and spore masses. C,G: Conidiophores and phialides.
D,H: Conidia. E,I: Chlamydospores and germination. (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn.,
16:264–267. With permission.)
1118-ch06B-Frame Page 437 Sunday, February 24, 2002 7:32 PM
Trichurus spiralis. A,B: Synnema and spore masses (C: top view). D: Part of the synnema and conidia.
1118-ch06B-Frame Page 439 Sunday, February 24, 2002 7:32 PM
Trinacrium Riess
Fres. Beitr. Mykol. 2: 42, 1852.
Type species: T. subtile Riess
Trinacrium iridis. A: Macroconidia. B,C: Branched macroconidia. D,F,G: Conidiophores elongated from
macroconidia and microconidia. E: Macroconidia and conidiophore. H: Chlamydospores. (From Watanabe, T.
1992g. Mycologia, 84:794–798. With permission.)
1118-ch06B-Frame Page 441 Sunday, February 24, 2002 7:32 PM
Tripospermum Speg.
Physis B. Aires 4:295, 1918.
Type species: T. acerinum (Syd.) Speg.
Tritirachium Limber
Mycologia 32:23, 1940.
Type species: T. dependens Limber
A Tritirachium sp.
References: Hoog 1972; Limber 1940; MacLeod 1954; Watanabe et al. 1986a.
Morphology: Conidiophores hyaline, simple or branched, bearing single
conidia on sterigmata or directly on verticillate or zigzag-shaped branchlets,
often elongating sterile, twisted, curved branchlets. Conidia sympodulosporous,
hyaline, ovate or globose, 1-celled.
Dimensions: Conidiophores 19.8–30.6 × 2.1–2.2 µm. Conidia 2.1–3.8 × 1.8–2.5
µm.
B
Material: 83-16 (Japanese black pine seed, Kumano, Mie, Japan).
Remarks: This fungus is morphologically close to Beauveria species.
Tritirachium sp. A,C: Conidiophores and conidia. B: Fertile portions and conidia.
1118-ch06B-Frame Page 443 Sunday, February 24, 2002 7:32 PM
Ulocladium Preuss
Dtschl. Flora, Pilze 3, 3:83, 1851.
Type species: U. botrytis Preuss
Vermispora sp.
A, B
References: Deighton and Pirozynski 1972; Rajashekhar
et al. 1991; Vasant & de Hoog. 1986.
Morphology: Conidiophores simple, erect, with terminal
single primary macroconidia, conidiogenous cells of sec- B, C
ondary microconidia sympodially branched often develop-
ing from macroconidia. Conidia hyaline, of two kinds: the
long-fusiform, 3- to 5-septate primary macroconidia; and
clavate, cylindrical, aseptate or occasionally 1-septate,
D, E
sharpened at one end, secondary microconidia.
Dimensions: Conidiophores mostly 120–132.5 × 3.7–4 µm. Conidia: primary macroconidia, 70–130 ×
12.5–13.8 µm, and secondary microconidia 27.5–36.3 × 5 µm.
Material: 74-632 (Strawberry root, Shizuoka, Japan).
Remarks: This fungus resembles V. cauveriana Rajashekar, Bhat et Kaveriappa morphologically, but the
macroconidia of the latter are 160–180 µm long, 6- to 9-septate, and the microconidia are truncate at one end.
Vermispora sp. A: Habit. B: Conidiophores and microconidia. C,D: Microconidia and conidiophores.
E: Macro- and microconidia.
1118-ch06B-Frame Page 446 Sunday, February 24, 2002 7:32 PM
Key to Species
Verticillium dahliae. A: Conidiophore and spore masses. B: Thick-walled resting cells. C: Conidia. (From
Watanabe, T. et al. 1973. Ann. Phytopathol. Soc. Jpn., 39:344–350. With permission.)
1118-ch06B-Frame Page 448 Sunday, February 24, 2002 7:32 PM
A D
Verticillium hahajimaense. A: Sporulation habit. B,C: Apexes of phialides and conidia. D: Chlamydospores
in chains. (From Watanabe, T. et al. 2001b. Mycoscience. In press. With permission.)
1118-ch06B-Frame Page 450 Sunday, February 24, 2002 7:32 PM
Verticillium nubilum. A: Conidiophores and spore masses. B: Conidia and solitary chlamydospores.
C: Catenulate chlamydospores.
1118-ch06B-Frame Page 452 Sunday, February 24, 2002 7:32 PM
Verticillium sphaerosporum. A,B: Sporulation on dead nematode. C: Conidia attached around the buccal
region of a nematode. D: Hyphae inside disintegrated body. E: Sporulation on the disintegrated nematode
body. F: Simple conidiophore bearing cylindrical conidia and spore mass composed of globose conidia.
G: Phialide and globose conidium. H: Verticillate conidiophore. I: Cylindrical and globose conidia. (From
Watanabe, T. 1980. Ann. Phytopathol. Soc. Jpn., 46:598–606. With permission.)
1118-ch06B-Frame Page 453 Sunday, February 24, 2002 7:32 PM
Verticillium sp.
References: Domsch et al. 1980a,b; Watanabe 1975d.
Morphology: Conidiophores erect, hyaline, bearing spore masses at verti-
cillate, alternate, opposite, or simple phialides: phialides tapering from base
toward apex. Conidia phialosporous, hyaline, cylindrical, 1-celled. Chlamydo-
A
spores pale brown, subglobose, minutely echinulate marginally.
Dimensions: Phialides 26.7–53.5 µm long. Conidia 3.6–4.9 × 1.2–2 µm.
Chlamydospores 5.6–7.8 µm in diameter.
Material: 72-X25 (Sugarcane root, Taiwan, ROC).
Remarks: Minutely echinulate globose chlamydospores are characteristic
for this fungus.
B D, E
Verticillium sp. A,D: Conidiophores and spore masses. B: Chlamydospores. C: Apical part of creeping hypha.
E: Conidiophores and conidia. (From Watanabe, T. 1975d. Trans. Mycol. Soc. Jpn., 16:264–267. With permission.)
1118-ch06B-Frame Page 454 Sunday, February 24, 2002 7:32 PM
Volutella ciliata. A: Sporodochia and conidia. B: Part of sporodochial tissue. C: Part of the setae. D: Conidia
in Acremonium or Verticillium state.
1118-ch06B-Frame Page 455 Sunday, February 24, 2002 7:32 PM
Wiesneriomyces Koorders
Verh. K. Akad. Wet. Amsterd. 2, 13(4):246, 1907.
Type species: Wiesneriomyces javanicus Koorders
Wiesneriomyces javanicus. A: Habit. B–D: Part of sporodochia showing fertile portions and setae. E: Conidia.
F: Sclerotia.
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Appendix
List of Living Cultures of Soil Fungi Deposited and Publicized
471
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Appendix 473
Appendix 475
Appendix 477
Note: The fungi described in this text are mostly deposited as the living stock cultures with MAFF, ATCC, IFO, or
CBS code number in addition to the list (TW) in this table. Consult the homepage for the MAFF list, at the e-mail
address: http://www.gene.affrc.go.jp./micro/T list/index.html.
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Index
479
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Index 481
Index 483
V W
Vermispora sp., 439, 445 Wiesneriomyces javanicus, 455–456
Verticillium albo-atrum, 447
Verticillium cauveriana, 445
Verticillium dahliae, 447
X
Verticillium fungicola, 448 Xylaria sp., 326
Verticillium hahajimaense, 449
Verticillium lecanii, 450
Verticillium malthousei, See Verticillium fungicola Z
Verticillium nubilum, 451
Zopfiella curvata, 174
Verticillium sp., 453
Zopfiella latipes, 175
Verticillium sphaerosporum var. bispora, 452
Zopfiella pilifera, 176
Volutella ciliata, 454
Zygomycetes, 23
Zygomycotina, 23, 93–131
Zygorhynchus moelleri, 131
1118-Afterword-Frame Page 485 Monday, March 4, 2002 1:27 PM
About 800 species of fungi are being catalogued in the Index of Fungi every year according to
Hawksworth et al. (1995). If 10% of them are assumed to be soilborne, more than 500 species
must be added to the number of soil fungi for the 7 years since the publication of the English
edition of this book in 1994. In the revised second edition, some new species include such recent
knowledge on soil fungi.
Biodiversity of soil fungi and their knowledge are significant in relation to environmental
problems practically and scientifically. Most soil fungi in any location may be identified if we know
nearly 300 fungal species because any soil fungus flora may not include more than 200 species
(see Supplement, Chapter 1). Therefore, the revised second edition, which includes more than
350 species belonging to at least 153 genera, may meet the demand for identification with quanti-
tative and qualitative improvement.
Tsuneo Watanabe
September 20, 2001
485
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Afterword
About 600 species of soil fungi were described by Gilman in 1945, but no one knows how many
species have been described as soil fungi since then. Total fungal species have increased by 1.7 times
from 1950 to 1983. Based on records on the number of fungi in the 3rd and 7th edition of Ainsworth
& Bisby’s Dictionary of the Fungi published in 1950 and 1983, respectively, at least 1200 species
of soil fungi have been described. Among 308 species described in this book, at least 20 are newly
described as soil fungi.
Although there are many mycological taxonomists worldwide, all of them have specialties in
particular fields, and they may be amateurs in other fields. Therefore, it is almost impossible to
describe various fungi covering the whole fungal fields alone, and would probably result in
misinterpretation or misidentification. However, bearing this in mind, this book was summarized
on the basis of the author’s own data. Therefore, all data and observations are not systematic and
purposive, including both detailed observations and data, and unsatisfactory observations and poor
data. However, all pictures and illustrations are based on the author’s own work, collected from
various sources during the past 25 years, irrespective of time, collection sites, hosts, types of works,
etc. Among materials, some are common, but others are different in having compared or summarized
all the data together, even if originally they were thought to be the same.
Although the results obtained are not always satisfactory, the author is happy to contribute to
influencing mycological interest in soil fungi. By summarizing this book, some new species could
be described, including the fresh data recently submitted for printing.
The author hopes finally for a prosperous and successful future in soil mycology, including
soil fungus taxonomy.
Tsuneo Watanabe
486