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Plant Disease
R. Kenneth Horst
Eighth Edition
1 3Reference
Westcott’s Plant Disease Handbook
R. Kenneth Horst

Westcott‘s Plant Disease

Eighth Edition

With 87 Figures and 2 Tables

R. Kenneth Horst
Department of Plant Pathology and Plant-Microbe Biology
Cornell University
Ithaca, NY, USA

ISBN 978-94-007-2140-1 ISBN 978-94-007-2141-8 (eBook)

ISBN 978-94-007-2142-5 (print and electronic bundle)
DOI 10.1007/978-94-007-2141-8
Springer Dordrecht, Berlin, Heidelberg, New York
Library of Congress Control Number: 2013934975

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This 8th Edition of the Plant Disease handbook is dedicated to the mentoring
experiences I have had the pleasure of experiencing starting with Pleasant View
Grade School, North Lawrence, OH, to Massillon Washington High School,
Massillon, OH, to Ohio University, Athens, OH, to The Ohio State University,
Columbus, OH, to Yoder Bros., Inc., Barberton, OH, to Cornell University, Ithaca,
NY (Professor, Department of Plant Pathology and Plant-Microbe Biology).
Although I felt in those early years that I was doing all the learning, I soon found that
mentoring was a two-way phenomenon. Not only was I mentoring my students at
Cornell University, but I found I was learning from them as well.
I was stimulated to reflect on this by my two youngest grandchildren, Madeline
Turner and Trevor Horst to whom I dedicate this 8th Edition as well as the students
who taught me while I was teaching them. Madeline initiated this process when
I asked her what she was learning in kindergarten. She listed all that she was
learning. I indicated she was really getting smart and that maybe Grampy should
go to kindergarten so that he could get smart. She said “No Grampy you can’t”
and I asked her “why not” and she said “Because you’re no kid anymore”. What
a great answer and also very profound since she was really telling me I needed to
continue moving beyond being a kid in my learning process.
Students (Masters, Doctoral and Post-Doctoral Students) I have mentored and
from whom I have also learned much.
Jamil Abu-Sadah Cristi Lynn Palmer
Richard Biamonte Hugh Allen Poole
Lester Burgess Leah LeEarle Porter
Eugene Oscar Erickson Ramona Ann Reiser
Donna Gardiner-Matteoni Charles Peter Romaine
Mary Handley Nancy Jane Schenk
Jeffrey R. Houge Margosheta Schollenberger
Hussein Ali Ahmed Hussein Gail Lynn Schumann
Catherine M. Klein Ann Finer Silverglate
Selin Kryzcynski Robert W. Stack
Randolph Edward McCoy Marek Szyndel
Robert J. McGovern Arnold T. Tschanz
Elzbieta Paduch-Cichal Sek-Man Wong
Preface to the Eighth Edition

It was a compliment to me to be asked to prepare the fourth edition of

Wescott’s Plant Disease Handbook, and the decision to accept the responsi-
bility for the fourth edition, the fifth edition, the sixth edition, the seventh
edition, and now the eighth edition was not taken lightly. The task has
been a formidable one. I have always had great respect professionally
for Dr. Cynthia Westcott. That respect has grown considerably with the
completion of the five editions. I now fully realize the tremendous amount
of effort expended by Dr. Westcott in developing the Handbook. A book such
as this is never finished, since one is never sure that everything has been
included that should be. In the 4 years since the seventh edition there were
more than 600 new reports of diseases on plants. I would quote and endorse
the words of Dr. Westcott in her preface to the first edition: “It is easy enough
to start a book on plant disease. It is impossible to finish it . . .” Dr. Cynthia
Westcott passed away March 22, 1983.
This revision of the Handbook retains the same general format contained
in the previous editions. The chemicals and pesticides regulations have been
updated; major taxonomic changes have been made in the bacteria, fungi,
nematodes and viruses; the changing picture in diseases caused by viruses
and/or virus-like agents have been described. New host plants have been
added, and many recently reported diseases as well as previously known
diseases listed now on new hosts have been included in the Handbook. In
addition photographs have been retained from the seventh edition as well as
the color photograph section. For the photography work I am grateful for the
help and expertise of Kent E. Loeffler. I also had access to the Cornell Plant
Pathology Herbarium, which contains a wealth of photographic work on plant
diseases that has been supplied by numerous scientists over many years.
This book should be useful to gardeners, master gardeners, botanical
gardens, landscape architects, florists, nurserymen, seed and fungicide
dealers, pesticide applicators, arborists, cooperative extension agents and
specialists, plant pathologists, diagnostic laboratories and consultants. The
book should also be a useful reference book for plant pathology classrooms
and in some cases used as a textbook.

March 2013 R. Kenneth Horst


I am indebted to many people for advice and suggestions for the 8th Edition.
The reviewers acquired by Kluwer Academic Publisher to review the
5th Edition and to advise on significance of a 6th Edition provided many
helpful suggestions which were used in the 7th and 8th Editions. Moreover,
a few individuals who were particularly helpful in my tasks of updating and
putting together the revision for the 8th Edition into an appropriate format
were J. Esnard, K. Hodge, S. J. Ingalls, K. Loeffler, C. Palmer, K. Snover,
R. E. Stall, B. Szyndel and M. S. Szyndel. Finally, I recognize and appreciate
the professional and efficient job of typing the manuscript by Margaret Haus
and her dedicated efforts in aiding me in proofreading, which was a major task
with the increasing size of the book and the changing scientific names of the
pathogenic organisms.

How to Use This Book

This is a reference manual. You will certainly not read it through from cover
to cover, but I hope you will read the first and last section of ▶ Part I on garden
chemicals. The chemicals themselves are listed in alphabetical order, by
common names where possible, by trade names where these are used in lieu
of approved common names. A few materials still in the experimental stage
but very promising are included. A few uses are suggested, but many more,
with correct dosages, will be found on the labels or in recent publications.
▶ Part II, on the classification of plant pathogens, can be taken or not as
desired. It provides a mycological, bacteriological, nematological and viro-
logical background for students and a review for professional workers. The
bibliography gives some of the taxonomic references consulted in preparing
this very condensed treatment.
The rest of the book is in two main sections. ▶ Part III describes specific
diseases and gives remedies when known. The diseases are grouped
according to their common names into forty types treated in alphabetical
order. ▶ Part IV gives over 1300 host plants in alphabetical order, from
Abelia to Zoysia, according to common names except where the Latin
name may mean less confusion. Under the hosts the diseases are sorted out
according to types, given in small capitals, and you can quickly thumb back to
the corresponding section, Anthracnose, Blight, Wilt, etc., in ▶ Part III by
means of the running head at the top of each page.
The book works like a dictionary. In both the disease and host section the
Latin name of the pathogen causing the disease is given in boldface type. The
individual diseases in the host section are listed in alphabetical order
according to the common name of the diseases.
You may be able to find the information you are seeking directly from the
index, which includes common and Latin names of hosts plants, Latin names
of pathogens and common names of the diseases described in ▶ Part III. More
than 4000 diseases are included in that chapter and some additional pathogens
are listed under Host Plants without a corresponding description of disease.
Website addresses of state universities and agricultural experiment sta-
tions, which are sources of help for every gardener, are given following
▶ Part IV. The very best way to use this book is to take it in small doses as
needed. Do not let the hundreds of diseases you will never meet worry you too
much. And remember that most plants survive, despite their troubles!


Part I Garden Chemicals and Their Application . . . . . . . . . . . 1

Fungicides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Bactericides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Nematicides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Virocides ............................................. 17
Applying The Chemicals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Spraying Vs Dusting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Mixing The Chemicals .................................. 25
All-Purpose Sprays and Dusts . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Integrated Pest Management . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

Part II Classification of Plant Pathogens . . . . . . . . . . . . . . . . . 31

Fungi ................................................ 33
Bacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Viruses, Viroids, Phytoplasmas . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Nematodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

Part III Plant Diseases and Their Pathogens . . . . . . . . . . . . . . 57

Anthracnose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Bacterial Diseases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Black Knot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Blackleg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Black Mildew . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Blackspot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Blights ............................................... 103

xiv Contents

Blotch Diseases ........................................ 143

Broomrapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Cankers and Diebacks .................................. 149
Club Root . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
Damping-Off . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Dodder . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179
Downy Mildews . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Fairy Rings ........................................... 187
Fruit Spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
Galls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191
Leaf Blister and Leaf Curl Diseases . . . . . . . . . . . . . . . . . . . . . . . . 195
Leaf Scorch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Leaf Spots ............................................ 201
Lichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237
Mistletoe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239
Molds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243
Needle Casts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
Nematodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
Nonparasitic Diseases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
Powdery Mildews ...................................... 285
Rots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295
Rusts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341
Scab ................................................. 363
Scurf . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 369
Slime Molds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 371
Smuts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 373
Snowmold . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Sooty Mold . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383
Spot Anthracnose ...................................... 385
Virus, Viroid, Phytoplasma – Pathogens and Diseases . . . . . . . . . 389
White Rusts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 431
Contents xv

Wilt Diseases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 433

Witchweed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 443

Part IV Host Plants and Their Diseases . . . . . . . . . . . . . . . . . . 445

Host Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 447
Color Plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 701
List of Land-Grant Institutions and Agricultural
Experiment Stations in the United States . . . . . . . . . . . . . . . . . . . 717
Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 719
Selected Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 725
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 733

The chief hazard any garden plant has to endure is its owner or gardener.
Moreover, many plants will suffer undue hardship from the publication of this
handbook. It is human nature to read symptoms of an ailment and immedi-
ately assume it is your own affliction. Jumping to conclusions is as dangerous
to plants as to humans. A sore throat does not necessarily mean diphtheria.
Only a trained physician can diagnose probable diphtheria, and for positive
identification a laboratory culture is necessary.
A spotted or yellowed rose leaf does not necessarily mean rose black spot.
Mite injury, spray injury, or reaction to weather conditions may also cause
spotted or yellow rose leaves; yet gardeners blithely continue increasing the
spray dosage, confident that more and stronger chemicals will control the
“disease” and seldom notice they are nearly killing the patient in the process.
A browning azalea flower does not necessarily mean the dreaded petal blight.
Some years ago a Westcott article on possible azalea troubles appeared in
print about the time azalea blooms in a Northern region were turning brown
from a combination of unusual weather conditions. Some gardeners immedi-
ately assumed the worst, thought that the southern blight had arrived in the
North, and started spraying. The poor plants, suffering from drought and
a heat wave, suffered additional injury from the additional stress of sprays.
All chemicals used as sprays or dusts are injurious to plants under some
conditions, the injury varying with the chemical and the dosage, with the
species and even the variety of plant, with temperature, soil moisture, and
many other factors. Plants suffering from drought are commonly injured by
So please, don’t jump to conclusions. Don’t do anything in a hurry because
the plants are getting sick fast and there is no time for a proper diagnosis. Don’t
rush to the seed store to buy some chemical you vaguely remember reading
about. Relax! You have all the time in the world for proper identification, since,
by the time the disease is serious enough for you to notice, it is probably too late
for protective spraying this season anyway.
Browning of an azalea flower means nothing as a diagnostic symptom. It
could just as well come from frost, heat, or old age as from a pathogen. If the
flowers are limp and collapsed with a slimy feel, these are good symptoms, but
signs of the fungus are needed as well. Thin, slightly curved black bodies
(sclerotia) formed at the base of petals are distinctive, but even more conclu-
sive are spores taken from the inside of the petals and examined under

xviii Introduction

a microscope. If these are one-celled, with a little box-like appendage, then

you may reasonably conclude that you have the true azalea petal blight.
This is a book of garden diseases, but it is not expected that anyone,
amateur or professional, can read a brief description, look at an unfamiliar
disease in the garden, and make a very reliable diagnosis. I certainly cannot,
and after compiling this tome I am less likely to try than ever before. I have
written “water-soaked” or “reddish brown” too many hundreds of times for
different diseases to make such symptoms seem very distinctive.
However, if you are a gardener, you can narrow the field down consider-
ably by consulting ▶ Part IV, where host plants are listed in alphabetical
order, and under each the type of disease –Blight, Canker, Leaf Spot,
etc. – and then the organisms causing these diseases by their scientific
names and the states where they have been reported. Eliminating the types
of disease that are obviously different from yours and eliminating diseases
that are reported only on the West Coast when you live in New York, you may
find only two or three possibilities to look up in ▶ Part III, which lists, under
the different disease groups, the pathogens in alphabetical order, followed by
a discussion of each disease. In situations where pathogen names have been
changed due to critical investigations of spore formation and development,
the original name is listed in alphabetical order followed by “see new name”.
Under the new name in parenthesis “formerly old name” is indicated.
Don’t let all the scientific names worry you. It is the only way to make this
a quick and easy reference, for there are very few common names of plant
diseases that can be used without confusion. It works just like the telephone
book. While thumbing your way down to Smith, John, you do not worry about
spelling Smiecinski, C., which you pass on the way.
If you are a quasi-professional, with little or no formal mycology but trying
to keep abreast of a flood of miscellaneous specimens, there is a brief review
for you of the salient microscopic characteristics of each genus, together with
its classification. This is in small type and may be readily passed over by those
interested solely in macroscopic characteristics.

What is Plant Disease?

There are many definitions of plant disease, the simplest being any deviation
from the normal. The concept of the late professor H. H. Whetzel, a great
teacher of plant pathology who influenced many students including
Dr. Cynthia Westcott, is valid and appropriate even today. “Disease in plants
is an injurious physiological process, caused by the continued irritation of
a primary causal factor, exhibited through abnormal cellular activity and
expressed in characteristic pathological conditions called symptoms.” The
causal factor may be a living organism or an environmental condition. Injury
differs from disease in being due to the transient irritation of a causal factor,
as the wound of an insect, sudden freezing or burning, application of a poison.
Plant diseases may be necrotic, with dying or death of cells, tissues, or
organs; hypoplastic, resulting in dwarfing or stunting; or hyperplastic, with an
overgrowth of plant tissue, as in crown gall or club root.
Introduction xix

Plant Diseases Are Not New

All species of plants, wild and cultivated, are subject to disease. Fossil
remains suggest that plant diseases were present on earth before man himself.
Certainly man has been punished by them ever since the Garden of Eden.
“I smote you with blasting and with mildew and hail in all the labors of your
hands yet ye turned not to me, saith the Lord” (Haggai 2:17).
Man’s attempts at controlling plant disease go back at least to 700 B.C. when
the Romans instituted the Robigalia to propitiate the rust gods with prayer and
sacrifice. About 470 B.C. Pliny reported that amurca of olives should be
sprinkled on plants to prevent attacks of blight, this being our earliest known
reference to a fungicide, although Homer, 1000 B.C., wrote of “pest-averting
In 1660 at Rouen, France, a law was passed calling for eradication of the
barberry as a means of fighting wheat rust, two centuries before anyone knew
the true nature of rust or how barberry affected wheat.
In the latter part of the eighteenth century the Englishman Forsyth dis-
coursed on tree surgery and treatment of wounds and cankers. His seemingly
fantastic recommendation of a paste of cow dung to promote healing of tree
wounds has modern corroboration in research showing that urea speeds up
healing of such wounds.
Much of our progress in dealing with plant disease has followed spectac-
ular catastrophes. Modern plant pathology had its start with the blight that
swept the potato fields of Europe in 1844 and 1845, resulting in the Irish
famine. This lesson in the importance of plant disease to the economic
welfare of mankind marked the beginning of public support for investigations
into the cause of disease. Two men, both German, laid the firm foundations of
our present knowledge. Mycologist Anton de Bary, 1867 to 1888, first proved
beyond doubt that fungi associated with plant diseases were pathogenic,
while Julius Kuhn, farmer with a doctor’s degree in science, first showed
the relation between science and practice in the problems of plant disease
control. His textbook on Diseases of Cultivated Plants, published in 1858, is
still useful.
The accidental discovery of bordeaux mixture in France in 1882 marks the
beginning of protective spraying for disease control, but the use of drugs goes
back to 1824, when sulfur was recommended as an eradicant for powdery
mildew. The development of synthetic organic fungicides was sparked by
World War II, partly as a result of a search for chemicals to mildew-proof
fabrics used by the armed forces. Antibiotics for plant disease control
followed their use in medical practice, with a great deal of research in this
field since 1949.
Since the establishment of the Environmental Protection Agency in 1972
there has been increased concern on the use of toxic chemicals for controlling
plant disease. Moreover, this concern has generated renewed interest in
integrated pest management (IPM) and biological control strategies in the
1980’s. IPM utilizes all available pertinent information regarding the crop or
plant, its pathogens, the environmental conditions expected to prevail,
xx Introduction

locality, availability of materials, and costs in developing the control pro-

gram. Biological control is the total or partial destruction of pathogen
populations by other organisms. This phenomenon occurs routinely in nature.
There are several diseases in which the pathogen cannot develop because the
soil, called suppressive soils, contain microorganisms antagonistic to the
pathogen, or because the plant that is attacked has been naturally inoculated
before or after the pathogen attack, with antagonistic microorganisms. Even
higher plants may reduce the amount of pathogen inoculum by trapping
available pathogens (trap plants) or by releasing substances toxic to the
pathogen into the soil. Although biological antagonisms are subject to numer-
ous ecological limitations it can be expected to become an important part of
control measures employed against many more diseases in future years.

Plant Pathology in the United States

Organized plant pathology in the United States started in 1885 with a section
of Mycology in the U.S. Department of Agriculture. In 1904 the start of the
great epiphytotic of chestnut blight, which was to wipe out our native trees,
stimulated more public interest and support for plant pathology. In 1907 the
first university Department of Plant Pathology was established at Cornell
The United States Quarantine Act of 1912 officially recognized the possi-
bility of introducing pests and diseases on imported plants, after low-priced
nursery seedlings from Europe had brought in the white pine blister rust. This
was our first attempt at control by exclusion.
In 1917, during World War I, the Plant Disease Survey was organized as an
office of the Bureau of Plant Industry “to collect information on plant diseases
in the United States, covering such topics as prevalence, geographical distri-
bution, severity, etc., and to make this information immediately available to
all persons interested, especially those concerned with disease control.”
During World War II the Plant Disease Survey was in charge of the emer-
gency project “to protect the country’s food, feed, fiber and oil supplies by
ensuring immediate detection of enemy attempts at crop destruction through
the use of plant diseases and providing production specialists and extension
workers with prompt and accurate information regarding outbreaks of plant
diseases whether introduced inadvertently or by design while still in incipient
stages.”As a by-product of these wartime surveys we accumulated a good
deal of evidence on the prevalence of new and established diseases across the
country, in home gardens as well as on farms.
In 1946, a century after Phytophthora infestans had made history with the
potato blight, a strain of the same fungus started an unprecedented epiphytotic
of tomato blight. This disaster led to the forecasting service warning dealers
and growers when certain diseases are imminent.
The Plant Disease Survey has now become the Epidemiology Investiga-
tions Section of the Agricultural Research Service of the U.S. Department of
Agriculture. The Agricultural Research Service became a part of the Science
and Education Administration in 1978. It issues a monthly bulletin, The Plant
Introduction xxi

Disease Reporter, based on reports from qualified volunteer collaborators all

over the country. The American Phytopathological Society assumed the
responsibility for publishing this journal in 1980 and the journal was renamed
Plant Disease. Much of the material in this handbook is taken from these

Principles of Control

Control of a plant disease means reduction in the amount of damage caused.

Our present annual toll from disease is nearly four billion dollars. Perfect
control is rare, but profitable control, when the increased yield more than
covers the cost of chemicals and labor, is quite possible. Commercial growers
now average a return of four dollars for each dollar so invested. Keeping
home plantings ornamental yields a large return in satisfaction and increased
property value.
The five fundamental principles of control are exclusion, eradication,
protection, resistance, and therapy.
1. Exclusion means preventing the entrance and establishment of pathogens
in uninfested gardens, states, or countries. For home gardeners it means
using certified seed or plants, sorting bulbs before planting, discarding any
that are doubtful, possibly treating seeds or tubers or corms before they are
planted, and, most especially, refusing obviously diseased specimens from
nurseryman or dealer. For states and countries, exclusion means quaran-
tines, prohibition by law. Sometimes restricted entry of nursery stock is
allowed, the plants to be grown in isolation and inspected for one or two
years before distribution is permitted.
2. Eradication means the elimination of a pathogen once it has become
established on a plant or in a garden. It can be accomplished by removal
of diseased specimens, or parts, as in roguing to control virus diseases or
cutting off cankered tree limbs; by cultivating to keep down weed hosts
and deep ploughing or spading to bury diseased plant debris; by rotation of
susceptible with nonsusceptible crops to starve out the pathogen; and by
disinfection, usually by chemicals, sometimes by heat treatment. Spraying
or dusting foliage with sulfur after mildew mycelium is present is eradi-
cation, and so is treating the soil with chloropicrin to kill nematodes and
3. Protection is the interposition of some protective barrier between the
susceptible part of the suspect or host and the pathogen. In most instances
this is a protective spray or dust applied to the plant in advance of the
arrival of the fungus spore; sometimes it means killing insects or other
inoculating agents; sometimes it means the erection of a windbreak or
other mechanical barrier.
▶ Part I gives an alphabetical list of chemicals used in present-day
protective spraying and dusting, along with eradicant chemicals, and
includes notes on compatibility and possibilities of injury. It is here
that home gardeners, sometimes commercial growers, can do their plants
irreparable harm instead of the good they intend. Spraying is never to be
xxii Introduction

undertaken lightly or thoughtlessly. Stop and think! Read all of the fine
print on the label; be sure of your dosage and the safety of that particular
chemical on the plant you want to protect, to say nothing of precautions
necessary for your own safety.
4. Resistance is control by the development of resistant varieties. Resistant
varieties are as old as time. Nature has always eliminated the unfit, but since
about 1890 man has been speeding up the process by deliberately breeding,
selecting, and propagating plants resistant to the more important diseases.
Resistant ornamental plants have lagged behind food plants, but we do have
wilt-resistant asters, rust-resistant snapdragons, wilt-resistant mimosas. Here
is the ideal way for home gardeners to control their plant diseases – in the
winter when the seed order and the nursery list is made out – so easy, and
so safe!
5. Therapy is control by inoculating or treating the plant with something that
will inactivate the pathogen. Chemotherapy is the use of chemicals to
inactivate the pathogen, whereas heat is sometimes used to inactivate or
inhibit virus development in infected plant tissues so that newly develop-
ing tissue may be obtained which is free of the pathogen. The use of this
procedure is discussed in ▶ Part II.
Part I
Garden Chemicals and Their Application

A fungicide is a substance that destroys or inhibits the growth of fungi. It may

be an eradicant, applied to a plant, plant part, or the environment as a curative
treatment to destroy fungi established within a given area or plant; or prefer-
ably it may be a protectant, applied to protect a plant or plant part from
infection by killing, or inhibiting the development of, fungal spores of
mycelium that may arrive at the infection court. A bactericide is a
substance that destroys or inhibits bacteria and nematicide for nematodes.
Among the more recent bactericides are antibiotics, products of other living
organisms. They also have value against certain fungi. There are few
virocides, which are toxic or poisonous to viruses.
A pesticide is any chemical that is used to kill pests, especially insects
and rodents.
An insecticide is an agent used to kill insects.
A disinfectant is an agent that frees a plant or plant part from infection by
destroying the pathogen established within it. A disinfectant kills or inacti-
vates organisms present on the surface of the plant or plant part or in the
immediate environment. Chemicals for seed treatment can be either
eradicants or protectants, but most of them are disinfestants, in that they kill
organisms on the surface of the seed rather than those within. In common
usage, however, they are called disinfectants.
A nematicide is, of course, a chemical that kills nematodes in the soil or in
the plant. Most nematicides are fumigants, chemical toxicants that act in
volatile form.
Not so long ago the chemicals on the garden medicine shelf consisted
of copper and sulfur for protectants, lime sulfur as an eradicant, mercuric
chloride as a disinfectant, and formalin and carbon bisulfide as
fumigants. You sometimes got plant injury; you did not always get the best
2 I Garden Chemicals and Their Application

possible control, but at least you did not have to be an organic chemist. Now
we have the following classes of fungicides:

Inorganic Organic
Sulfur Dithiocarbamates
Copper Thiazoles
Salts Triazines
Sterol Inhibitors

The search for new fungicides goes on, with hundreds of synthetic organic
compounds being screened each year. This screening is often a cooperative
venture between manufacturers, state experiment stations, and the U.S.
Department of Agriculture. After safety precautions for the operator and the
environment, and the effectiveness of a compound for certain diseases have
been determined, the chief question is whether the material is phytotoxic, that
is, injurious to plants, at concentrations required for control. Phytotoxicity is
an elusive factor, not to be pinned down in a few tests. It varies not only with
the kind of plant but with the particular variety, the amount of moisture in the
soil when the spray is applied, the temperature, whether or not the application
is followed by rain or high humidity, the section of the country, and the
compatibility of the chemical with spreaders or wetting agents, as well as with
other fungicides or insecticides. Coordinated tests with new materials in
many different states are extremely valuable. Some compounds give rather
uniform results over the country; others vary widely with climatic conditions.

The 1947 Federal Insecticide, Fungicide and Rodenticide Act (FIFRA)

provides that all fungicides must be registered with the U.S. Department of
Agriculture before being marketed. Materials highly toxic to humans must be
prominently marked, instructions given for avoiding injury to plants or
animals, the toxicant chemical named, and the percentage of active and
inactive ingredients given. All labels submitted for registration must be
accompanied by proof that the claims for performance are valid.
In 1954, Public Law 518, known as the Miller Bill, was passed, providing
for tolerances. A tolerance is the legal limit of a poisonous residue, expressed
in parts per million (ppm), that may remain on an edible product at the time it
is distributed for consumption. In 1958, The Food Additives Amendment was
passed, which also controls pesticides residues in processed foods. It included
the Delaney clause, which states that any chemical found to be a carcinogen in
laboratory animals may not appear in a human food, a zero tolerance. In 1959,
the FIFRA was amended to include nematicides, plant growth regulators,
Garden Chemicals and Their Application 3

defoliants, and desiccants as pesticides. Since that time, poisons and repel-
lents used against all classes of animals (from invertebrates to mammals)
have been brought into the approval process.
FIFRA was further amended in 1972 as the Federal Environmental Pesti-
cide Control Act (FEPCA), making violations by growers, applicators, or
dealers subject to heavy fines and/or imprisonment. All pesticides had been
classified into either general-use or restricted-use categories by October 1977,
with anyone applying restricted pesticides required to be state-certified.
Pesticide manufacturing plants are to be registered and government-
inspected. All pesticide products must be registered whether shipped in
interstate or intrastate commerce. Other provisions are of various degrees of
importance to concerned persons or companies.
Additional modifications were made in FIFRA in 1989. The modifications
specifically will (1) accelerate re-registration of older pesticides (those reg-
istered prior to November 1984) and impose fees on chemical manufacturers
for re-registration; (2) essentially eliminate indemnification payments to
those holding inventories of suspended or canceled pesticides, except farmers
and certain end users; and (3) shift part of the burden for storage and disposal
of banned pesticides from the government to the manufacturer. The 1989
FIFRA also empowers the Environmental Protection Agency (EPA) to
change regulations on how applicators handle, rinse, and dispose of pesticide
containers. The Worker Protection Standard (WPS), passed in 1992, required
labels to carry re-entry intervals (REI) and personal protection equipment for
certain end-uses. Labels having both WPS and non-WPS uses are required to
have recommendation for PPE and REI for both categories.
In 1996, the Food Quality Protection Act (FQPA)was passed, requiring
a 10 year review process of all currently registered pesticides based on a risk
cap of aggregate exposure, in other words all possible ways people, particu-
larly children, can come into contact with pesticides whether through residues
on food or backyard applications. Part of FQPA determined that an acceptable
level of risk is a reasonable certainty of no harm, essentially eliminating the
Delaney Clause from 1958. The FQPA also provided for accelerated regis-
trations of safer pesticides, such as biological control organisms or active
ingredients with the potential for minimal environmental impact.
The federal government considers these to be minimum pesticide regula-
tions. Any state may choose to establish more rigid pesticide regulations
within its boundaries than those legislated by the federal government, and
some have done so. Some states require notification to be posted prior to
commercial pesticide application including chemicals used. Thus, pesticide
applicators must be familiar with individual state pesticide regulations as well
as federal pesticide regulations.
Consumers, therefore, are well protected against fraud, but they must
be willing to read the fine print on labels if they are to choose intelligently
from the bewildering array of proprietary compounds on dealers’ shelves. In
the following pages there are lists of available products, cross-referenced
by product name and the common name of active ingredients. In many
cases, several product names may exist for the same active ingredient; those
readily available to consumers may not be listed so pay particular attention to
4 I Garden Chemicals and Their Application

the name of the active ingredient present on the available products. Con-
sumers must also read the fine print and follow directions exactly if their
homegrown vegetables are to be as safe for consumption as those from
commercial growers who have to comply with the law in the matter of
Even if you follow exactly the directions for dosage given on the label, you
may have some plant injury under your particular combination of soil,
weather, and kinds of plants. Keep a notebook. Put down the date you
sprayed, the dosage used, the approximate temperature and humidity,
whether it was cloudy or sunny, in a period of drought or prolonged wet
weather. Go around later and check for burning; for leaf spotting and defo-
liation from the spray or from failure to control the disease; for leaf curling or
stunting; for too much unsightly residue. Note which varieties can take the
spray and which cannot.
The following alphabetical list includes chemicals now commercially
available, a few that are rather outmoded but still found in textbooks, a few
that were marketed in the past but have now been discontinued, and a few that
will probably be marketed before this text is published. By that time there will
be many more that should have been included, for the search for better
chemicals is unending. There will also be more that will be discontinued.
The list presented herein must be considered only as a guide. Exclusive
reliance must be placed on directions and information supplied by the man-
ufacturer or by agricultural specialists, agents, or advisors. Be sure to read
the label. Because so many of the new compounds have long, complex
chemical names, they have been given short common names by the American
Standards Association. Such common names are listed first in the following
listing. Frequently, however, the trade name is used as a common name; trade
names are listed in parenthesis following the common names. The Crop
Protection Handbook, which is published each year by Meister Publishing
Co., 37841 Euclid Ave., Willoughby, OH, 44094, gives an up-to-date listing
of pesticides. Vance Publishing publishes the Turf and Ornamental Reference
and the Crop Protection Reference (Greenbook).
As always, read and follow label directions carefully. If unsure whether
a listed product is registered in your area, contact local, state, and federal
In the following lists, the common chemical name is given first, in bold,
followed by trade names available for professional growers of agricultural,
turf and ornamental crops and products available for consumers. Then there is
a brief description of target pathogens.

Azoxystrobin of water. Stock solutions are made up for each

chemical (1 pound per gallon of water), the lime
Abound, Amistar, Bankit, Heritag, Ortiva, Priori, solution placed first in the sprayer, diluted to
Quadris nearly the full amount, and the copper sulfate
Systemic fungicide for control of foliar and solution added. Or, for power sprayers, finely
soil borne diseases. divided copper sulfate can be washed through the
strainer into the spray tank, and when the tank is
two-thirds full the weighed amount of hydrated
Bacillus subtilis lime can also be washed through the strainer
while the agitator is running. Casein or other
Serenade, Subtilex, Taego, Companion, Kodiak spreader is added toward the end. Phytotoxicity
Used as a seed treatment to suppress Rhizoc- comes from both the lime and the copper. Plants
tonia and Fusarium diseases. are often stunted, with yield reduced; fruit-setting
of tomatoes may be delayed. Bordeaux is not safe
on peaches during the growing season, may burn
Benalaxyl and russet applies (both foliage and fruits), may
cause red spotting, yellowing, and dropping of
Galben, Tairel rose leaves (often confused with blackspot by
Systemic fungicide. amateur and sometimes professional gardeners),
and may cause defoliation of Japanese plums.
Injury is most prominent early in the season
Bitertanol when temperature is below 50  F and in dull,
cloudy weather when light rain or high humidity
Bacseal, Baycor, Baycoral, Baymat, Zaron prevents rapid drying of the spray. Late summer
Fungicide. use of bordeaux is credited with making some
plants more susceptible to early fall frosts. For
ornamentals, a 4-4-100 mixture is usually strong
Bordeaux Mixture enough and can be made in small amounts by
dissolving 2 ounces of copper sulfate in 1 gallon
Comac of water, 2 ounces of hydrated lime in 2 gallons of
Bordeaux mixture is made in varying concen- water, pouring the copper sulfate solution into the
trations. The most usual formula is 8-8-100 (often lime water, and straining into the spray tank
stated as 4-4-50), which means 8 pounds copper through fine cheesecloth. For some plants, such
sulfate, 8 pounds hydrated lime to 100 gallons as stone fruits, the proportion of lime is increased;

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_1, 5

# Springer Science+Business Media Dordrecht 2013
6 Fungicides

for others, such as azaleas, a low-lime bordeaux is

used. Once the two solutions have been mixed, the Chloranil
preparation must be used immediately. Fresh lime
is essential, not some left over from a previous Foliar fungicide and seed treatment.
season. Somewhat less effective than homemade
bordeaux but easier for the home gardener are the
various powders and pastes available under trade Chloroneb
names; to these add only water at the time of use.
Terraneb, N-Flow D, Proturf

Nimrod Chloropicrin
Systemic fungicide.
Soil fumigant.
Burgundy Mix

A soda bordeaux formerly used, prepared with Chlorothalonil

washing soda instead of lime.
Bravo, Bravo Ultrex, Bravo Weatherstik
Burkholderia cepacia

Fungicide, nematicide. Copper Acetate

First developed in 1889; became the first factory

Captan made basic copper fungicide.

Captan 30 DD, Captan 400

Protectant, eradicant fungicide. Copper Ammonium Carbonate

Carbendazim Bactericide, Fungicide.

Bavistin, Decrosol
Systemic fungicide. Copper Carbonate

Foliar fungicide, Seed treatments.

Carbon Disulfide

Soil fumigant. Copper Compounds

More stable than bordeaux mix, less phytotoxic,

Carboxin easier to use, and with less objectionable residue.
Copper sprays control many blights, leaf spots,
Vitavax downy and powdery mildews. They are incompat-
Fungicide, seed protectant. ible with lime sulfur, questionable with cryolite,
Dimethomorph 7

benxene hexachloride, tetraethyl pyrophosphate,

organic mercuries, and thiocarbamates. They Cyproconazole
may injure plants in cool, cloudy or moist weather.
Injury to apple and rose foliage varies from red- Alto, Atemi, Sentinel
dish spots to yellowing and defoliation. Systemic and eradicative fungicide.

Copper Hydroxide Cyprodinil

KOP Hydroxide Chieftain, Chorus, Unix, Vangard
Fungicide, bactericide. Fungicide, seed treatment.

Copper Oxide
Chem Copp, Cuprocop, Cuprox, Caocobre, Cop-
per Sandoz Basimid
Protective fungicide. Soil fumigant.

Copper Oxychloride Dichlofluanid

KOP Oxy-85, Recop, Coprantol, Cupramar, Elvaron, Euparen

Cupravit Fungicide.
Protective fungicide.

Copper Sulfate
Allisan, Botran, Clortran
Tribasic (Flurane, Idrorame, King, Diachum) Fungicide.

Cufraneb Difenoconazole

Bardos, Bogard, Dividend, Geyser, Score, Sico

Fungicide. Miticide, Seed Dressing.
Systemic fungicide.

Antibiotic fungicide from Streptomyces griseus,
the first antibiotic introduced (1949) for control Milcurb
of plant disease. Systemic fungicide.

Cymoxanil Dimethomorph

Curzate Acrobat, Forum

Fungicide. Oomycete fungicide.
8 Fungicides

Diniconazole Fenbuconazole

Mitazole Enable, Govern, Indar

Systemic fungicide. Systemic fungicide.

Dithianon Fenpiclonil
Delan Beret, Electer, Gambit

Dodemorph Acetate
Mildin, Patrol, Sorilan, Tern
Meltatox Systemic fungicide.

Systemic fungicide.

Epoxiconazole Ferbam

Opal, Opus Ferman Granuflo

Systemic fungicide. Fungicide.

Etridiazole Filipin

Terrazole, Koban, Truban Antibiotic. Fungicide for seed-rot fungi.

Soil fungicide.

Protects germinating seeds and seedlings in corn, Fungicide, Miticide.
beans, peas, spinach, cucumbers, and ornamentals.

Celest, Geoxe, Maxim, Medallion, Saphire, Sav-
Used for powdery mildew and rust on ornamen- ior, Scholar
tals, turf, and tree crops. Contact fungicide.
Lime Sulfur 9

Flusilazole Hydroxyquinoline Sulfate

Fungicide. Chinosol
Systemic fungicide, bactericide.

Folistar, Moncut, Prostar
Systemic fungicide. Bromazil, Deccozil, Double R11, Flo Pro IMZ,
Freshgard 700, Impala, Nu-Zone
Systemic fungicide.

Folpet Iprodione
Protective fungicide.
Chipco, Kodan, Rovral, ProTurf, Fungicide X
Contact/locally systemic fungicide.

Aliette, Chipco
Systemic fungicide, bactericide.

Alliage, Candit, Cygnus, Discus, Sovran, Stroby

Hexaconazole Surface systemic fungicide; protective and
curative effects.
Anvil, Planete Aster
Lime Sulfur

Hydrated Lime and Copper Sulfate Polysulfides formed by boiling together sulfur
and mild of lime. The standard liquid has
Bordeaux Mix a specific gravity of 32 Baume and the commer-
Prepared from copper sulfate and lime to form cial product is far superior to the homemade.
a membranous coating over plant parts, the first Lime sulfur dates back to 1851, when the head
protective spray and still widely used. About gardener, Grison, at Versailles, France, boiled
1878, French vineyard were threatened with together sulfur and lime for a vegetable fungicide
downy mildew, which had been introduced from called “Eau Grison.” In 1886, this fungicide was
the U.S. Millardet, one of the workers assigned to used in California as a dormant spray for San Jose
the problem, noticed that where grapes near the scale and later for peach leaf curl. A self-boiled
highways to Bordeaux had been treated with lime sulfur made without heat was produced in
a poisonous-looking mixture of copper and lime 1908 as a summer spray for sensitive plants, but it
to prevent stealing, there was little or no downy was later replaced by wettable sulfurs for most
mildew. A description of the preparation of bor- fruit-spray programs. A dry form of lime sulfur
deaux mixture was published in 1885, and it was marketed about 1908. Used as a dormant
remains a most efficient fungicide. It does, how- spray for fruits, roses, and some other plants for
ever, have a most conspicuous residue and is mildews, Volutella blight of boxwood, and other
injurious to some plants. diseases. Do not use above 85  F.
10 Fungicides

Mancozeb Nitrothal-Isopropyl

AgriSolutions, Mancozeb, Penncozeb, Pennfluid, Pallitop

Tridex, Trimanin, Trimanoc, Trimanzone, Fungicide.
Triziman, Triziman D, Vandozeb, Protect T/O,
Mancozin, Manzin, Dithane, Fore, Manzate, Pen-
tathlon DF, Pentathlon LF Oxadixyl
Recoil, Ripost, Sandofan, Wakil
Systemic fungicide.

Trimangol, Vondac, M, Manex, Manox, Man-Zox,

Pentathon, Maneb Spritzpulver, Manex, Manzi
Systemic fungicide.

Metax, Allegiance, ProTurf Soil and foliar fungi- Quintozene

cide; seed dressing fungicide.
Blocker, Parflo, Winflo, Defend, Terraclor,
Turfcide, FF11, Penstar, RTU, PCNB
Metalaxyl M Soil fungicide; seed dressing agent.

Quell, Apron XL, Ridomil Gold, Subdue 2X,

Subdue MAXX Penconazole
Omnex, Topas
Systemic fungicide.

Metam 426, Polefume, Turfcure, Vapam, Vapam

HL, Busan 1236, Trimaton, Sectagon 42, Metam
CLR Soil fumigant –fungicide, insecticide,
Monceran, Trotis

Metiram Piperalin

Polyram Pipron
Contact fungicide. Fungicide.

Myclobutanil Potassium Bicarbonate

Eagle, Nova, Rally, Systhane Armicarb, GreenCure, Milstop Broad spectrum

Fungicide. fungicide; K bicarbonate plus surfactants.
Thiophanate Methyl Plus Etridiazole 11

Potassium Bicarbonate Quarternary Ammonium Compounds

Kaligreen Barquat, Hyamine

Fungicide; K bicarbonate, no surfactants. Fungicide, Bactericides, Surfactants.

Prochloraz Sulfur

Abavit, Ascurit, Oczave, Omega, Prelude, Kumulus, CSC Dusting, CSC Thioben, CSC
Sporgon, Sportak Thiosperse, Crisazufre, Sulfox, Suffa, Sulfa,
Fungicide. Red Ball, Microsulf, Thiolux, Thiovit, Bensul,
Golden Demo Signal, Special Electric, Comoran,
Comoran Supra, Cosan, Elosal, Kolodust,
Propamocarb Kumulus, Microsperse
Fungicide, Miticide.
Hydrochloride (Banol, Dynone, Filex, Prevex,
Previcar, Previcur N, Win)
Fungicide. Tebuconazole

Corail, Elite, Folicur, Horizon, Horizon Arbo,

Horizon T, Lyux, Raxil
Systemic fungicide.
PropiMaX, Alamo, Banner, Break, Orbit, Tilt,
Systemic fungicide; seed treatment. Tetraconazole

Propineb Eminent
Systemic fungicide.
Antracol, Inicol
Contact fungicide.

Pyrifenox Arbotect, Mertect, Storite, Storite Excel, Tectab,

Tecto, APL-Luster, APL Luster T, Decco 205,
Corona, Dorado Brogdex, Freshgard 598, Gustafson SP
Systemic fungicide. Systemic fungicide.

Pyroquilon Thiophanate Methyl

Coratop, Fongorene Domain, Fungo, Mildothane, Topsin M, 3336,

Systemic fungicide. Fansin-M, OHP 6672

Thiophanate Methyl Plus Etridiazole
Afugan, Curamil
Systemic fungicide. Banrot Soil fungicide.
12 Fungicides

Thiram Trifloxystrobin

Tripomol, Defiant, Thiram Granuflo, Aatack, Compass, Flint, Gem, Twist, Compass O
Aules, Chipco, Thiram 75, Pomarsol, Forte, Fungicide.
42-S Thiram, Rhodiasan Express, Spotrete
Fungicide, Seed protectant. Triflumizole

Procure, Terraguard
Systemic fungicide.

Bayleton, Strike
Systemic fungicide.

Denarin, Funginex, Saprol


Bayton, Bayfidan, Cereous, Trisan, Bayton 30 Triphenyltin Hydroxide

Systemic fungicide.
Brestanid, Agri Tin Fungicide.

Trichoderma Harzianum
Plant Shield, Root Shield, Top Shield
Foliar, Soil fungicide. Ronilan, Curalan, Vorlan

Tricyclazole Zineb
Cuprothex, Super Mixy
Systemic fungicide.

Tridemorph Ziram

Calixin Tricarbamix, Triscabol, Cuman, Pomarsol 2

System fungicide. Fungicide.

Agrobacterium radiobacter Copper Hydroxide

Galltrol-A, Strain 84, Nogall, Strain K-1026 Cudrox, Cuidrox, Blue Shield, Kocide, Spin Out,
Crown gall preventative bactericide. KOP Hydroxide, NuCop

Benzalkonium Chloride
Copper Sulfate
Barquat, Hyamine, Nacco San
Bactericide. Basic (Cuprofix, Disperss, Cuprofix, MZ
Disperss, Basic Cooper 53, Cop-O-Zinc 25–25.
Basicop) Bactericide. ▶ Fungicides.

Bronotak Dazomet
Bactericide, bacteriostat.
Bactericide, Soil fumigant.

Copper Ammonium Carbonate

Dimanin A
Bactericide. Bayclean

Copper, Fixed
Includes Cu hydroxide, Cu oxide, Cu oxychloride
sulfate, Cu oxychloride, Cu sulfate. Bactericide, crown gall eradicant paint.

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_2, 13

# Springer Science+Business Media Dordrecht 2013
14 Bactericides

Hydroxyquinoline Sulfate Quarternary Ammonium Compounds

Chinosol Arquad 2C-75, Barquat, Hyamine

Bactericide. Bactericide.

Streptomycin (Nitrate or Sulfate)

Oxytetracycline Hydroxide
Agricultural Streptomycin, Streptrol, Agr-Mycin
Mycoshield 17, As-50
Bactericide. Bactericide.

Carbofuran Du Nema

Furadan, Curaterr Nematicide, used on turf.


Chlorpicrin Ethylene Dibromide

Chlor-O-Pie Nematicide, Soil fumigant.

Nematicide, Soil fumigant.

Dazomet Metam-Sodium

Basamid Busan 1236, Trimaton

Nematicide, Soil fumigant. Nematicide, Soil fumigant.

Dichloropropene Triazophos

DD-92, In Line, Telone, Telone II Hostathion, Trelka

Nematicide, Soil fumigant. Nematicide.

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_3, 15

# Springer Science+Business Media Dordrecht 2013


Virus inhibitor.

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_4, 17

# Springer Science+Business Media Dordrecht 2013
Applying The Chemicals

Spraying is the application of a chemical to and also in order to see the distribution of the
a plant in liquid form; dusting the application of concentrates, they often have to be used at night.
a fine dry powder. The difference between They are not too efficient for very tall trees, and
spraying and dusting was very clear-cut before the droplet size has to be rather carefully regu-
aerosol bombs, mist blowers, and fog machines lated. Too large drops may fall out before they
were developed to apply liquids in such concen- reach a tree, and too small drops may not settle
trated form that the particles are practically dry down but go on past.
before they reach the plant and before spray- Although we usually think of mist blowers on
dusters were made to deliver wetted dusts. trucks for large scale operations, there are now
Sprayers vary from a flit gun or pint atomizer, some about the size of knapsack sprayers that,
which takes an hour to discharge a gallon, to engine and all, are worn on the back around the
power apparatus that discharges 60 gallons a garden. They weigh around 35 pounds and will
minute at 800 pounds pressure from a 600-gallon cover foliage up to 30 feet. They cost, however,
spray tank. Dusters vary from the small somewhat more than the hydraulic power
cardboard or plastic carton in which the dust is sprayers of small estate size.
purchased to helicopters. Applicators for pressur-
ize sprays or aerosols vary from the one-pound
“bomb” to truck-mounted fog generators or air Hydraulic Sprayers
blast machines. See Fig. 1 for various applicators.
Mist blowers will probably never entirely out-
mode hydraulic sprayers, which can place the
Mist Sprayers spray more accurately, at a greater height, and
can operate under more unfavorable weather con-
In orchards and in shade tree work there has been ditions. For trees, high gallonage per minute and
increasing use of mist blowers, air blast machines enough pressure to drive sprays high in the air
that carry droplets of concentrated pesticides to have advantages, but for garden plants the
plants in air rather than water. They are speedier emphasis should be on cutting down gallonage
than hydraulic sprayers, use far less water, which and pressure.
may be scarce in times of drought, and do not Power sprayers for home gardens are available
leave puddles or poisonous run-off which may be in almost any size, from 5-gallon capacity on up,
dangerous to pets and birds. They cannot, how- and may have gasoline or electric motors
ever, be operated in much wind; for that reason, (see Fig. 2). For the orchard a spray gun is

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_5, 19

# Springer Science+Business Media Dordrecht 2013
20 Applying The Chemicals

Figure 1 Aerosol
pressurized spray

Shaker Pressure Trigger Continuous Hose End

Can Can Sprayer Sprayer Sprayer

Bucket Slide Compressed Wheel-

Pump Pump Air Knapsack barrow
Sprayer Sprayer Sprayer Sprayer Sprayer

Plunger Crank Drop Centrifugal Root

Duster Duster Spreader Spreader Irrigator

satisfactory, but for flowering shrubs–azaleas,

roses, etc.–a spray rod, curved at the end, or
with an angle nozzle, is easier on the plants and
more effective, allowing for better coverage of
underside of foliage.
The size of the hole in the nozzle disc and the
pressure determine the amount of spray used. The
volume of spray ejected per minute doubles or
triples with each small increase in the hole size
or pressure used. This means that in a home garden
where the objective is to cover a few rose bushes
effectively a large amount of spray can be wasted
at too high pressure, an expensive item with many
pesticide mixtures costing 20 to 30 cents a diluted
Figure 2 Spray application techniques gallon. Most chemicals are corrosive, and even if
Hand-Operated Sprayers 21

you start with a mist nozzle with a small hole at the

beginning of the season, you will soon be deliver-
ing more spray per minute because the hole is
enlarging. This usually means more conspicuous
residue left on the plant as well as more expense.

Hand-Operated Sprayers

“Aerosol bombs” are pressurized sprays in push-

button containers. A gas propellant reduced to
liquid form is added to a pesticide concentrate
and a fine mist is released when the button is
Figure 3 Pesticide application equipment (Modified
pushed. Unless the container is held 12 to 18 from the National Sprayer and Duster Association)
inches away from plants, to allow the gas to
evaporate, there will be some burning (more lit-
erally a freezing) when the liquid gas hits foliage. Slide or trombone sprayers have a telescoping
Such cans are good for house plants and for spot plunger, operated with two hands. They draw
treatment of insects outdoors, but air currents material from an attached jar or separate pail
make it difficult to place fungicides effectively. and discharge it as a continuous spray. They
Aerosols are also used for the application of develop good pressure and can be used for small
wound dressings to trees. (See Fig. 3). trees, but are tiring to use.
Household sprayers of the atomizer type are Wheelbarrow sprayers are manually operated
intermittent, discharging spray material with hydraulic sprayers, holding 7 to 18 gallons, that
each forward stroke of the pump, or continuous, are mounted on a frame with wheelbarrow-type
maintaining constant pressure. They are too small handles and one or two wheels. Pressures up to
and too tiresome to operate for more than a few 250 pounds may be developed, providing excel-
plants, and it is hard to get adequate coverage of lent coverage for shrubs and small trees. This
underside of foliage. type works best with two people, one to control
Compressed air sprayers are adequate for the pump, the other to operate the spray rod.
small gardens and are relatively inexpensive. Hose-end sprayers are attached to the garden
Capacity varies from 1 to 6 gallons. They are hose so that water supplies the pressure. The
meant to be carried slung over one shoulder, but action is that of a siphon. The concentrated
some come mounted on a cart. Air is compressed pesticide is placed in a jar, and as water under
into the tank above the spray liquid by a pressure is passed over the metering jet a small
hand-operated pump. A short hose, extension amount of chemical is drawn into it. This is a
rod, and adjustable nozzle make it possible very easy way to spray, and some models are
to cover undersurfaces. Such sprayers are a bit relatively accurate in materials discharged. Be
hard to pump up, and some models have sure to purchase a type with an extension tube
carbon dioxide cylinders to provide operating and deflector, so that spray can be directed to
pressure. underside of the foliage, with a shut-off at the
Knapsack sprayers, of 2 to 6 gallons capacity, jar, not just back at the hose, and with a device
are carried on the back of the operator and are to prevent back-siphonage. Hose-end sprayers
pumped by moving a lever up and down with the can be used for roses and other shrubs and
right hand as you spray with the left. These are for low trees. The droplets may be somewhat
more expensive than compressed air sprayers, but larger than those from a wheelbarrow or
deliver a fine continuous mist and are excellent knapsack sprayer, and slightly more chemical
for larger gardens. may be used.
22 Applying The Chemicals

to make a good formulation with relatively little

Dusters or no diluent. From a toxicity standpoint, it is
desirable to have a very small particle size, since
Pesticide dusts are most often made with talcs, immediate toxicity is generally inversely propor-
pyrophyllite, clays, calcium carbonate, precipi- tional to particle size. There are several important
tated hydrated silicates and silicon dioxides, syn- disadvantages to extremely small particle size:
thetic calcium silicate and diatomaceous earth as high wind losses, more or less rapid volatilization
the diluents although finely ground plant material and the prohibitive cost of extremely fine grind-
such as tobacco dust or walnut shell flour is ing. Also, to obtain better toxicant exposure of
sometimes used. technical concentrates absorbed on a carrier, it is
In some cases, a solution of the toxicant in a desirable to have the extender or diluent in as large
volatile organic solvent such as acetone or a particle size as possible and still give good
benzene is mixed with the dust diluents, the sol- dusting characteristics. In a 5 % dust effective
vent allowed to evaporate, and the mixture then toxicant exposure is obtained with the extender
ground. A solution of toxicant may be sprayed on averaging 10 times the size of the toxicant parti-
the dust diluent during mixing and grinding or the cles. At present, particle size specifications are
toxicant dissolved in a nonvolatile solvent and usually 10 to 30 mm for ground dusters and 20 to
mixed with the diluent. When this is done, care 40 mm for aircraft units. For use in fertilizer mix-
must be taken to avoid an excess of solvent that tures, granulated powders of 20 to 80 mesh are
might impair dusting qualities of the finished prepared by impregnation of Fuller’s earth and
product. Many technical pesticides in solid form bentonite fractions with the desired toxicants.
lend themselves to direct grinding with a sorptive Some dusts are sold in a can with a shaker top,
clay carrier in adequate milling equipment. Field meant to be applied like salt, which is certainly
strength dusts may be produced by diluting or not going to place a fungicide where it will do the
cutting down dust concentrates which contain most good. Some dusts are sold in small card-
from 10 to 50 % a.i. (Dust Bases). Because of their board cylinders to be used as dusters, which work
good dusting properties, attapulgites, diatomite, for a little while if the cardboard is well
talc, pyrophyllite, kaolins, and treated calcium paraffined to slide easily; but the dust soon gets
carbonate are used as diluents to provide the damp and clogs. Many more dusts are sold in
volume per acre needed to facilitate metering of plastic containers, with the dust supposedly com-
the dust through the duster mechanism. Since ing out in clouds as you squeeze, but more often it
many formulations contain more than one a.i., doesn’t after the first few days. Dusts are tricky to
dry concentrates must have the proper qualities use because of these disadvantages.
Spraying Vs Dusting

There is really no answer to the question of the next application a bit sooner. It is easier to
whether it is better to dust or to spray. In most spray than to dust on a windy day. Also, in dusting
gardens you will do both, depending on the you are somewhat more likely to get possibly
weather, the plant, the fungicide you want to toxic materials into your lungs than in spraying.
use, and how much time and help you have. The chief points in favor of dusting are the ease
Some orchardists prefer dusting because they and speed of application and the fact that you do
can get around the trees quickly in a rain, whereas not have to clean out the duster after each dusting.
to apply a spray they must wait until the foliage is Sprayers have to be cleaned, often between
dry. But for ornamentals exactly the opposite is different sprays, and they must be rinsed with at
true! You cannot dust a shrub even slightly wet least two changes of water pumped through the
with rain or dew without having a hideous system at the end of every day. Occasionally they
splotchy effect that persists for a long time. If must be taken apart, the tank soaked in trisodium
absolutely necessary you can spray while the phosphate or washing soda, the strainers and noz-
plants are still slightly wet, though the spray may zles in kerosene, wire run through the spray rods,
not stick quite as well, and you may want to make then all put together and rinsed with water.

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Mixing The Chemicals

It still seems incredible that so many gardeners scales. It also works out at about 1 level tablespoon
continue to treat their plants in a haphazard of the Dithane powder per gallon, and it is easier to
fashion. Buy a set of measuring spoons and measure than to weigh. There is, however, a good
a measuring cup, marked in ounces. Buy a large deal of volume variation, depending on how
pail and mark it off in gallons. Then measure, fluffed up the material is at the time you measure
exactly! it; so weighing is preferable.
Dosage directions are usually given in pounds When you buy chemicals in small packages
per 100 gallons of water, with or without transla- designed for the home garden, the dosages
tion on the label into small amounts. Not much given on the labels will probably be in terms
arithmetic is required to figure a smaller dosage, of tablespoons per gallon, and you need only
if you remember a few measurements: follow directions. When, to save a good deal of
Conversion table expense, you buy the larger sizes intended for
farmers, the directions may be given only in
3 teaspoons ¼ 1 tablespoon terms of pounds per 100 gallons. As a very
2 tablespoons ¼ 1 fluid ounce rough rule of thumb, you can figure 1 table-
16 tablespoons, 8 fluid ounces ¼ 1 cup
spoon per gallon where directions call for 1
16 ounces, 2 cups ¼ 1 pint
pound per 100 gallons, but the different mix-
2 pints, 4 cups ¼ 1 quart
tures have different weights so this is not very
16 cups, 8 pints ¼ 4 quarts ¼ 1 gallon
1 acre ¼ 43,560 square feet
At the rate of 1 pound to 100 you would use,
Suppose 3 gallons of a 2 to 100 dilution of lime accurately, 3/4 T captan 50 %, 1 T chloranil
sulfur is desired. That is the same as a 1 to 50 (Spergon), 1/3 T copper sulfate, 2/3 T dichlone
dilution. Three gallons constitute 48 cups; so if 1 50 % (Phygon), 1 1/4 T ferbam, 1/2 T maneb,
cup of liquid lime sulfur is added to 3 gallons, you 1 T spray lime, 3/4 T thiram, 1/2 T sulfur,
will have a 1 to 49 dilution, and that is close 2/3 T zineb (Dithane Z-78 or Parzate), 1 1/4 T
enough. ziram to 1 gallon of water.
Or suppose you want to make 4 gallons of Sometimes materials for soil treatment are
Zineb at the rate of 1 1/2 pounds per 100 gallons. given in pounds per acre. Knowing that one
That is 24 ounces per 100 gallons, or .24 ounce for acre contains 43,560 square feet, you can
1 gallon and .96 ounce for 4 gallons. That is make a proportion to find out how many pounds
approximately 1 ounce to weigh on your small are required per 1,000 square feet.

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All-Purpose Sprays and Dusts

The practicability of combination insecticide- other pests in check and also damage the envi-
fungicide mixtures is sometimes argued. The pro- ronment. DDT is no longer available. Some of the
prietary compounds are more expensive, but they new fungicides leave a rather conspicuous resi-
are more properly prepared than can be done at due; some are somewhat phytotoxic under certain
home and certainly save a lot of time. Nobody conditions. Some of these pesticides are no lon-
today could put on in separate applications all the ger available owing to new federal pesticide leg-
materials needed. The trouble is that the mixtures islation; however, new materials are available
follow fads, as in human medicine. Just as peni- that will replace those whose use is illegal.
cillin was given for most human ills some years Every mixture must be evaluated for particular
ago, so DDT was put in almost all pesticide climatic situations and kinds of plants. There are
mixtures, followed a little later by malathion. hundreds and hundreds of combinations on the
Both are excellent insecticides. The trouble is market under brand names. In order not to be out
they are somewhat too efficient, killing the para- of date before this text is printed, I have used as
sites and predators that keep mites and some few brand names as possible.

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Integrated Pest Management

Pesticides have been constantly scrutinized since • Use biological controls when available and
Rachel Carson’s book Silent Spring in the early when appropriate; and
1960’s and the birth of the U.S. Environmental • Use cultural practices which are favorable to
Protection Agency (EPA) in the early 1970’s. healthy plant growth.
Registrations of many pesticides have been can- A successful IPM program depends on four
celed and more will be canceled with the passage basic techniques.
of the Food Quality Protection Act (FQPA) in • Scouting. Regular and random visual
1996. The diminished availability of pesticides observations provide early warning to disease
may limit choices to more costly materials. In problems.
addition, there is growing concern about ground- • Disease Identification. The first and most
water contamination by pesticides and fertilizers, important step is to identify the problem;
consumer exposure to pesticide residue on food misdiagnosis results in use of improper
and plant material, pesticide resistance in plant control.
pathogens, insects and weeds, destruction of ben- • Timing. Improper timing of control measure
eficial organisms, atmospheric contamination by will result in disease control failure; the con-
pollutants, and concern for endangered species, trol measure must be timed correctly to the
all of which combine to make the problem of pest stage of disease development.
control more serious. • Records. Brief accurate records are a good
For the past 30 years integrated pest manage- tool for disease control decisions.
ment (IPM) has received increased interest. Although entomologists have achieved some
Investigations have concentrated on enhance- success with biological controls, the successes by
ment of a broad arsenal of integrated strategies plant pathologists with biological control has
for control of pests and diseases on selected com- been somewhat sparse. While use of classical
modities. A key goal of IPM strategies is the biological control has aided pest control, most
reduction of pesticide use to the absolute mini- biocontrol products have not yet proved to be
mum and the reliance on other strategies to assist preferred treatments for disease control. Intense
in controlling pests. IPM strategies which can be research in biological control of root diseases
used include: has been proceeding in the United States and
• Apply pesticides only when necessary; in Europe. Some microbial agents, although
• Make use of application methods that apply sometimes sensitive to environmental variation,
less pesticide or use a more efficient spray can be effective in controlling soil-borne plant
system; pathogens. Although there are many promising
• Use biocompatible chemicals as they become fungal and bacterial biocontrol agents, and exper-
available; iments demonstrate successful biocontrol in the

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_9, 29

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30 Integrated Pest Management

greenhouse and field, there are few commercially There have been searches in recent years for
available biocontrol products. The reasons may “natural” substances that may bear profound anti-
be due to: fungal/antibacterial properties and that exhibit
• An insufficient understanding of the mode of low mammalian and environmental toxicities.
action of most biocontrol agents; These chemicals are termed biocompatible and
• To need to develop mass production and there are four of interest:
delivery systems; • Neem from the neem tree (Melis azedarach)
• Little methodology for integrating biocontrol • Bicarbonates (used in baking)
with other control strategies and crop • Horticultural oils
production methods; and • Strobilurins (from fungal extracts)
• Competition of the biocontrol agent with other Some of these have now been formulated
microorganisms. for the commercial market and are exhibiting
It should also be recognized that biocontrol excellent disease control. There are numerous
products are effective against specific pathogens biocompatible chemicals under investigation for
and that the use of pesticides on foliage or soil their efficacy in disease and pest control. The
may have detrimental effects on the biocontrol great benefit of these products is their safety for
agent. the user and the environment.
Part II
Classification of Plant Pathogens

The plant diseases described in this handbook are caused by bacteria, fungi,
nematodes, a few seed plants (such as dodder, mistletoe, and witchweed),
viruses, physiological disturbances, and air and water pollutants. The classi-
fication of bacteria, fungi, and viruses is somewhat involved and is given here
as a background for the specific descriptions in ▶ Part III. There are many
classifications of bacteria, fungi, and viruses, with numerous disagreements
among mycologists, bacteriologists, virologists and plant pathologists.
Names and groups have been chosen that are widely accepted and most
readily adapted to the alphabetical requirements of a reference that works
like a dictionary.
Classification of the bacteria is based on that given in the Volume 1
(1984) and Volume 2 (1986) of Bergey’s Manual of Systematic Bacteriology,
and Laboratory Guide for Identification of Plant Pathogenic Bacteria
(1980) by N. W. Schaad (Editor). Classification of the viruses is based on
that given by Murphy et al. 1995. Virus Taxonomy, Classification, and
Nomenclature of Viruses. 6th Rept. ICTV, Brunt, A. A. et al. 1996. Virus of
Plants. Descriptions and List from the VIDE Database, Van Regenmortel
et al. 1999. Virus Taxonomy. 7th Report. ICTV and Descriptions of Plant
Viruses, published by the Commonwealth Mycological Institute and Associ-
ation of Applied Biologists. Other helpful sources were European Handbook
of Plant Diseases (1988) by I. M. Smith, J. Dunez, R. A. Lelliott, D. W.
Phillips and S. A. Archer and A Textbook of Plant Virus Diseases, 1972
edition, by Kenneth M. Smith.
So far as possible, the genera, orders, and families of fungi agree with
those given in Plant Pathogenic Fungi (1987) by J. A. von Arx. Helpful
sources included A Dictionary of the Fungi, 1961 edition, by G. C. Ainsworth
and G. R. Bisby, which includes G. W. Martin’s Key to the Families of
the Fungi; The Genera of Fungi, by F. E. Clements and C. L. Shear;
The Fungi, by Frederick A. and Frederick R. Wolf; The Lower Fungi:
Phycomycetes, by H. M. Fitzpatrick; Morphology and Taxonomy of Fungi,
by E. A. Bessey; and Illustrated Genera of Imperfect Fungi, by H. L. Barnett
and B. B. Hunter.
32 II Classification of Plant Pathogens

Classification in accordance with convention or law is called taxonomy.

Common names vary from locality to locality and country to country. Scien-
tific names are international and are based on the binomial system. Each kind
of bacterium, fungus, nematode, or higher plant is a species, and it has two
Latin words for its name. The first name indicates the genus to which the
species belongs, and the second the species itself. The latter name is usually
descriptive. Diplocarpon rosae means that Diplocarpon, the blackspot fun-
gus, is found on rose. Sometimes the species name honors a person, as
Coniothyrium wernsdorffiae for the fungus causing brand canker of rose.
Such a species name, derived from a proper name, has sometimes been
written with a capital, but present custom is to decapitalize all species
names. The names of genera should always be written with a capital.
Correctly, the author of the name should be written after the species. Then,
if someone else places the species in a new genus, the name of the first author
is put in parentheses followed by the name of the second author. When
a number of taxonomists have worked on a group, the list of authors gets
quite unwieldy. For simplicity all authors have been omitted from the scien-
tific names in this text. The correct name for a fungus with more than one
stage is that first given, with a valid description, for the teleomorph or sexual
stage. That rule is followed here with a few exceptions–as when a fungus is
almost universally recognized by another name.
Species are grouped into genera, related genera into families, designated
with the suffix aceae, as Erysiphaceae, and families into orders with the suffix
ales, as Erysiphales. Groups of related orders form classes.
Strange as it may seem, scientists are not yet agreed on what constitutes
a plant or even a living organism. The old definition of bacteria as unicellular
plants is disputed, and some question if fungi are truly plants. Bacteria are
prokaryotes. Prokaryotes are generally single-celled microorganisms that
have a cell membrane or a cell membrane and a cell wall surrounding the
cytoplasm and no organized nucleus. Eukaryotes contain membrane-bound
nuclei, mitochondria and – in plants only – chloroplasts. Although viruses are
known to multiply inside their hosts or vectors, the question of their being
a living entity has not been resolved. The arguments continue. Meanwhile
entities have to be grouped into some sort of order. Whittaker in 1969
introduced the five kingdom classification for all living organisms: Monera
(or Procaryota), Protista, Animalia, Plantae and Fungi (or Mycota). The
Monera are organisms with small cells lacking nuclei, mitochondria and
plastids, viz. the bacteria. The Protista include Microorganisms with one-
celled, often motile thalli (cells contain nuclei). The plants, animals and fungi
are believed to have evolved from Protista. The Fungi are characterized as
heterotrophic organisms, dependent on organic food, which they absorb. The
following scheme, adapted from the Plant Pathogenic Fungi, Nova Hedwigia
87: 288 pp. by J. A. von Arx, is an attempt to show the position of fungal plant
pathogens in the Kingdom Mycota. The listing of families is restricted to
those containing such pathogens.

Fungi are organisms having no chlorophyll, of fungi. The mycelium of these three phyla has
reproducing by sexual and asexual spores, not many nuclei which are not marked off by cross-
by fission like bacteria, and typically possessing wells (or nonseptate mycelium) except where
a mycelium or mass of interwoven threads reproductive structures arise, a condition known
(hyphae) containing well-marked nuclei. as coenocytic. Asexual reproduction is by means
According to Hawksworth (1991), there are of spores borne in sacs called sporangia. The
about 4,300 valid genera, and many more that Zygomycota have sexual spores called zygo-
are synonyms, and about 70,000 species living spores which are formed by the union of two
as parasites or saprophytes on other organisms or similar sex cells or gametes; the Oomycota have
their residues. More than 8,000 species cause sexual spores called oospores formed from dis-
plants disease. Fungi are divided into three king- similar gametes; the Chytriodiomycota have nei-
doms and eleven phyla. ther type of sexual spore; the Ascomycota have
Kingdom: Fungi septate mycelium and sexual spores in asci; the
Phylum: Chytridiomycota Basidiomycota have septate mycelium, fre-
Phylum: Zygomycota quently with clamp connections, and sexual
Phylum: Ascomycota spores; the Myxomycota have thalli as a motile
Phylum: Basidiomycota mass of protoplasm (a plasmodium or
Kingdom: Stramenopila myxamoeba – no mycelium) which is
Phylum: Oomycota transformed into a mass of small, aseptate
Phylum: Hyphochytriomycota resting spores that on germination form motile
Phylum: Labyrinthulomycota (slime molds) cells with or without flagella. The Myxomycota
Kingdom: Protists include protists with amoeboid thalli and their
Phylum: Plasmodiophoromycota (endoparasitic status as fungi often has been questioned. The
slime molds) thalli of the Myxomycota are naked, amoeboid,
Phylum: Dictyosteliomycota (Dictyostelid slime plasmotic masses without cell walls and are
molds) termed plasmodia or pseudoplasmodia. They are
Phylum: Acrasiomycota (Acrasid slime molds) also able to move by the formation of pseudopo-
Phylum: Myxomycota (true slime molds) dia and by plasma-streaming. The Plasmodio-
Oomycetes, Zygomycetes and phoromycetes is the only class of the
Chytridiomycetes were formerly listed as Myxomycota which includes parasites of vascu-
subclasses within the class Phycomycetes. lar plants. The best known species is
Oomycota, Zygomycota and Chytridiomycota Plasmodiophora brassicae, which causes “club
are now generally accepted as separate phyla root” of cabbage.

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34 Fungi

Saprophytes in water, most of which grow on
The thalli are usually vesicular, occasionally fil- submerged twigs and fruit; thallus of much-
amentous, and are transformed to sporangia, branched delicate hyphae.
gametangia or resting spores. The
Chytridiomycetes are the only members of the Plasmodiophorales
kingdom Fungi that produce motile cells. Motile
cells may function as zoospores, or as gametes, The placement of this order has always been
are radially symmetrical, with a single, posteri- uncertain. Some put it with the Myxomycetes,
orly directed whip-lash type flagellum. The the slime molds, others between the Myxomy-
Chytridiomycetes are the only class in this cetes and the true fungi. Some have considered
phylum. it a family in the Chytridiales. This single class is
placed in the kingdom, Protists, phylum,
Plasmodiophoromycota which contains a single
Chytridiales class, Plasmodiophoromycetes. There is also
only one family, Plasmodiophoraceae, in this
This order is defined on the basis of zoospore order. Parasitic, assimilative phase a
ultrastructure. Most members are water-or soil- multinucleate thallus within host cells, chiefly
inhabiting fungi; many of the former are parasitic of vascular plants, often causing hypertrophy;
on algae and water molds, many of the latter on germinating in place by amoeboid, occasionally
vascular plants. A few parasitize animal eggs and uniciliate, zoospores.
protozoa while others are saprobic on the Plasmodiophoraceae The only family in this
decaying remains of dead plants. Simple fungi order but with two important genera:
which have almost no mycelium, the thallus at Plasmodiophora, causing club root, and
maturity acting as a single sporangium, or divid- Spongospora, causing potato scab.
ing to become a sorus of sporangia; zoospores
posteriorly uniflagellate.

Spizellomycetales The thalli may be vesicular, often irregular, but

are usually filamentous. Sporangia on germina-
Members of this order are diverse and include tion release biflagellate zoospores. One flagellum
plant and fungal parasites and free-living is the whip-lash type and the other the tinsel type.
saprobes inhabiting both soil and water. There is Motile sex cells are absent. Sessile gametangial
great morphological variation in the group, and cells conjugate and form an oogonium containing
examples of both endogenous and exogenous one or several egg cells (see Fig. 1).
development. Most species are monocentric. The Oomycota are related to autotrophic algae
with similar characters. The Oomycetes are the
only class in this phylum; however, the small
Blastocladiales classes Hyphochytriomycetes and Labrinthu-
lomycetes may also be included.
Saprophytes in water or soil; genera are charac-
terized by thick-walled, resistant sporangia, usu- Hyphochytriales
ally with pitted walls. Another feature is the
prominent membrane-bound nuclear cap present Zoospores anteriorly uniflagellate, usually
in zoospores and planogametes. formed outside the sporangium. The order
Zygomycota 35



a1 c1

Fig. 1 Reproduction of an Oomycete (Phytophthora, oogonium. B asexual reproduction by sporangium (a)

order Peronosporales). A multinucleate oogonium (a) formed on sporangiophore (b); a1 sporangium germinat-
and male antheridium (b) in contact; fertilization tube ing by formation of ciliate zoospores; c1 zoospores ger-
formed between gametes after all nuclei except one has minating with germ tube
disintegrated; thick-walled oospore (c) formed inside

Hyphochytriales has now been placed in the phy- plants; in the latter case, zoosporangia function
lum Hyphochytriomycota which contains as conidia.
a single order, Hyphochytriales. Albuginaceae The white rusts. Conidia
(sporangia) in chains on club-shaped conidio-
Saprolegniales phores borne in dense sori beneath epidermis of
host, the sori forming white blisters; intercellular
Marine forms, parasites of diatoms and algae, or mycelium with globose haustoria.
in fresh water and soil, the water molds, with Pythiaceae Conidiophores differing little from
abundant mycelium; hyphae without constric- assimilative hyphae; mycelium saprophytic or
tions; oogonium with several oospores. parasitic, but if latter within cells and without
haustoria. Two genera, Phytophthora, which
includes the potato blight and other pathogens,
and Pythium, causing damping-off, are especially
Water forms; hyphae constricted, with cellulin
Peronosporaceae Downy mildews. Conidia are
plugs; oogonium with a single oospore.
borne singly or in clusters at tips of usually
branched, rarely clavate, conidiophores emerging
Lagenidiales through stomata; haustoria various.

Primarily aquatic, mostly parasitic on algae and

water molds; thallus simple; zoospores formed by Zygomycota
cleavage within sporangium or partly or wholly
in an evanescent external vesicle. The thalli are vesicular, or more often represent
a coenocytic, multinucleate mycelium (with
Peronosporales aseptate hyphae). The gametangial cells conju-
gate and form a thick-walled, persistent resting
Downy mildews and white rusts. Primarily ter- spore, called a zygospore (see Fig. 2). Motile sex
restrial, living in soil or parasitic on vascular cells are absent, but sporangiospores and conidia are
36 Fungi

A B a
a b

a1 e d1

Fig. 2 Reproduction of a Zygomycete (Rhyzopus, order formed inside a sporangium (a) formed on
Mucorales). A suspensors (a) from different hyphae cut a sporangiophore (b) around a columella (c). Hyphae are
off gametes (b) ofequalsize which fuse (b1) to form attached to substratum by rhizoids (e). Sporangiospore
a spinyzygospore (b2). B asexualsporangiospores (d) germinates by a germ tube (d1)

usually formed and dispersed by air. There are two as conidia; zoospores free within a gametangial
classes, the Zygomycetes, and the Trichomycetes vesicle.
(mainly parasitic on insects). These classes differ by
morphological and chemical characteristics.

The thalli may consist of aseptate yeast cells or

septate hyphae. Following meiosis, endogenous
spores (ascospores) form within a cell called an
ascus. There are three groups: Archiascomycetes
Profuse mycelium, much branched; asexual
(members lack ascogenous hyphae and ascocarps,
reproduction by sporangia or conidia; sexual
and asci sometimes homologized with sporangia),
reproduction by zygospores from union of two
Saccharomycetales (Ascomycetes, Yeasts: contain
branches of the same mycelium or from different
no ascogeneous hyphae and ascocarps; asci
mycelia. Some species damage fruits and vegeta-
thin walled and may release ascospores by deli-
bles in storage. Only two families are of much
quescing or breaking) and Filamentous Ascomy-
interest to plant pathologists.
cetes (with functional sex organs – possess
Mucoraceae Sporangiophores liberated by
ascogonium, ascogenous hyphae and crosiers that
breaking up of thin sporangial wall; zygospores
become enclosed in an ascocarp). The asci in Asco-
rough. Mucor and Rhizopus cause storage molds.
mycetes are aggregated in fructifications called
Choanephoraceae Both sporangia and conidia
ascomata (apothecia, cleistothecia, perithecia).
present, the latter borne on swollen tips; zygo-
The asexual states (anamorphs) of the Ascomycetes
spores naked. Choanephora is a weak parasite
usually are classified in a separate class called
causing blossom blight or blossom-end rot of
young fruits.

Entomophthorales Ascomycetes
Profuse mycelium, species frequently parasitic
on insects or other animals, rarely on plants; The diagnostic characteristics of this class are
anamorph spores modified sporangia functioning a septate mycelium (hyphae with cross walls)
Ascomycota 37



Fig. 3 Sexual reproduction in the Ascomycetes. A asci opening with a mouth or ostioles (Spaheriales).
borne singly in locules in stroma (Myriangiales). B peri- D Discomycetes (Heliotiales), ascus (a) and paraphyses
thecia with long necks or beaks immersed in stroma (a) formed in a hymenial layer in a cuplike apothecium (c)
(Sphaeriales). C papillate perithecium in host tissue, and (c1); ascospore (d) germinates by germ tube

and the ascus, a sac, typically club-shaped or Protomycetaceae Chlamydospores thick-

cylindrical, bearing the sexual spores, asco- walled, germinating after a rest period, the exo-
spores, usually eight in number. Asci may be spore splitting and the endospore emerging to
formed on or in hyphae or cells but are usually form a large multispored spore-sac. Parasitic on
grouped in structures, ascocarps, either in locules vascular plants.
in a stroma or lining a cup-shaped fruiting body Taphrinales Chlamydospores thin-walled; asci
called an apothecium or the walls of an enclosed eight-spored but may become multispored by
round or flaska-shaped perithecium. The young budding. Genera Exoascus and Taphrina cause
ascus has two nuclei, which fuse and then leaf curl and leaf blisters and now Exoascus is
undergo generally three divisions to give the usually considered a synonym of Taphrina.
eight spores. In many genera paraphyses, thin Subclass Euascomycetidae. Asci borne in
sterile clubs, are formed between the asci (see ascocarps.
Fig. 3).
Many ascomycetes have both a parasitic and Eurotiales
a saprophytic stage. In their parasitic stage they The order is characterized by (1) asci free on
usually produce conidia or anamorph spores, mycelium or within sessile or stipitate ascocarps;
sometimes on groups of conidiophores growing (2) sexual fusion, usually by trichogyne and
out of the mycelium, sometimes in a special pyc- undifferentiated hypha; (3) spherical-to-ovoid
nidium. Similar structures sometimes found are evanescent asci; (4) single-celled ascospores
spermagonia containing spermatia, small sex cells. that are oblate, spherical, globosa, or smooth or
with reticulations, spines, or thickened rings;
Taphrinales sometimes with appendages; (5) dry usually
This order is now placed in the class phialidic conidia; and (6) being known from
Archiascomycetes. Hyphae bearing terminal a variety of often starchy, oily, or cellulosic
chlamydospores or ascogenous cells, each of substrates.
which produces a single ascus, usually forming Trichocomaceae Diversity of coverings are
a continuous hymenium-like layer on often found over the asci: pseudoparenchymatous
modified tissues of hosts. Spore sac compound cleistothecia; stromata in which cleistothecia
(a synascus) regarded as equivalent of numerous develop; stromata without cleistothecia; or
asci. Parasitic on vascular plants. wefty hyphal coverings over the asci.
38 Fungi

Pseudeurotiaceae Species have coiled ascocarp branches, resembling perithecia. This family is
initials, hyaline, to dark cleistothecia, and now placed in the order Capnodiales.
scattered globose asci that are evanescent. Coryneliaceae Stroma lobed, each lobe with
a single locule which is finally wide open. Martin
Ouygenales places this in the Coryneliales.
This order characterized by asci free on myce- Dothideaceae Stroma not markedly lobed, loc-
lium; fusion of gametangial hyphae variable ules immersed in groups; at maturity stroma is
without trichogyne but with ascogonium some- erumpent and superficial.
times coiling; spherical to ovoid deliquescent Acrospermaceae Stroma typically uniloculate,
asci; single-celled ascospores variously shaped. clavate, erect; dehiscence by a fimbriate, often
Gymnoascaceae Cloistothecia around asci of spreading, tip. Martin places this family in the
loosely interwoven hyphae. Coryneliales and adds, under Dothideales,
Onygenaceae Ascocarp stalked and capitate, Pseudo-sphaeriaceae, with asci more or less sep-
small to medium; peridium tough, opening arated by stromatic tissue.
above; ascospores pitted. Microthyriaceae (including Asterineae and
Trichopelteae) Stromatic cover of radial or par-
Erysiphales (Perisporiales) allel hyphae; chiefly tropical species.
Parasites of higher plants; mycelium generally on
surface of host; perithecia without true ostioles.
The powdery mildews. White mycelium, with Microthyriales
conidia in chains; perithecia rupturing with an Mycelium largely superficial; stroma flattened;
apical tear or slit. dimidiate; opening by a pore or tear, simulating
the upper half of a perithecium.
Meliolales Polystomellaceae (including Stigmateae) Myce-
Dark or black mildews. Mycelium dark; stroma lium largely internal, forming a hypostroma;
unilocular, resembling a perithecium. Mycelium fruiting stroma subcuticular or superficial.
dark; asci exposed by gelatinization of upper Trichothyriaceae Superficial mycelium irregu-
portion of ascocarp. lar or lacking; base of stroma well developed;
parasitic on other fungi.
Stroma well developed, often gelatinous; asci Pleosporales
borne singly in locules. Nearly all are parasites Ostiole an elongated slit on a usually flattened,
on higher plants. elongate apothecium, bearing asci in a flat, basal
Piedraiaceae Tropical fungi invades cuticle of layer.
hair of primates, including humans. Hysteriaceae Ascocarps superficial from the
Myriangiaceae Stroma pulvinate, often with first; black, carbonaceous, round or elongate.
lobes, nearly homogeneous. Micropeltaceae (Hemisphaeriaceae) Internal
Elsinoaceae Stroma effused, with gelatinous mycelium scanty; stromatic cover not of radially
interior and crustose rind. arranged hyphae; chiefly tropical species. This
family now placed in order Pleosporales; for-
Dothideales merly in Microthyriales.
Mycelium immersed in substratum; stroma with
hard, dark rind, soft and pale within; locules more Hypocreales
or less spherical, resembling perithecial cavities. Perithecia, and stromata if present, bright col-
Capnodiaceae Sooty molds. Often on living ored, soft, and fleshy. Martin gives two families.
plants associated with insect secretions. Stroma Nectriaceae Asci elliptical to cylindrical;
massive, carbonaceous, often excessively inoperculate; ascospores various but never long-
branched; fruiting bodies borne singly at tips of filiform.
Ascomycota 39

Clavicipitaceae Asci long-cylindrical, with conidial stage; perithecia develop under bark;
a thickened tip, ascospores long-filiform. ascospores small, allantoid, hyaline to yellow-
brown. This family now placed in order
Sphaeriales (Pyrenomycetes) Xylariales.
Mycelium well developed; perithecia dark, more Melogrammataceae Conidia typically borne in
or less hard, carbonaceous, with an ostiole typi- hollow chambers in stroma composed of fungal
cally circular in section; with or without stro- elements; ascospores one-to many-celled, hya-
mata; asci inoperculate (without a lid) but line or brown.
spores discharged with force; paraphyses and Xylariaceae Conidia borne in superficial layer
periphyses usually present. on surface of stroma; ascospores one-to two-
Chaetomiaceae Perithecia superficial, hairy, celled, blackish brown.
walls membranous; asci deliquescent; ascospores Martin does not use the order Sphaeriales. He
dark; paraphyses wanting. Now placed in order places some of the above families in separate
Sordariales. orders. This family now placed in order Xylariales.
Sordariaceae (Fimetariaceae) Perithecia super-
ficial, walls membranous, naked or sparsely Laboulbeniales
setose; asci discharging spores forcibly. Now Minute parasites on insects or spiders; mycelium
placed in order Sordariales. represented by a small number of basal cells
Sphaeriaceae Perithecia superficial, walls car- functioning as haustorium and stalk.
bonaceous, mouths papillate.
Ceratostomataceae Perithecia superficial, car- Phacidiales (=Rhytismatales)
bonaceous, with long, hairlike beaks. Discomycetes in which the hymenium is covered
Cucurbitariaceae Stroma present but perithe- by a membrane until ascospores are mature, then
cia completely emergent at maturity; formed in splitting stellately or irregularly.
groups. Phacidiaceae Ascocarps leathery or carbona-
Amphisphaeriaceae Bases of perithecia persis- ceous, black, remaining embedded in host tissue
tently immersed in stroma; mouths circular. or in stroma; hypothecium thin. Martin includes
Lophiostomataceae Bases of perithecia persis- Tryblidiaceae, ascocarps leathery, immersed,
tently immersed in stroma; mouths compressed, hypothecium thick; but Ainsworth and Bisby
elongate. place members of this family in the Helotiales.
Sphaerellaceae (Mycosphaerellaceae) Perithe-
cia immersed in substratum; stroma lacking or Helotiales
poorly developed; asci not thickened at tips; Discomycetes without a membrane; asci
mouths of perithecia papillate. inoperculate, opening with a definite pore. Cup
Gnomoniaceae Perithecia immersed in substra- fungi.
tum; usually beaked; asci thickened at tips. This Geoglossaceae Ascocarps calvate or caplike,
family has been eliminated: Gnomonia species hymenium covering convex upper portion.
are in the order Dothideales. Ascocorticiaceae Fructification effused, inde-
Clypeosphaeriaceae Stroma a shieldlike crust terminate, without excipulum; paraphyses
(clypeus) over perithecia, through which necks lacking.
protrude. Stictidiaceae Ascocarps first immersed in sub-
Valsaceae Stroma composed of mixed host and stratum, then erumpent; asci long-cylindrical
fungal elements; perithecia immersed, with long with thickened apex; ascospores filiform, break-
necks; conidia borne in cavities in stroma. ing up into segments at maturity.
Melanconidiaceae Like Valsaceae but conidia Cyttariaceae Ascocarps compound, in form of
borne superficially on the stroma. subglobose stromata bearing numerous apothe-
Diatrypaceae Stroma composed wholly of fun- cial pits. Now placed in order Cyttariales; no
gus elements; in some genera present only in family.
40 Fungi

Patellariaceae Apothecia leathery, horny, carti-

laginous, or gelatinous; tips of paraphyses united Basidiomycota
to form an epithecium; asci thick-walled. Now
placed in order Patellariales; no family. The thalli may contain budding cells which are
Mollisiaceae Apothecia waxy or fleshy; perid- formed successively by new inner layers which
ium of rounded or angular, mostly thin-walled, burst through the outer layers. After meiosis, the
dark cells forming a pseudoparenchyma. haploid cells are formed exogenously by budding
Helotiaceae Apothecia soft, fleshy, stalked; and are called basidiospores or sporidia. Endog-
peridium of elongate, thin-walled, bright-colored enous spores (sporangiospores or ascospores) are
hyphae, arranged in parallel strands. absent in Basidiomycota.
Sclerotiniaceae Apothecia arising from The structures on which haploid spores
a definite sclerotium or stromatized portion of resulting from meiosis are formed are termed
the substratum; stalked, cup-shaped, funnel- basidia and usually bear a constant number of
form, or saucer-shaped; usually brown; asci spores, 2 or 4, occasionally more. The basidia
inoperculate, usually eight-spored; spores ellip- are differentiated on dikaryotic hyphae usually
soidal, often flattened on one side, usually hya- in or on fruiting bodies called basidiomata. The
line; spermatia globose to slightly ovate; conidial basidia may also be formed on resting spores
forms lacking in many genera. These families are called teliospores (see Fig. 4). Dikaryotic resting
from Martin’s 1954 Key to Families. His 1961 list spores may also germinate with a shorter or longer
puts Ostropaceae in the Ostropales and tube, which is termed promycelium. The three
Patellariaceae in the Hysteriales. Ainsworth and classes now distinguished are the Ustomycetes,
Bisby list Geoglossaceae and put all other genera the Urediniomycetes and the Basidiomycetes.
under “other Helotiales.” The Ustomycetes propagate mainly by bud-
ding cells; septate hyphae may be present, but
are rare. After meiosis, resting spores form
Pezizales short, often septate promycelia, which produce
Asci operculate, opening by a lid; hymenium budding cells laterally or terminally. Character-
exposed before maturity of spores; apothecia istic basidia or basidiospores are absent.
often brightly colored; most forms saprophytic. The Urediniomycetes form basidia, which
Pezizaceae Apothecia cup-shaped or discoid; after meiosis form uninucleate cells by transverse
sessile or stalked. septation. Each cell forms a single, stalked
Helvellaceae Fruit bodies upright, columnar or basidiospore.
with a stalk and cap; sometimes edible. Nearly all Uredinomycetes are obligate para-
sites of vascular plants and are known as rust fungi.
Tuberales The Basidiomycetes form basidia, which usu-
Ascocarp hypogeic, remaining closed; hyme- ally remain aseptate after meiosis; the basidio-
nium covered with a pseudo-tissue or hymenium spores are arranged in an apical whorl and are
lacking and asci filling cavities; mostly subterra- sessile or stalked. The septa of the hypha have
nean; includes edible truffles. characteristic central pores termed dolipores, with
Tuberaceae Interior waxy at maturity; asci per- thickened walls and caps. Dolipores are not present
sistent. This family now placed in order in the Ustomycetes and the Urediniomycetes.
Pezizales. The order Tuberales has been
Elaphomycetaceae Interior powdery at matu- Ustomycetes
rity; asci disappearing early, leaving interior
filled with spores. This family now placed in Ustomycetes include about 500 species belong-
order Pezizales. The order Tuberales has been ing to two orders; the plant parasitic Ustilaginales
eliminated. (smut fungi), and the Sporidiales (red yeasts).
Basidiomycota 41




Volva Pore

Fig. 4 Reproduction in Basidiomycetes. a, mushroom conk, in Polyporaceae where basidia line pores instead of
(Agaricaceae) with cap of pileus lined with gills bearing gills. Mycelium in basidiomycetes sometimes have
basidia germinating by basiodiospores. b, sporophore, or a structure around a septum called a clamp connection

Ustilaginales into four cells, each producing a single basidio-

The smuts. Spore masses are usually black; spores spore on a sterigma; spore masses are yellowish
are heavy-walled chlamydospores, germinating or orange, and there are several spore forms.
by a promycelium (basidium) and four or more Melampsoraceae Teliospores sessile, in crusts,
sporidia (basidiospores). cushions, or cylindrical masses, or solitary, or in
Ustilaginaceae Smuts. Basidiospores are pro- clusters, in mesophyll or epidermis of host. Now
duced on sides of a four-celled promycelium. placed in the order Melanosporales.
Tilletiaceae Smuts. Elongated basidiospores Pucciniaceae Teliospores usually stalked, sepa-
produced in a cluster at tip of a non-septate rate, or held together in gelatinous masses; some-
promycelium or basidium. times several on common stalks; less frequently
sessile, catenulate, breaking apart.
Auriculariaceae Basidia with transverse septa;
Urediniomycetes typically gelatinous. The genus Helicobasidium
causes violet root rot and the genus
Urediniomycetes – have cylindrical, often Herpobasidium causes blight of lilac. Now
slightly curved, transversely septate basidia. placed in the order Auriculariales.
Each cell forms a sterigma with a basidiospore, Septobasidiaceae (Felt fungus) Arid, lichenoid,
which is forcibly discharged when mature. Usu- parasitic on scale insects; probasidia often with
ally basidia develop on resting spores called thickened walls. Now placed in the order
teliospores. The Urediniomycetes contain two Septobasidiales. There are six other families, of
orders, the Uredinales (rust fungi, obligate para- no particular interest from the standpoint of plant
sites on vascular plants) and the Auriculariales. disease.

The rusts. More than 5,000 species have been Basidiomycetes
described in about 300 genera. Always parasitic
in vascular plants; teliospores or probasidia ger- Basidiomycetes – About 10,000 species have
minate with a promycelium divided transversely been described and includes the mushrooms and
42 Fungi

the bracket fungi formed on trees. Most grow in Hydnaceae Hymenium covering downward-
the soil and many form mycorrhiza with roots of directed spines, warts, or teeth. Now placed in
forest trees. The hyphae in general are septate and order Aphyllophorales.
dikaryotic. The septa of the hyphae often have Polyporaceae Hymenium lining pores (pits or
clamp connections, hyphal outgrowths formed dur- tubes); hymenophore woody, tough or membra-
ing cell division and forming a connection between nous, rarely subfleshy but never soft. Martin
two cells. The basidia are formed in or on places this family and the preceding three in
basidiomata on dikaryotic hyphae or on dikaryotic another order, Polyporales. Now placed in order
resting spores (teliospores). At maturity they are Aphyllophorales.
arranged either in a free, open layer termed hyme- Boletaceae Fruiting surface poroid or occasion-
nium or enclosed in fungal structures termed gleba. ally pitted; basidiocarp fleshy to tough or
The basidiospores are sessile or more often develop membranous.
on sterigmata. Young basidia are dikaryotic, until Agaricaceae The mushrooms. Fruiting bodies
the nuclei fuse and meiosis follows. The two, four usually fleshy, sometimes tough or membranous,
or more haploid nuclei migrate into the basidio- often with a stipe and cap; hymenophore lamel-
spores, which usually are uni-, occasionally binu- late, with gills.
cleate. Those orders containing plant parasitic
species are included below. Hymenogastrales
Hymenium present in early stages, lining cham-
Graphioliales bers of the gleba, closed fruiting body, which is
Graphiolaceae False smuts. Black, erumpent fleshy or waxy, sometimes slimy and fetid at
sori and spores in chains; on palms in warmer maturity.
Tremellales Gleba slimy and fetid; exposed at maturity on an
Trembling fungi Basidiocarp usually well elongated or enlarged receptacle.
developed, often gelatinous varying to waxy or
leathery hornlike when dry; mostly saprophytic, Lycoperdales
sometimes parasitic on mosses, vascular plants, The puffballs. Gleba powdery and dry at matu-
insects, or other fungi. rity; spores usually small, pale.

Agaricales Sclerodermatales
Hymenium (fruiting layer) present, exposed from Gleba powdery at maturity; chambers not sepa-
beginning or before spores are matured. rating from peridium or each other; spores usu-
Exobasidiaceae Hymenium on galls or ally large, dark.
hypertrophied tissues of hosts, which are vascular
plants. Martin places this in a separate order, Nidulariales
Exobasidiales. Bird’s nest fungi. Gleba waxy; chamber with
Thelephoraceae Hymenium smooth or some- distinct walls forming peridioles (the eggs in the
what roughened or corrugated; basidiocarp web- nest), which serve as propagules of
like or membranous, leathery or woody; dissemination.
hymenium on lower side. Now placed in order
Clavariaceae Hymenium smooth, pileus more Deuteromycetes: Fungi Imperfecti
or less clavate or club-shaped, erect, simple or (Mycelia Sterilia)
branched, fleshy or rarely gelatinous; hymenium
on all surfaces. Now placed in order Anamorph fungi are those for which
Aphyllophorales. a teleomorph state is not yet known or does not
Basidiomycota 43

b c2





Pycnidium Sporodochium Synnema

Fig. 5 Spore formation in the Deuteromycetes. a Tuberculariaceae; c2 dard conidiophores and conidia of
Sphaeropsidales, conidia in pycnidum. b Melanconiales, Dematiaceae; c3 hyaline conidia in chains, Moniliaceae;
conidia in acervulus. c Moniliales – c1 sporodochium of c4 conidiophores grouped into a synnema, Stilbaceae

b c d e
f g

1 2 2

Fig. 6 Spore forms in the Deuteromycetes, commonly c Phragmosporae, spores with two or more cross septa;
designated by letters and figures. a Amerosporae, one- c1 Hyalophragmiae, hyaline or light; c2 Phaeophragmiae,
celled; a1 Hyalosporae, spores hyaline; a2 Phaeosporae, dark. d Dictyosporae, muriform spores. e Scolecosporae,
spores dark. b Didymosporae, two-celled; b1 filiform spores. f Helicosporae, spirally coiled spores.
Hyalodidymae, hyaline; b2 Phaeodidymae, dark. g Staurosporae, starlike spores

exist. Most of them are in the Ascomycetes. The Sphaerioidaceae (Sphaeropsidaceae

groupings are based on conidia: hyaline or col- Phyllostictaceae) Pycnidia more or less globose,
ored; with one, two, or several cells; formed in ostiolate or closed; walls dark, tough, leathery or
pycnidia, on acervuli (little cushions of hyphae carbona ceous.
breaking through the host epidermis), or free on Nectrioidaceae As above but walls or stroma
the surface of the host (see Figs. 5 and 6). bright-colored, fleshy or waxy.
Leptostromataceae Pycnidia dimidate (having
Sphaeropsidales the outer wall covering only the top half); usually
Conidia borne in pycnidia or chambered cavities. radiate, sometimes long and cleft.
44 Fungi

Excipulaceae Pycnidia discoid or cupulate. Sporobolomycetaceae False yeasts. Reproduc-

tion by budding and germination by repetition;
Melanconiales probably anamorph species of the Tremellales, in
Conidia borne in definitely circumscribed acervuli; the Basidiomycetes.
erumpent (breaking through the substratum). Moniliaceae Hyphae and spores hyaline or
Melanconiaceae Conidia are slime-spores; brightly colored; conidiophores not grouped
cause anthracnose diseases. together.
Dematiaceae Same as Moniliaceae but hyphae
Moniliales or conidia, or both, brownish to black.
Conidiophores (specialized hyphae bearing Stilbaceae (Stilbellaceae) Conidiophores united
conidia) superficial, entirely free or bound in into a coremium or synnema, an upright group of
tufts or in cushionlike masses (sporodochia). hyphae.
Pseudosaccharomycetaceae (Cryptococcaceae) Tuberculariaceae Hyphae and conidiophores
False yeasts. Hyphae scanty or nearly lacking; repro- combined in a sporodochium, a tight, spore-
duction by budding but not germinating by repetition. bearing mass.

The fact that bacteria can cause plant diseases was Pathogenic bacteria apparently cannot enter
discovered almost simultaneoualy in four different plants directly through unbroken cuticle but get in
countries, with the United States claiming first through insect or other wounds, through stomata,
honors. In 1878 Professor T. J. Burrill of the through hydathodes, possibly through lenticels,
University of Illinois advanced the theory that fire and often through flower nectaries. They can sur-
blight of apple and pear was due to the bacteria that vive for some months in an inactive state in plant
he found constantly associated with blighted tissues. tissue, as in holdover cankers of fire blight, and
In 1879, the French scientist Prillieux published perhaps years in the soil, although claims for
a paper on bacteria as the cause of rose-red disease extreme longevity of the crown-gall organism in
of wheat; in 1880 the Italian Comes recognized soil are discounted.
bacteria as pathogenic to plants; in 1882 Burrill Most of these plant disease bacteria have had
named his fire-blight organism Micrococcus their genus names changed several times since
amylovorus; and in 1883 Walker in Holland reported they were first described, and some species have
the bacterial nature of yellows disease of hyacinth. been combined. Classification of bacteria will
It remained, however, for Erwin F. Smith, of the probably change further in future years. Where
U.S. Department of Agriculture, to do most of the genus and/or species names have been changed,
pioneer work in this field and to convince the world the old name is given in parentheses. The genera
that bacteria were to blame for so many diseases. He and species used in this text agree with those given
spent a lifetime in the process, starting with peach in Ninth Edition of Bergey’s Manual of Determi-
yellows, and going on to a study of crown gall and its native Bacteriology (1994) and recent articles
relation to human cancer. In 1905 the first volume of in the J. Systematic Bacteriology. Walter H.
his monumental work Bacteria in Relation to Plant Burkholder, of Cornell University, who revised
Diseases was published. the portions of the Manual dealing with plant
There are about 80 species of bacteria which pathogens, followed in the footsteps of Erwin F.
cause plant disease and many of them consisting Smith by spending his life with bacterial diseases
of numerous pathovars. Bacterial diseases fall of plants, as did Charlotte Elliott of the U.S.
into three categories: (1) a wilting, as in cucum- Department of Agriculture, from whose Manual
ber wilt, due to invasion of the vascular system, of Bacterial Plant Pathogens much information on
or water-conducting vessels; (2) necrotic blights, disease symptoms have been taken.
rots, and leaf spots, where the parenchyma tissue Two kinds of prokaryotes (organisms that lack
is killed, as in fire blight, delphinium black spot, a true nucleus) cause disease in plants. Bacteria have
soft rot of iris and other plants with rhizomes or a cell membrane, a rigid cell wall, and often one or
fleshy roots; (3) an overgrowth or hyperplasia, as more flagella. The mollicutes, or phytoplasmas lack
in crown gall or hairy root. a cell wall and have only a single-unit membrane.

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_11, 45

# Springer Science+Business Media Dordrecht 2013
46 Bacteria

A general classification of plant pathogenic Part III: Irregular, Gram-positive, nonsporing

prokaryotes is shown below: rods.
Kingdom: Prokaryotae – Organisms with genetic FAMILY: Mycobacteriaceae –The plant pathogens
material not organized into a nucleus that is formerly classified into the genus Corynebacte-
not surrounded by a membrane. rium were separated into the genera:
Bacteria: Have a cell membrane and cell wall. Clavibacter, Curtobacterium, Rhodococcus,
Anthrobacter, and Rathoyibacter. Bacteria in
Part I: Gram-negative aerobic rods and cocci. all genera are pleomorphic rods and form yel-
FAMILY: Pseudomonadaceae low colonies that are slow growing. Separation
GENUS: Pseudomonas, rod-shaped, one or several of the genera is based primarily on the presence
polar flagella, colonies white. of specific amino acids in the cell walls. Posi-
Xanthomonas, rod-shaped, one polar flagel- tive identification of the genera is very difficult.
lum, colonies yellow. GENUS: Clavibacter. Contains important phyto-
Xylella, rod-shaped, under some cultural con- pathogenic bacteria formerly classified as
ditions filamentous; nonmotile, aflagellate, Corynebacterium.
Burkholderia, rod-shaped bacteria, motile by Part IV: Actinomycetes, bacteria forming
one or more polar flagella, colonies white, branching filaments.
nutrition very versatile. FAMILY: Streptomycetaceae
Acidovorax, rod-shaped bacteria, motile by sin- GENUS: Streptomyces. Gram-positive, aerial myce-
gle polar flagellum, colonies white, limited lium with chains of nonmotile condidia.
number of sugars are used for growth.
Rhizomonas, small rod-shaped bacteria, Part V: Mollicutes, prokaryotes that have a cell
motile by single polar flagellum, colonies membrane but no cell wall.
slow growing and white, causes corky root FAMILY: Mycoplasmataceae, the plant
of lettuce. mycoplasmalike organismsphytoplasmas.
Rhizobacter, rod-shaped bacterium with one Spiroplasmataceae.
lateral or polar flagellum, colonies white to GENUS: Spiroplasma, helical, motile but lacking
yellowish-white depending upon the flagella. Phytoplasma, pleomorphic cells not
medium, causes carrot bacterial gall. culturable in artificial medium.
Ralstonia, rod-shaped bacterium with single The taxonomy of the plant pathogenic fastid-
polar flagellum, colonies white and usually ious phloem-limited bacteria is still unknown,
pleomorphic in shape, causes bacterial wilt and the taxonomy of the phytoplasmas, and
in over 200 species of plants. of the spiroplasmas is tentative. Furthermore,
Xylophilus, rod-shaped bacterium with single Richettsialike organisms (RLOs) have been
polar flagellum, colonies yellow and slow- reported to be associated with a number of
growing, causes cankers on grape vines. plant diseases. RLOs are also cultured with diffi-
FAMILY: Rhizobiaceae culty which is a characteristic similar to the
GENUS: Agrobacterium, rod-shaped sparse lateral phytoplasmas. On this basis, both phytoplasmas
flagella, colonies white, rarely yellow. and RLOs are referred to as ‘fastidious prokary-
otes’. There are more than 200 distinct plant
Part II: Gram-negative facultative anaerobic diseases affecting several hundred genera of
rods. plants which have been shown to be caused by
FAMILY: Enterobacteriaceae the Mollicutes.
GENUS: Erwinia, peritrichous flagella, colonies The taxonomic scheme for mollicutes and
white or yellow. phytoplasmas is difficult to present in this hand-
Pantoea, peritrichous flagella, colonies yel- book since morphological criteria are limited and
low. Nutritionally restricted. both the criteria used in bacteriology and the
Bacteria 47

serological methods used in virology are difficult other cases identification must be left to the
to apply because phytoplasmas (except for technically trained bacteriologist. It involves
spiroplasma) have not been cultured. Thus the special staining technique, for examination of
true nature of phytoplasmas and RLOs, and form and motility under the microscope, and to
their taxonomic position among microorganisms see whether it is Gram-negative or Gram-posi-
is uncertain. In practice, the diseases caused by tive, and special culture technique to determine
mollicutes have been taxonomically been treated shape, color, and texture of colonies on agar and
individually. The elucidation of true relatedness gelatin, production of gases, fermentation of
among these organisms awaits further research. sugar, coagulation of milk, etc. If you are in
The general nature of the symptoms and the doubt about a plant disease, and the absence of
name of the host plant will, in many cases, leave fungus fruiting bodies leads you to believe that
little doubt as to the identity of a bacterial dis- bacteria may be at work, send a specimen to
ease. In the case of the soft rot due to Erwinia your State Experiment Station for expert
carotovora the nose alone is a reliable guide. In diagnosis.
Viruses, Viroids, Phytoplasmas

Viruses viruses contain deoxyribonucleic acid (DNA)

instead of RNA. X-ray diffraction and electron
The word virus means poison or venom. When it microscopy have shown something of the mor-
is used in connection with a plant disease, it phology of virus particles. Some are rods, some
means a filterable virus, an infective principle or filiform, and some are isometric, but polyhedral
etiological agent so small it passes through filters rather than spherical. They apparently act not as
that will retain bacteria. Virus diseases in man organisms but as disturbances in the host metab-
range from infantile paralysis to the common olism of nucleic acid.
cold and in plants from “breaking” of tulip There are over 850 described plant virus spe-
flowers to the deadly raspberry ringspot disease cies. Many of the described viruses are definitive
on the Malling Jewel variety of raspberry. members of genera, whose names have been
Viruses are obligate parasites in that they are approved by the ICTV (International Committee
capable of increasing only in living cells. They on Taxonomy of Viruses) to be viruses and could
are not organisms in the usual sense because they be agents of other kinds. For example, aster yel-
do not multiply by growth and fission, and they lows and elm phloem necrosis were thought for
are too complex to be chemical molecules. F. C. some time to be caused by viruses, but have now
Bawden, in the 1964 edition of his Plant Viruses besen determined to be caused by phytoplasmas.
and Virus Diseases, defines viruses as “submi- Moreover, in Part 4 some phytoplasma may still
croscopic infective entities that multiply only be positioned under bacterial or viral caused dis-
intracellularly and are potentially pathogenic.” ease since their true identity is not yet known or
Virus diseases are old; our knowledge of them that the confirmation of identity has been made
is relatively recent. Tulip mosaic, shown as but missed for inclusion in the 7th edition. In
breaking of flower color, was described in addition, potato spindle tuber and chrysanthe-
a book published in 1576. In 1892 it was shown mum stunt disease were long thought to be caused
that the cause of tobacco mosaic could pass by viruses, but have now been determined to
through a bacteria-proof filter, and in 1935 be caused by viroids. Viroids consist solely of
a crystalline protein was prepared from tobacco small RNAs with no protein coat. There are now
mosaic virus juice. At present we believe that about 30 plant diseases that have been identified
virus particles contain only two major compo- as having viroid causal agents including potato
nents, nucleic acid embedded in a protein struc- spindle tuber, chrysanthemum stunt, citrus
ture, and that they are built of uniform-sized exocortis, chrysanthemum chlorotic mottle, and
subunits arranged in a fixed and regular manner. cadangcadang of palm. More diseases caused by
Many plant viruses contain ribonucleic acid viroids will probably be identified in future years.
(RNA). Some plant viruses and many animal There are now about 68 identified phytoplasmas

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_12, 49

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50 Viruses, Viroids, Phytoplasmas

and finally, some plant diseases formerly thought the growing season and found free of
to be caused by viruses have now been deter- certain diseases. Virus-free foundation stock
mined to be caused by spiroplasma, such as citrus can be built up from heat treatment –rowing
stubborn disease. Thus the field of virology has plants at high temperatures for weeks or even
changed somewhat in recent years. In order to months – and/or meristem tip cultured plants.
simplify the discussion of these viruses and virus- Virus-free stock is tested by “indexing”, bioas-
like agents and the diseases they cause, these says and/or serological assays, before using stock
agents are grouped under virus diseases, since for propagating. Controlling insect vectors (by
the symptoms which they cause in plants are spraying plants or treating soil with systemic
similar. insecticides), eliminating weed hosts, roguing
Some viruses attack a large number of differ- diseased plants before insects can transmit the
ent plants and are of great economic importance; virus, and using resistant varieties are all ways
others are confined to a single host. Virus symp- of combating virus diseases.
toms fall into several categories, but commonly This handbook does not deal predominantly
there is loss of color due to the suppression of with the characteristics of the causal viral agent,
chlorophyll development. Foliage may be mot- but with the disease caused by the virus, viroids
tled green and yellow, mosaic, or have yellow and phytoplasmas.
rings (ring spot); or there may be a rather uniform Viruses are classified now in the traditional
yellowing (yellows). Stunting is common. The taxonomic system (family – genus – species).
reduction in manufactured food from the chloro- Formal use of a virus species name should be
phyll loss leads to smaller size, shorter inter- printed in italics with the first word capitalized;
nodes, smaller leaves and blossoms, and an acronym, when used, should also be capital-
reduced yield. There may be various distortions ized. In this book however, we used bold font for
of leaves and flowers, witches’ brooms, or the generic name of viruses. Generally, the spe-
rosettes. There may be necrotic symptoms with cies name consists of the vernacular plus the
death as the end result, and sometimes symptoms generic names. For example, Tomato spotted
are “masked,” not showing up under certain con- wilt tospovirus is the species name, tomato
ditions, such as hot weather, or latent, not spotted wilt virus is the vernacular name and
appearing until another virus is also present. Tospovirus the generic name. The scheme of
Viruses are transmitted from plant to plant by: virus and viroid classification (shown below)
insects, mites, fungi, and nematodes; rubbing, according to Murphy et al. (1995) and Brunt
abrasion, or other mechanical means (sometimes et al. (1996) include the following data: genus
handling tobacco and merely touching a healthy of virus, family (if designated), kind of nucleic
plant spreads mosaic); grafting or propagation by acid in genome, shape of virions, presence of
cuttings and bulbs; occasionally seeds; some- envelope and the type species:
times soil and water; and dodder, parasitic vines
whose tendrils link one plant to another. About
half of the insect vectors are aphids; a third are
leafhoppers. Mealybugs and whiteflies transmit Alfamovirus
some viruses, and six, including tomato spotted
wilt, are transmitted by thrips. In some cases the Bromoviridae; (+)ssRNA; isometric particles;
virus multiples within the insect as well as in the not enveloped; alfalfa mosaic virus.
plant. Some viruses have many different vectors,
50 being recorded for onion yellow dwarf, and
some have but a single known vector. Alphacryptovirus
Control of virus diseases starts with obtaining
healthy seed, cuttings, or plants. “Certified” Partitiviridae; dsRNA; isometric particles; not
means that plants have been inspected during enveloped; white clover cryptic virus 1.
Fabavirus 51

Badnavirus Caulimovirus

dsDNA; bacilliform particles; not enveloped; dsDNA; filamentous particles; not enveloped;
commelina yellow mottle virus. cauliflower mosaic virus.

Betacryptovirus Closterovirus

Partitiviridae; dsRNA;isomet-ric particles; not (+)ssRNA; filamentous particles; not enveloped;

enveloped; white clover cryptic virus 2. beet yellows virus.

Bigeminivirus Comovirus

Geminiviridae; ssDNA; isometric particles; not Comoviridae; (+)ssRNA; isometric particles; not
enveloped; bean golden mosaic virus. enveloped; cowpea mosaic virus.

Bromoviridae; (+)ssRNA; isometric particles;
Bromoviridae; (+)ssRNA; isometric particles;
not enveloped; cucumber mosaic virus.
not enveloped; brome mosaic virus.

Rhabdoviridae; Mononegavirales; ( )ssRNA;
Potyviridae; (+)ssRNA; filamentous particles; bacilliform particles; enveloped; lettuce necrotic
not enveloped; barley yellow mosaic virus. yellows virus.

Capillovirus Dianthovirus
(+)ssRNA; filamentous particles; not enveloped; (+)ssRNA; isometric particles; not enveloped;
apple stem grooving virus. carnation ringspot virus.

Carlavirus Enamovirus

(+)ssRNA; filamentous particles; not enveloped; (+)ssRNA; isometric particles; not enveloped;
carnation latent virus. pea enation mosaic virus.

Carmovirus Fabavirus

Tombusviridae; (+)ssRNA; isometric particles; Comoviridae; (+)ssRNA; isometric particles; not

not enveloped; carnation mottle virus. enveloped; broad bean wilt virus 1.
52 Viruses, Viroids, Phytoplasmas

Fijivirus Machlomovirus

Reoviridae; dsRNA; isometric particles; not (+)ssRNA; isometric particles; not enveloped;
enveloped; Fiji disease virus. maize chlorotic mottle virus.

Furovirus Macluravirus

(+)ssRNA;rod-shaped particles; not enveloped; (+)ssRNA; filamentous particles; not enveloped;

soil borne wheat mosaic virus. maclura mosaic virus.

Hordeivirus Marafivirus

(+)ssRNA;rod-shaped particles; not enveloped; (+)ssRNA; isometric particles; not enveloped;

barley stripe mosaic virus. maize rayado fino virus.

Hybrigeminivirus Monogeminivirus

Geminiviridae; ssDNA; isometric particles; not Geminiviridae; ssDNA; isometric particles; not
enveloped; beet curly top virus. enveloped; maize streak virus.

ssDNA; small isometric particles; not enveloped;
(+)ssRNA; isometric particles; not enveloped;
subterranean clover stunt virus.
raspberry bushy dwarf virus.

(+)ssRNA; isometric particles; not enveloped;
Bromoviridae; (+)ssRNA; isometric particles; tobacco necrosis virus.
not enveloped; tobacco streak virus.

Comoviridae; (+)ssRNA; isometric particles; not
Potyviridae; (+)ssRNA; filamentous particles; enveloped; tobacco ringspot virus.
not enveloped; sweet potato mild mottle virus.

Rhabdoviridae; Mononegavirales; ( )ssRNA;
(+)ssRNA; isometric particles; not enveloped; bacilliform particles; enveloped; potato yellow
barley yellow dwarf virus. dwarf virus.
Tymovirus 53

Oryzavirus Sobemovirus

Reoviridae; dsRNA; isometric particles; not (+)ssRNA; isometric particles; not enveloped;
enveloped; rice ragged stunt virus. southern bean mosaic virus.

Ourmiavirus Tenuivirus
(+)ssRNA; bacilliform particles; not enveloped; (+/ )ssRNA; thin filamentous particles; not
ourmia melon virus. enveloped; rice stripe virus.

Phytoreovirus Tobamovirus
Reoviridae; dsRNA; isometric particles; not (+)ssRNA;rod-shaped particles; not enveloped;
enveloped; wound tumor virus. tobacco mosaic virus.

(+)ssRNA; filamentous particles; not enveloped;
(+)ssRNA;rod-shaped particles; not enveloped;
potato virus X.
tobacco rattle virus.

Potyviridae; (+)ssRNA; filamentous particles;
not enveloped; potato virus Y. Tombusviridae; (+)ssRNA; isometric particles;
not enveloped; tomato bushy stunt virus.

Potyviridae; (+)ssRNA; filamentous particles;
not enveloped; ryegrass mosaic virus. Bunyaviridae; ( )ssRNA; large isometric parti-
cles; enveloped; tomato spotted wilt virus.

ssRNA or DNA; the satellite viruses depended on
helper viruses, but produce their own virions – small (+)ssRNA; filamentous particles; not enveloped;
isometric; not enveloped; tobacco necrosis virus. apple chlorotic leaf spot virus.

Sequivirus Tymovirus

Sequiviridae; (+)ssRNA; isometric particles; not (+)ssRNA; isometric particles; not enveloped;
enveloped; parsnip yellow fleck virus. turnip yellow mosaic virus.
54 Viruses, Viroids, Phytoplasmas

Umbravirus Phytoplasma

(+)ssRNA; isometric particles; enveloped; carrot Classification, presented below, based on restric-
mottle virus. tion fragment length polymorphism or putative
restriction site analysis of 16s rRNA gene
sequences according to Davis and Sinclair
Varicosavirus Aster yellows group: tomato big bud, Michi-
gan aster yellows, clover phyllody, Paulownia
dsRNA; rod-shaped particles; not enveloped; let- witches’-broom, blueberry stunt, apricot chlo-
tuce big-vein virus. rotic leafroll, strawberry multiplier.
Peanut witches’-broom group: peanut witch’s-
broom, witches’-broom of lime – “Candidatus
Phytoplasma aurantifolia”, faba bean phyllody,
Waikavirus sweet potato little leaf.
X – disease group: X – disease, clover yellow
Sequiviridae; (+)ssRNA; isometric particles; not edge, pecan bunch, goldenrod yellows, Spirea
enveloped; rice tungro spherical virus. stunt, milkweed yellows, walnut witches’-broom,
poinsettia branch-inducing, Virginia grapevine
Coconut lethal yellows group: coconut lethal
Viroids yellowing, Tanzanian coconut lethal decline.
Elm yellows group: elm yellows, cherry lethal
Unencapsidated, small circular ssRNAs. Viroid yellows, flavescence doree. Clover proliferation
replication parasitizes plant host transcription. group: clover proliferation, “Multicipita” phyto-
Known described viroids are: apple dimple fruit plasma. Ash yellows group: ash yellows.
viroid, apple scar skin viroid, Australian grape- Loofah witches’-broom group: loofah
vine viroid,. avocado sunblotch viroid, chrysan- witches’-broom.
themum chlorotic mottle viroid, chrysanthemum Pigeon pea witches’-broom group: pigeon pea
stunt viroid, citrus bent leaf viroid, citrus witches’-broom.
exocortis viroid, citrus III viroid, citrus IV viroid, Apple proliferation group: apple proliferation,
Coleus blumei 1 viroid, Coleus blumei 2 viroid, apricot chlorotic leafroll, pear decline, Spartium
Coleus blumei 3 viroid, Columnea latent viroid, witches’-broom, black alder witches’-broom.
coconut cadang-cadang viroid, coconut tinangaja Rice yellow dwarf group: rice yellow dwarf,
viroid, grapevine yellow speckle 1 viroid, grape- sugarcane white leaf, leafhop-per-borne.
vine yellow speckle 2 viroid, hop latent viroid, Stolbur group: stolbur phytoplasma, Austra-
hop stunt viroid, Iresine viroid 1, Mexicana lian grapevine yellows – “Candidatus Phyto-
papita viroid, peach latent mosaic viroid, pear plasma australiense”.
blister canker viroid, potato spindle tuber viroid, Mexican periwinkle virescence group: Mexi-
tomato apical stunt viroid, tomato planta macho can periwinkle virescence. Bermudagrass white
viroid. leaf group: Bermudagrass white leaf.

In the more than five decades since the first FAMILY: Belonolaimidae
edition of this book was prepared, nematodes GENUS: Belonolaimus, sting nematode on wide
have become of major importance in plant variety of plants.
pathology. Several hundred species are known Tylenchorhynchus, stunt nematode tobacco, corn
to cause plant disease. All plant parasitic nema-
todes are in the animal kingdom and belong to the
phylum Nematoda. Some examples are given FAMILY: Heteroderidae
after each genus. GENUS: Globodera, cyst nematode of potato.
PHYLUM: Nematode Heterodera, cyst nematode on wide variety of
ORDER: Tylenchida plants.
SUBORDER: Tylenchina Meloidogyne, root-knot nematode on wide
SUPERFAMILY: Tylenchoidea variety of plants.
FAMILY: Anguinidae
GENUS: Anguina, seed gall nematode.
Ditylenchus, stem or bulb nematode of onion, FAMILY: Tylenchidae
narcissus. GENUS: Tylenchus
FAMILY: Pratylenchidae SUPERFAMILY: Criconematoidea
GENUS: Pratylenchus, lesion nematode of nearly FAMILY: Criconematidea
all plants. GENUS: Criconemella, ring nematode of citrus,
Radopholus, burrowing nematode of citrus. fig, zoysia.
Naccobus, false root-knot nematode of garden Hemicycliophora, sheath nematode of beet, bean,
beets, cacti, crucifers, lettuce. blueberry, dracaena.
FAMILY: Hoplolaimidae
GENUS: Hoplolaimus, lance nematode of corn,
turf grass, carnation. FAMILY: Tylenchulidae
Rotylenchus, spiral nematode of turf grass, GENUS: Tylenchulus, citrus nematode of citrus,
tomato, gardenia. grapes, lilac.
Helicotylenchus, spiral nematode of turf grass, Paratylenchus, pin nematode of carnation,
gardenia, azalea, apple, grape. celery, fig.
Rotylenchulus, reniform nematode of turf grass,
tomato, gardenia.
FAMILY: Dolichodoridae ORDER: Aphelenchida
GENUS: Dolichodorns, and nematode of turfgrass. SUBORDER: Aphelenchina

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56 Nematodes

SUPERFAMILY: Aphelenchoidea ORDER: Triplonchida

FAMILY: Aphelenchoididae FAMILY: Trichodoridae
GENUS: Aphelenchoides, foliar nematode of GENUS: Paratrichodorus, stubby root nematode
chrysanthenum, strawberry, lilly, begonia. of apple, vegetables.
Bursaphelenchus, pine wood nematode. Trichodorus, stubby root nematode of vegeta-
Rhadinaphelenchus, coconut red ring nematode. bles, turf grass, dahlia, azalea.

ORDER: Dorylaimida
FAMILY: Longidoridae
GENUS: Longidorus, needle nematode of grape,
celery, leek, lettuce, parsley.
Xiphinema, dagger nematode of rose, trees, many
Part III
Plant Diseases and Their Pathogens

Because this is a reference book and not one to be read for pleasure or
continuity, most of you will come to the material you need in this section
by way of the index or the lists of diseases given under the different hosts in
▶ Part IV. At the beginning of ▶ Part IV you will find a list of headings
under which diseases are grouped and described, from Anthracnose to
Witchweed. In the Host section, ▶ Part IV, the key word, for example, rot
or blight, is given in capital and small capitals, followed by the name of
the pathogen (agent causing disease) in boldface. In this Diseases section,
Chap. 3, the pathogens are listed in boldface in alphabetical order under
each heading such as ROTS or BLIGHTS and so on, followed by the common
name of the disease. This system was adopted for quick and easy reference
because trying to alphabetize hundreds of similar common names would lead
to endless confusion. Also, it allows a very brief summary of the classification
and diagnostic characters of each genus before going on to a consideration
of diseases caused by the various species. This brief summary is in small
type, so that it can be readily skipped by readers uninterested in the technical
details. Perhaps I am the only one who feels the need for this quick review, to be
used in conjunction with the classification given in ▶ Part II; perhaps others
who have to answer questions over a broad field instead of their own specialty
can make use of these capsules sandwiched in between nontechnical
An alphabetical arrangement has the great disadvantage of being thrown
out of alignment every time the name of a fungus is changed, as it so
frequently is. In some such cases the old name is retained to avoid change
in order, but the present accepted name is also given. Sometimes names have
been changed under several hosts and the old name inadvertently retained
under others. And sometimes the old name is purposely retained because it is
so familiar to everyone. This is particularly true of a few fungi far better
known by their anamorph states than by the correct name of the teleomorph
state. A fungus not only can have several names; it also can cause more than
one type of disease. For instance, Pellicularia filamentosa is the present name
of the fungus formerly known as Corticium vagum when causing Rhizoctonia
rot of potatoes and Corticium microsclerotia when causing web blight of
beans. As Rhizoctonia solani, the name given to the sclerotial stage, the same
58 III Plant Diseases and Their Pathogens

fungus causes damping-off of seedlings, root rots of many plants, and brown
patch of lawn grasses. There are lots of plant diseases, and there are lots of
fungi causing them, but there are not nearly as many separate pathogenic
organisms as all the names would indicate. Thus, a Linkage Reference guides
the user to two or more common disease sites, i.e. “Canker” or “Blight”, where
the user searches for the pathogen alphabetically or the link may guide the user
directly to a pathogen in another chapter or section.
I cannot think of anything more deadly than ploughing straight through
this section from Anthracnose to Wilts. By doctor’s orders, take it in small
doses, as needed. But do read the few introductory remarks as you look up
each group, and please, please, before starting any control measures, read the
opening remarks in ▶ Part I on Garden Chemicals, and look up, in the list of
chemicals, any material you propose to use, noting precautions to be taken
along the lines of compatibility, weather relations, and phytotoxicity.
Although the disease descriptions, fungus life cycles, and general principles
of control given here will remain fairly valid, it must be stressed that
chemicals suggested for control are constantly changing. Today’s discovery
may be obsolete tomorrow. This Plant Disease Handbook should, therefore,
be used in conjunction with the latest advice from your own county agent or
experiment station. Addresses of the state agricultural experiment stations are
given following ▶ Part IV.

The term “anthracnose” has been used for two Sphaceloma, and are treated, in this revised text,
distinct types of disease, one characterized by as a separate group. ▶ Spot Anthracnose.
a typical necrotic spot, a lesion of dead tissue, Anthracnose in the modern sense is a disease
and the other by some hyperplastic symptom, characterized by distinctive limited lesions on
such as a raised border around a more or less stem, leaf, or fruit, often accompanied by dieback
depressed center. The word was coined in France and usually caused by a Gloeosporium or
for the latter type, to differentiate a grape disease a Colletotrichum, anamorph fungi producing
from a smut of cereals, both of which were called slime spores oozing out of fruiting bodies
charbon. The new word was taken from the (acervuli) in wet, pinkish pustules. These spores
Greek Anthrax (carbuncle) and nosos (disease), (conidia) on germinating form an appressorium
and was first used for the grape disease, caused by (organ of attachment) before entering the host
Sphaceloma ampelina, the chief symptom of plant. The teleomorph state of the fungus, when
which was a bird’s-eye spot with a raised border. known, is Gnomonia or Glomerella (see Fig. 1).
A disease of brambles, raspberry and black-
berry, was then named anthracnose because it
looked like the grape disease. The fungus, how- Apiognomonia
ever, instead of being correctly placed in the
genus Sphaceloma, was mistakenly named Apiognomonia errabunda (Anamorph,
Gloeosporium venetum. The next disease enter- Gloeosporium quercinum). Oak Anthracnose.
ing the picture was a bean trouble, and, because See ▶Discula umbrinella and Fig. 2.
the fungus was identified as Gloeosporium Apiognomonia errabunda (formerly
(though later transferred to the genus Gnomonia quercina). Oak Anthracnose. The
Colletotrichum), this common bean disease with fungus is closely related to Gnomonia platani,
typical necrotic symptoms was also called usually reported as G. veneta, but is now consid-
anthracnose and came to typify diseases so ered a separate species. The anthracnose appears
designated. as brown areas adjacent to midribs and lateral
The term “spot anthracnose” has been given to veins.
those diseases similar to the original hyperplastic Apiognomonia tiliae (formerly Gnomonia
grape disease. Those with slight hyperplastic tiliae). Linden Anthracnose, Leaf Spot, Leaf
symptoms are still commonly called anthracnose, Blotch, Scorch on American and European lin-
and those with pronounced overgrowth of tissue den. Small, circular to irregular brown spots with
are commonly called scab. Both types are caused dark margins form blotches along main veins
by the genus Elsinoe¨, anamorph state in leaves, leaf stalks, and young twigs, with

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_14, 59

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60 Anthracnose

Fig. 1 Bean Anthracnose.

Pod and seeds with dark,
sunken areas; section
through bean seed showing
spores formed in an
acervulus marked with
prominent black setae

infected later in the season have irregular brown

areas along the veins. Conidia ooze out from
acervuli on underside of veins in flesh-colored
masses, in rainy weather, and are splashed to
other leaves. Twigs and branches have sunken
cankers with more acervuli. Native sycamores
may be nearly defoliated, with smaller twigs
killed. Larger branches die with several succes-
sive wet springs. The trees usually put out
a second crop of leaves after defoliation, but
this is a devitalizing process. Dead twigs and
branches give a witches’ broom effect to the
On white oaks anthracnose appears as brown
areas adjacent to midribs and lateral veins.
Control Although raking up and burning all
fallen leaves has been stressed for years, the
overwintering of the fungus on twigs makes this
measure rather ineffective. The spray schedule
has called for three applications of bordeaux mix-
ture; a dormant spray, one when the buds swell,
and another 7 days later. Trees should be fertil-
ized to stimulate vigorous growth.
Fig. 2 Oak Anthracnose

rose-colored pustules. In wet seasons, defoliation Colletotrichum

in early summer may be followed by wilting and
death of branches. Cut out and burn such Deuteromycetes, Coleomycetes
Apiognomonia veneta (formerly Gnomonia Spores are formed in acervuli, erumpent,
platani) (G. veneta). Sycamore Anthracnose, cushionlike masses of hyphae bearing conidio-
Twig Blight, general on American and Oriental phores and one-celled, hyaline, oblong to fusoid
planes (London plane is rather resistant) and on conidia. Acervuli have stiff marginal bristles
California and Arizona sycamores. The fungus (setae), which are sometimes hard to see. Conidia
winters as mycelium in fallen leaves, producing (slime-spores), held together by a gelantinous
perithecia that discharge ascospores when young coating, appear pinkish in mass. They are not
foliage is breaking out. Mycelium also winters in wind-borne but can be disseminated by wind-
twig cankers. Young sycamore leaves turn brown splashed rain. On landing on a suitable host, the
and die, looking as if hit by late frost. Leaves conidium sends out a short germ tube, which,
Colletotrichum 61

on contact with the epidermis, enlarges at the tip Colletotrichum dematium f. sp. truncata An-
into a brown thick-walled appressorium. From this, thracnose on tomato. Found in Georgia on
a peglike infection hypha penetrates the cuticle. Dolichos.
Colletotrichum acutatum Anthracnose on Colletotrichum erumpens (▶Glomerella
almond, strawberry and black gum. cingulata). Rhubarb Anthracnose, Stalk Rot.
Colletotrichum antirrhini (▶Glomerella Colletotrichum fragariae (▶Glomerella
cingulata) Snapdragon Anthracnose, on snap- cingulata). Strawberry Anthracnose found in
dragon, chiefly in greenhouses, sometimes out- Florida and Louisiana.
doors in late summer. Colletotrichum fuscum Foxglove Anthrac-
Colletotrichum atramentarium (or C. nose small spots to 1/8 inch, circular to angular,
coccodes). Potato Anthracnose, Black Dot Dis- brown to purple brown, on leaves; sunken, fusi-
ease on potato stems and stolons following wilt form lesions on petioles and veins; minute black
and other stem diseases, occasionally on tomato, acervuli, with bristles, in center of spots. Seed-
eggplant, and pepper; general distribution but lings damp-off, older plants are killed or stunted
minor importance. Starting below the soil surface, in warm moist weather. Use clean seed or treat
brown dead areas extend up and down the stem. with hot water (131  F for 15 min).
The partial girdling causes vines to lose their fresh Colletotrichum gloeosporioides (▶Glomerella
color and lower leaves to fall. Infection may cingulata). Lime Anthracnose, Withertip, only
extend to stolons and roots. The black dots embed- on lime in southern Florida.
ded in epidermal cells, inside hollow stems and on Colletotrichum graminicola (formerly
tubers, are sclerotia to carry the fungus over winter Colletotrichum sublineola). Anthracnose on
and to produce conidia the following spring. wild rice (Zizania).
The fungus is a wound parasite ordinarily not Colletotrichum graminicola Cereal Anthrac-
serious enough to call for control measures nose widely distributed on barley, oats, rye,
other than cleaning up old refuse and using wheat, sorghum, wild rice (Zizamia) and also on
healthy seed potatoes. cultivated lawn grasses, causing a root decay and
Colletotrichum bletiae (▶Glomerella stem rot. Leaf spots are small, circular to elliptical,
cingulata) and other species. Orchid Anthrac- reddish purple, enlarging and fading with age; cen-
nose, Leaf Spot on orchids coming in from the ters have black acervuli. The fungus winters on
tropics. seed and plant refuse in or on soil. Improved soil
Colletotrichum capsici Ripe Fruit Rot of fertility reduces damage from this disease. This
pepper. pathogen also causes fruit anthracnose of tomato.
Colletotrichum coccodes Anthracnose on Colletotrichum higginsianum Turnip Anthrac-
soybean. nose, also on rutabaga, mustard greens, radish,
Colletotrichum dematium (formerly and Chinese cabbage in southeastern states. Very
Colletotrichum omnivorum). Anthracnose on small, circular gray spots on leaves, and elongate
aspidistra and hosta. Large, whitish spots with brown or gray spots on midrib, petiole, and stem,
brown margins are formed on leaves and stalks. show pink pustules in centers of dead tissue.
Remove and burn infected plant parts. Heavily infected leaves turn yellow and die;
Colletotrichum dematium Anthracnose on young seeds in diseased pods may be killed.
spinach. Mustard variety Southern Curled Giant is highly
Colletotrichum dematium f. sp. resistant.
spinaciae Spinach Anthracnose. Known on Colletotrichum lagenarium (see
spinach since 1880 but unimportant in most ▶Colletotrichum orbiculare). Melon Anthrac-
years. Leaves have few to many circular spots, nose on muskmelon, watermelon, cucumber,
water-soaked, turning gray or brown, with setae and other cucurbits.
prominent in spore pustules. The fungus is seed- Colletotrichum liliacearum (see
borne. ▶Colletotrichum lilii). Found on dead stems of
62 Anthracnose

daylilies and many other plants and perhaps the centers of such spots are covered with gelat-
weakly parasitic. inous masses of salmon-colored spores. Infected
Colletotrichum lilii (formerly Colletotrichum fruit has a bitter taste or the flesh is tough and
liliacearum). Found on dead stems of daylilies insipid. Soft rots often follow the anthracnose.
and many other plants and perhaps weakly Epiphytotics occur only in periods of high rainfall
parasitic. and temperature, near 75  F.
Colletotrichum lindemuthianum (▶- Control Treating seed before planting is essen-
Glomerella lindemuthianum). Bean Anthrac- tial. Use a three year crop rotation with non-
nose, a major bean disease, sometimes cucurbits; destroy plant refuse. Watermelon
mistakenly called “rust,” generally present in varieties Charleston Gray, Congo, Fairfax, and
eastern and central states, rare from the Rocky Black Kleckly are resistant but not to all races
Mountains to the Pacific Coast. of the fungus.
Colletotrichum malvarum Hollyhock Colletotrichum phomoides (▶Glomerella
Anthracnose, Seedling Blight on hollyhock, cingulata). Tomato Anthracnose, common rot
mallow, and abutilon, particularly destructive to of ripe tomatoes, most frequent in Northeast and
greenhouse seedlings. Black blotches are formed North Central districts. Symptoms appear late in
on veins, leaf blades, petioles, and stems. the season, causing more loss to canning crops.
Remove and burn all old plant parts in autumn. Small, circular sunken spots, increasing to an
Colletotrichum omnivorum (see inch in diameter, penetrate deeply into the flesh.
▶Colletotrichum dematium). Anthracnose on At first water-soaked, the spots turn dark, with
aspidistra and hosta. Large, whitish spots with pinkish, cream, or brown spore masses in the
brown margins are formed on leaves and stalks. depressed centers, often arranged in concentric
Remove and burn infected plant parts. rings. The disease is worse in warm, moist
Colletotrichum orbiculare Anthracnose on weather. The fungus winters in tomato refuse,
watermelon. sometimes in cucumber and melon debris.
Colletotrichum orbiculare (formerly Control Clean up trash and rotting fruit.
Colletotrichum lagenarium). Melon Anthrac- Colletotrichum pisi (Glomerella cingulata).
nose on muskmelon, watermelon, cucumber, Pea Anthracnose, Leaf and Pod Spot commonly
and other cucurbits. This is our most destructive associated with Ascochyta blight and often
disease of watermelons, found everywhere that a secondary parasite.
melons are grown and particularly destructive in Colletotrichum schizanthi Anthracnose on
the South. There are at least three races of the butterfly-flower. Symptoms are small brown
fungus differing in ability to infect different spots on leaves and water-soaked areas on
cucurbits. One race is virulent on cucumber, young stems. Cankers on stems and branches of
slight on watermelon, moderate on Butternut older plants may cause leaves to turn yellow,
squash; another is virulent on both watermelon branches to die back from the tip, and finally
and cucumber; Butternut squash is immune to death of all parts above the canker.
a third. Colletotrichum sublineola (see
Leaf symptoms are small yellow or water-soaked ▶Colletotrichum graminicola). Anthracnose on
areas, which enlarge and turn black on water- wild rice (Zizania).
melon, brown on muskmelon and cucumber. Colletotrichum trichellum Fruit Anthracnose
The dead tissue shatters; leaves shrivel and die. of tomato and Hedera.
Elongated, narrow, sunken lesions appear on Colletotrichum truncatum Stem Anthracnose
stems and petioles; vines may die. Young fruit prevalent in the South on bean, lima bean, and
darkens, shrivels and dies if pedicels are infected; soybean, also on clovers and on lentil in ND.
older fruit shows circular, black, sunken cankers Brick-red spots appear on veins on underside of
or depressions, from 1/4 to 2 inches across and leaves and on pods. Plants are chlorotic, stunted,
1/3 inch deep on watermelon. In moist weather may die prematurely; blossoms or pods may drop.
Gloeosporium 63

Use healthy seed grown in arid states; clean up Gloeosporium melongenae (▶Glomerella
plant refuse; rotate with non-legumes. cingulata). (possibly identical with G.
Colletotrichum violae-tricoloris (▶Glomerella piperatum). Eggplant Anthracnose, Ripe Rot,
cingulata). Anthracnose of violet, pansy. an occasional trouble.
Colletotrichum sp. Azalea Anthracnose. New Gloeosporium piperatum (▶Glomerella
disease serious on Indian and Kurume azaleas cingulata). Pepper Anthracnose, Fruit Spot,
in Louisiana since 1954. Very small rusty sometimes a leaf and stem spot but more often
brown spots appear on both surfaces of a disease of green or ripe fruit.
young leaves, followed by defoliation. Spores Gloeosporium quercinum (Telemorph,
appear on fallen leaves, which serve as ▶Apiognomonia errabunda). Oak Anthracnose.
source of inoculum for the next season. Cop- See ▶Discula umbrinella and Fig. 2.
per and organic fungicides are effective in Gloeosporium thuemenii f. sp. tulipi Tulip
control. Anthracnose found in California in 1939.
Lesions on peduncles and leaf blades of Darwin
tulips are small to large, elliptical, first water-
soaked then dry with black margins and numer-
ous black acervuli in center of spots.
Gloeosporium sp Peony Anthracnose on
▶ Blights.
stems, leaves, flowers, petals of peony. Stem
Discula campestris Anthracnose on maple.
lesions are sunken, with pink spore pustules,
Discula destructive Anthracnose on dogwood.
and may completely girdle the stalks, causing
Discula fraxinea (Teleomorph, Gnomoniella
death of plants. Also a destructive anthracnose
fraxini). Anthracnose on ash.
on strawberry.

Gloeosporium Glomerella

Deuteromycetes, Coleomycetes Ascomycetes, Phyllachorales

Perithecia are dark, hard, carbonaceous, usu-
Genus characters are about the same as for ally beaked, immersed in substratum so only the
Colletotrichum except that there are no setae neck protudes. Ascospores are hyaline, one-
around the acervuli. Conidia are hyaline, one- celled; asci are thickened at tips, inoperculate
celled, appearing in masses or pustules on leaves but spores sometimes discharged with force;
or fruit. Leaf spots are usually light brown, with paraphyses present.
foliage appearing scorched. Glomerella cingulata (formerly
Gloeosporium allantosporum (▶Phlyctema Colletotrichum violae-tricoloris). Anthracnose
vagabunda). Anthracnose, Dieback on raspberry of violet, pansy. Circular dead spots with black
in Oregon, Washington. margins, sometimes zonate, appear on leaves;
Gloeosporium apocryptum (▶Glomerella flowers have petals spotted or not fully developed
cingulata). Maple Anthracnose, Leaf Blight, an and producing no seed; entire plants are some-
important leaf disease of silver maple, common times killed. Remove and burn infected plants or
also on other maples and boxelder, appearing parts; clean up old leaves in fall. Copper sprays
from late May to August. may be injurious.
Gloeosporium aridum (Discula fraxinea) Glomerella cingulata (formerly
Anthracnose on ash. Colletotrichum gloeosporioides). Anthracnose,
Gloeosporium limetticolum (▶Glomerella Canker, Dieback, Withertip, Fruit Rot of a great
cingulata). Lime Anthracnose, Withertip, only many plants, generally distributed except on the
on lime in southern Florida. Pacific Coast, more common in the South.
64 Anthracnose

Infection is often secondary, in tissues weakened spread by tools, insects, and workmen. Rotation
from other causes. See also under ▶ Cankers and of crops and sanitary measures may be sufficient
Diebacks and under ▶ Rots. control.
On citrus, orange, lemon, grapefruit there is Glomerella cingulata (formerly Gloeosporium
a dying back or withertip of twigs. Leaf spots are limetticolum). Lime Anthracnose Withertip, only
light green turning brown, with pinkish spore pus- on lime in southern Florida. Shoots, leaves, and
tules prominent in wet weather. Decayed spots are fruits are infected when young; mature tissues are
produced on ripening fruits in storage. Similar immune. Twigs wither and shrivel from one inch
withertip symptoms may also appear on avocado, to several inches back from the tip; young leaves
aucuba, cherimoya, fig, loquat, roselle, have dead areas or are distorted; buds fail to open
rosemallow, royal palm, dieffenbachia, rubber- and may drop; fruits drop, or are misshapen, or
plant, strawberry and other ornamentals and fruits. have shallow spots or depressed cankers.
The disease has also been reported on European Control Spray with bordeaux-oil emulsion as
white birch in Virginia. Lack of water and nutrient fruit is setting, with two or three applications of
deficiency predispose plants to infection by this 1 to 40 lime sulfur at 7–14 day intervals.
weak parasite. The fungus attacks blue lupine and Glomerella cingulata (formerly Gloeosporium
statice or sea-lavender; peach anthracnose became apocryptum). Maple Anthracnose Leaf Blight, an
important in Georgia when lupine was used as important leaf disease of silver maple, common
a ground cover in orchards. Sweet pea anthracnose also on other maples and boxelder, appearing from
is often more severe near apple orchards where the late May to August. The leaf spots are light brown,
fungus winters on cankered apple limbs and in often merging with the leaves, appearing scorched.
bitter rot mummies. Whitish lesions disfigure The effect may be confused with the physiological
sweet pea leaves, shoots, and flower stalks. Leaves scorch caused by hot weather. On Norway maples
wither and fall; stalks dry up before blossoming; the leaf lesions are confined to purple to brown
seed pods shrivel. There may be general wilting lines along the veins. In rainy seasons there may
and shoot dieback. be severe defoliation.
Anthracnose and twig blight are widespread Control If trees have been affected more than
on privet. Leaves dry and cling to the stem; can- a year or so, feed to stimulate vigorous growth.
kers at the base of stems are dotted with pink Spray with a copper fungicide two or three times at
pustules. Bark turns brown and splits; death fol- 14-day intervals, starting when buds break open.
lows complete girdling of stems. European privet Glomerella cingulata (formerly
is highly susceptible; California, Amur, Ibota, Colletotrichum bletiae) and other species. Orchid
and Regal privets are fairly resistant. Also found Anthracnose Leaf Spot on orchids coming in from
on black locust in GA and SC and pecan in GA. the tropics. Lemon-colored acervuli are formed in
Control Remove infected twigs and branches soft, blackish spots in ragged leaves. Burn diseased
from trees and shrubs, taking care to make plants or parts. Spray with a copper fungicide.
smooth cuts at base of limbs and painting surfaces Glomerella cingulata (formerly
with a wound dressing. Plant sweet peas, from Colletotrichum pisi). Pea Anthracnose Leaf and
healthy pods, at a distance from apple and privet, Pod Spot commonly associated with Ascochyta
in clean soil; rake up and burn plant refuse at the blight and often a secondary parasite. Spots on
end of the season. pods, stems, and leaves are sunken, gray, circular,
Glomerella cingulata (formerly Gloeosporium with dark borders. Crop rotation is the best control.
melongenae). (possibly identical with G. Glomerella cingulata (formerly Gloeosporium
piperatum). Eggplant Anthracnose Ripe Rot, an piperatum). Pepper Anthracnose, Fruit Spot,
occasional trouble. Yellow to brown spots on sometimes a leaf and stem spot but more often
leaves and small to medium depressed spots on a disease of green or ripe fruit. Spots are dark,
fruit show pink spore masses following rain or sunken, with concentric rings of acervuli and
heavy dew. Spores are splashed by rain and pink masses of spores, which are washed to
Gnomonia 65

other fruit. Seed is infected internally and con- Pacific Coast. It may also affect lima bean, Scarlet
taminated externally. Harvest seed only from runner, tepary, mung, kudzu, and broad beans, and
healthy fruit. cowpea. It is worldwide in distribution, known in
Glomerella cingulata (formerly the United States since 1880. There are at least 34
Colletotrichum erumpens). Rhubarb Anthrac- strains of the fungus, in three different groups, but
nose, Stalk Rot. Oval, soft watery spots on petioles the disease has decreased in importance with the
increase until whole stalks are included; leaves wilt use of western-grown, anthracnose-free seed.
and die. Small dark fruiting bodies with setae sur- The most conspicuous symptoms are on the pods,
vive winter in stems, produce conidia in spring. small, brown specks enlarging to black, circular,
Clean up all rhubarb remains in fall. sunken spots, in moist weather showing the typical
Glomerella cingulata (formerly pinkish ooze of the slime-spores. Older spots often
Colletotrichum antirrhini). Snapdragon have narrow reddish borders. After the spores are
Anthracnose on snapdragon, chiefly in green- washed away, the acervuli look like dark pimples. If
houses, sometimes outdoors in late summer. pods are infected when young, the disease extends
Stems have oval, sunken spots, grayish white through to the seed, which turns yellow, then rusty
with narrow brown or reddish borders, fruiting brown or black under the pod lesion. The infection
bodies showing as minute black dots in center. may extend deep enough to reach the cotyledons.
Spots on leaves are circular, yellow green turning Leaf lesions are dark areas along veins on underside
dirty white, with narrow brown borders. Stem of the blade and on petioles. Seedlings may show
cankers may coalesce to girdle plant at base, stem spotting below diseased cotyledons. The fun-
causing collapse of upper portions, with leaves gus is spread by splashing rain, tools, and gardeners
hanging limp along the stem. working with beans when they are wet. Optimum
Control Take cuttings from healthy plants; provide temperature is between 63 and 75  F, with maxi-
air circulation; keep foliage dry; destroy infected mum around 85  F.
outdoor plants in autumn. Spray, every 7 to 10 days. Control Use western-grown seed. Saving home-
Glomerella cingulata (formerly grown seed is dangerous unless you can be sure of
Colletotrichum phomoides). Tomato Anthrac- selecting from healthy plants and pods. Clean up,
nose, common rot of ripe tomatoes, most frequent or spade under, old bean tops; rotate crops. Never
in Northeast and North Central districts. Symp- pick or cultivate beans when vines are wet. There
toms appear late in the season, causing more loss are some resistant varieties, but more reliance
to canning crops. Small, circular sunken spots, should be placed on obtaining seed grown
increasing to an inch in diameter, penetrate deeply where the disease is not present.
into the flesh. At first water-soaked, the spots turn Glomerella nephrolepidis Fern Anthracnose,
dark, with pinkish, cream, or brown spore masses Tip Blight of Boston and sword ferns. The soft
in the depressed centers, often arranged in concen- growing tips of fronds turn brown and dry. Keep
tric rings. The disease is worse in warm, moist foliage dry; remove and burn diseased leaves.
weather. The fungus winters in tomato refuse,
sometimes in cucumber and melon debris.
Control Clean up trash and rotting fruit.
Glomerella glycines Fruit Anthracnose of Gnomonia
tomato. Also, anthracnose on soybean.
Glomerella gossypii Fruit Anthracnose of Ascomycetes, Diaporthales
tomato. Also, anthracnose on cotton.
Glomerella lindemuthianum (formerly Perithecia innate, beaked, separate; paraphyses
Colletotrichum lindemuthianum). Bean Anthrac- absent; ascospores two-celled, hyaline;
nose, a major bean disease, sometimes mistakenly anamorph state Gloeosporium or Marssonina.
called “rust,” generally present in eastern and cen- Diseases caused by Gnomonia are classified as
tral states, rare from the Rocky Mountains to the anthracnose, scorch, or leaf spot.
66 Anthracnose

Gnomonia caryae Hickory Anthracnose, Leaf Monographella opuntiae (formerly

Spot, widespread. The disease is common in Mycosphaerella opuntiae). Cactus Anthracnose
eastern states, causing defoliation in wet seasons. on Cereus, Echinocactus, Mammillaria, and
Large, roundish spots are reddish brown on upper Opuntia. The curved spores of the anamorph
leaf surface, dull brown underneath. The fruiting state (Microdochium lunatum) form light pink
bodies are minute brown specks, and the fungus pustules on the surface of moist, light brown rotten
winters in dead leaves on the ground. areas. Cut out and destroy diseased segments.
Gnomonia leptostyla (Marssonina juglandis).
Walnut Anthracnose, Leaf Spot, general on but-
ternut, hickory, and walnut. Spring infection Pezicula
comes from ascospores shot from dead leaves
on the ground, secondary infection from conidia. Ascomycetes, Helotiales
Irregular dark brown spots appear on leaflets in
early summer; if these are numerous, there is This is one of the discomycetes, cup fungi. The
defoliation. An unthrifty condition of black wal- apothecia, formed on living plants, are fleshy,
nuts and butternuts is often due to anthracnose. bright-colored with a peridium of dark cells
forming a pseudoparenchyma. Spores are hya-
line, fusoid.
Pezicula malicorticis (formerly Neofabraea
malicorticis). Northwestern Apple Anthracnose
on apple, crabapple, pear, quince, chiefly in the
Deuteromycetes, Coelomycetes
Pacific Northwest, where it is a native disease,
serious in regions with heavy rainfall. Cankers are
Hyaline, two-celled spores are formed in acervuli
formed on younger branches – elliptical, dark,
without setae. Spores are rounded at ends and are
sunken, up to 3 or 4 inches wide and 10 to 12 inches
formed in pale to black masses on leaves.
long, delimited when mature by a crack in the bark.
Microdochium panattonianum (formerly
Conidia of the anamorph state (Gloeosporium
Marssonina panattoniana). Lettuce Anthrac-
malicorticis) are formed in cream-colored cushions,
nose. Small, dead, brown spots appear on blades
which turn black with age, in slits in the bark.
and petioles, centers often falling out leaving
Young cankers, reddish brown, circular spots
black margined shot holes. Spots progress from
appear on the bark in late fall. Fruit is infected,
older to young inner leaves; outer leaves are
usually through lenticels from either ascospores or
broken off and blown around by wind. The dis-
conidia in pustules on bark, but the disease may not
ease is important only during prolonged periods
show up until the apples are in storage.
of wet weather, when it may cause heavy losses.
Control Cut out diseased limbs or excise can-
Sanitary measures and treating seed before plant-
kers, burning all prunings and dead bark. Spray
ing suffice for control.
with bordeaux mixture before fruit is picked and
fall rains start; repeat after harvest, and again
about 2 weeks later.

Ascomycetes, Dothideales Phlyctema

Perithecia immersed in substratum, not beaked, Deuteromycetes, Coleomycetes

not setose, paraphyses lacking; spores hyaline,
two-celled. The genus contains more than 1,000 Pycnidia dark, separate or sometimes cofluent,
species, many destructive to plants, with conidial developing in or under the epidermis or bark.
stages in many genera. Conidiophores simle or forked; conidia hyaline,
Pseudopeziza 67

one-celled, cylindrical or log, spindle-shapped, crescent-shaped conidia are formed in moist,

mostly bent. flesh-colored masses in center of spots. In severe
Phlyctema vagabunda (formerly infections there is progressive defoliation from
Gloeosporium allantosporum). Anthracnose, below upward.
Dieback on raspberry in Oregon, Washington. Other occasional symptoms are black,
See ▶Elsinoe¨ veneta under Spot Anthracnose for sunken spots on leaf stalks, light brown to
the common raspberry disease called anthracnose. pale yellow lesions on canes, and black
flyspeck spots on green berries, with considerable
reduction in yield. Apothecia are formed
Pseudopeziza on fallen leaves; ascospores are forcibly
discharged in spring and carried by wind to
Ascomycetes, Helotiales young leaves.
Control Clean up and burn old leaves under the
Apothecia brown, cup-shaped, arising from bushes. Spray with bordeaux mixture (preferred
leaves on short stalks, not setose, paraphyses to the newer organics) shortly after leaves
present; spores one-celled, hyaline, ovoid. appear (about 3 weeks after blossoming) and
Drepanopeziza ribis (formerly Pseudopeziza immediately after picking. Include a good
ribis). Current Anthracnose, Leaf, Stem and spreader and cover both leaf surfaces
Fruit Spot generally distributed on currant, thoroughly.
flowering currant, and gooseberry, first Pseudopeziza ribis (see ▶Drepanopeziza
reported on black currants in Connecticut in ribis). Current Anthracnose, Leaf, Stem and
1873. Very small, brown, circular spots Fruit Spot generally distributed on currant,
appear first on lower, older leaves, which turn flowering currant, and gooseberry, first reported
yellow if spots are numerous. Hyaline, on black currants in Connecticut in 1873.
Bacterial Diseases

Rhizobiaceae spherical protuberances or elongated ridges of

white gall tissue, turning brown after several
Agrobacterium weeks. Canes often split open and dry out; pro-
duce small seedy berries. Cane gall is not as
Small, motile, short rods, with two to six peritri- important as crown gall, but one should use the
chous flagella or a polar or subpolar flagellum, same preventive measures. Avoid runner plants
ordinarily Gram-negative, not producing visible from infected mother plants.
gas or detectable acid in ordinary culture media; Agrobacterium tumefaciens Crown Gall on
growth on carbohydrate media usually accompa- a great variety of plants in more than 40 families,
nied by copious entracellular, polysaccharide general on blackberry, raspberry, and other bram-
slime; gelatin liquefied slowly or not at all; opti- bles, on grapes and on rose (see Fig. 1); on fruit
mum temperatures 25 to 30  C. Found in soil, or trees – apple, apricot, cherry, fig, peach and nec-
plant roots in soil, or in hypertrophies or galls on tarine, pear (rarely), plum; on nuts – almond very
roots or stems of plants. susceptible, walnut fairly susceptible, pecan
Agrobacterium rhizogenes Hairy Root of occasionally; on shade trees, willow and other
apple, also recorded on cotoneaster, hollyhock, hard woods; rare on conifers but reported on
honey locust, honeysuckle, mulberry, peavine, incense cedar and juniper; on many shrubs and
peach, quince, Russian olive, rose, and spirea. vines, particularly honeysuckle and euonymus;
“Woolly root” and “woolly knot” are other on perennials such as asters, daisies, and chrysan-
names given to this disease, which was long themums; and on beets, turnips, and a few other
considered merely a form of crown gall. Both vegetables, with tomato widely used in experi-
diseases may appear on the same plant and in ments. Crown gall was first noticed on grape in
early stages be confused. In hairy root a great Europe in 1853, and the organism was first iso-
number of small roots protrude either directly lated in 1904 in the United States from galls on
from stems or roots or from localized hard swell- Paris daisy. It is of first importance as a disease of
ings that frequently occur at the graft union. The nursery stock, but may cause losses of large pro-
disease is common on grafted nursery apple trees ductive trees in neglected orchards, especially
1, 2, or 3 years old, and the root development may almonds and peaches in California and other
be as profuse as witches’ broom. Control mea- warm climates. It is very important to rose
sures are the same as for crown gall. growers and to the amateur gardeners who some-
Agrobacterium rubi Cane Gall of brambles, on times receive infected bushes.
blackberry, black and purple raspberries, and, Symptoms The galls are usually rounded, with an
very rarely, red raspberry. Symptoms appear on irregular rough surface, ranging up to several
fruiting canes in late May or June as small, inches, usually occurring near the soil line,

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_15, 69

# Springer Science+Business Media Dordrecht 2013
70 Bacterial Diseases

more than a couple of years, and that sudden

outbreaks of crown gall on land not previously
growing susceptible crops are due to irrigation
water bringing in viable bacteria from other
infected orchards. The addition of lime to the
soil may encourage crown gall, for bacteria do
not live in an acid medium. The period of greatest
activity is during the warm months.
Control For home gardens rigid exclusion of all
suspected planting stock is the very best control.
Do not accept from your nurseryman blackberries,
raspberries, roses, or fruit trees showing suspicious
bumps. If you have had previous trouble, choose
a different location for new, healthy plants. Be
careful not to wound stems in cultivating.
For nurserymen, sanitary propagating prac-
tices are a must. Stock should be healthy.
Grafting knives should be sterilized by frequent
dipping in 10 % Chlorox solution, 1 ounce in 2
gallons of water, or in denatured alcohol. If nurs-
ery soil is infested, 2 years’ growth of cowpeas,
oats, or crotalaria between crops will minimize
crown gall.
Fruit and nut growers can perhaps plant less
Fig. 1 Crown Gall on Rose susceptible varieties, although fruit that is resis-
tant in one locality may be diseased in another.
commonly at the graft union, but sometimes on American grape varieties are considered more
roots or aerial parts. On euonymus, galls are resistant than European. Apples may be better
formed anywhere along the vine. This is primar- on mahaleb root-stock, nut trees on black walnut
ily a disease of the parenchyma, starting with understock. Budding rather than grafting reduces
a rapid proliferation of cells in the meristematic the chance of infection.
tissue and the formation of more or less convo- Painting galls with a solution of Elgetol-
luted soft or hard overgrowths or tumors. The methanol has given control of crown gall on
close analogy of the unorganized cell growth of peaches and almonds in California. One part
plant galls to wild cell proliferation in human Elgetol (sodium dinitrocresol) is shaken with 4
cancer has intrigued scientists for many years. parts synthetic wood alcohol and applied with
In some fashion bacteria provide stimulus for a brush, covering the surface of the gall and
this overdevelopment, but similar galls have extending 1/2 inch to 1 inch beyond the margin
been produced on plants experimentally by into healthy bark.
injecting a virus or growth-promoting substances.
Entrance of bacteria into plants for natural
infection is through wounds. In nurseries and Coryneform Group Mycobacteriaceae
orchards nematodes, the plow, the disc, or the
hoe may be responsible; on the propagating Clavibacter
bench grafting tools are indicted. Many claims
have been made for the longevity of crown gall Slender, straight to slightly curved rods, with
bacteria in soil, but it now seems to be established irregularly stained segments or granules, often
that they do not live in the absence of host plants with pointed or club-shaped swellings at ends;
Coryneform Group Mycobacteriaceae 71

nonmotile with a few exceptions (C. Control Use certified seed, a 2-or 3-year rota-
flaccumfaciens and C. poinsettiae). Gram- tion; clean up tomato refuse at end of season and
positive. diseased plants throughout season. Fermenting
Clavibacter agcopyri (see Corynebacterium tomato pulp for 4 days at a temperature near
agcopyei). Yellow Gum Disease on western 70  F will destroy bacteria on surface of seed;
wheat grass. hot-water treatment, 25 min at 122  F will kill
Clavibacter fascians (see Rhodococcus some, perhaps not all, of internal bacteria. Start
fascians). Fasciation, widespread on sweet pea, seedlings in soil that has not previously grown
also on carnation, chrysanthemum, gypsophila, tomato.
geranium, petunia, impatiens, Hebe sp. and Clavibacter poinsettiae (see Curtobacterium
pyrethrum. flaccumfaciens pv. poinsettia). Stem Canker and
Clavibacter flaccumfaciens (see Leaf Spot of Poinsettia, a relatively new disease,
Curtobacterium flaccumfaciens pv. first noted in greenhouses in 1941.
flaccumfaciens). Bacterial Wilt of bean, wide- Clavibacter sepedonicum (see Clavibacter
spread on kidney and lima beans and soybean, michiganense subsp. sepedonicum). Bacterial
causing considerable loss. Ring Rot of potato, widespread since 1931,
Clavibacter humiferum (see Corynebacterium when it probably was introduced from Europe.
humiferum). Reported from wetwood of poplar, Clavibacter michiganense subsp.
in Colorado. sepedonicum (formerly Clavibacter
Clavibacter michiganense (see Clavibacter sepedonicum). Bacterial Ring Rot of potato,
michiganense subsp. michiganense). Bacterial widespread since 1931, when it probably was
Canker of Tomato, widespread, formerly causing introduced from Europe. All commercial varie-
serious losses of tomato canning crops. ties are susceptible, with losses formerly in mil-
Clavibacter michiganense subsp. lions of dollars in decay of tubers in field and
michiganense (formerly Clavibacter storage. Now a single infected plant in a potato
michiganens). Bacterial Canker of Tomato, wide- field disqualifies the whole field for certification.
spread, formerly causing serious losses of tomato Symptoms appear when plants are nearly full
canning crops. The disease has now been grown, with one or more stems in a hill wilted
reported on browallia, brunfelsia, cestrum, and stunted while the rest seem healthy. Lower
Datura sp., eggplant, Jerusalem-cherry, leaves have pale yellow areas between veins;
bittersweet, pepper, painted-tongue, potato, these turn deeper yellow, and margins roll
ground-cherry, and butterfly-flower in Wyoming. upward. A creamy exudate is expelled when the
This is a vascular wilt disease, seedlings stem is cut across. This bacterium may also occur
remaining stunted. Symptoms on older in sugar beet which are symptomless.
plants start with wilting of margins of lower leaf- Tuber infection takes place at the stem end,
lets, often only on one side of a leaf. Leaflets curl and the most prominent symptoms appear some
upward, brown, and wither, but remain attached time after storage. The vascular ring turns creamy
to stem. One-sided infection may extend up yellow to light brown, with a crumbly or cheesy
through the plant and open cankers from pith to odorless decay followed by decay from second-
outer surface of stem. Fruit infection starts ary organisms. Bacteria are not spread from plant
with small, raised, snow-white spots, centers to plant in the field, but by cutting knife and
later browned and roughened but the white color fingers at planting. A knife used to cut one
persisting as a halo to give a bird’s-eye infected tuber may contaminate the next 20 seed
spot. Fruits can be distorted, stunted, yellow pieces.
inside. In the field, bacteria are spread by Control Use certified seed potatoes. Use several
splashed rain and can persist in soil 2 or more knives and rotate them in disinfestant. Commer-
years. Seeds carry the bacteria internally as well cial growers use a rotating knife passed through
as externally. a chemical or hot-water bath between cuts.
72 Bacterial Diseases

Disinfest tools, grader, digger, and bags; sweep Rhodococcus fascians (formerly Clavibacter
storage house clean and spray with copper sul- fascians). Fasciation, widespread on sweet pea,
fate, 1 pound to 5 gallons of water. also on carnation, chrysanthemum, gypsophila,
Clavibacter xyli subsp. cynodontis Stunting geranium, petunia, impatiens, Hebe sp. and pyre-
Disease of bermudagrass. thrum. Sweet pea symptoms are masses of short,
Clavibacter xyli subsp. xyli Ratoon Stunt of thick, and aborted stems with misshapen leaves
sugarcane. developing near the soil line at first or second
Corynebacterium agropyri (formerly stem nodes. The fasciated growth on old plants
Clavibacter agropyri). Yellow Gum Disease on may have a diameter of 3 inches but does not
western wheat grass. Enormous masses of surface extend more than an inch or two above ground.
bacteria form yellow slime between stem and The portion exposed to light develops normal
upper sheath and glumes of flower head; plants green color. Plants are not killed, but stems are
dwarfed or bent; normal seeds rare. dwarfed and blossom production is curtailed.
Corynebacterium humiferum formerly Control Sterilize soil or use fresh.
Clavibacter humiferum). Reported from Rickettsialike bacteria Bacterial Wilt on
wetwood of poplar, in Colorado. Toronto creeping bentgrass; bacteria found in
Curtobacterium flaccumfaciens pv. xylem of roots, crown, and leaves. Initially, leaf
flaccumfaciens (formerly Clavibacter blades wilt from tip down and within several days
flaccumfaciens). Bacterial Wilt of Bean, wide- entire leaf wilts, becomes dark green, shriveled,
spread on kidney and lima beans and soybean, and twisted; also leaf scorch of mulberry.
causing considerable loss. Plants wilt at any stage
from seedling to pod-production, with leaves
turning dry, brown, and ragged after rains. Plants Enterobacteriaceae
are often stunted. Bacteria winter on or in seed,
which appear yellow or wrinkled and varnished. Erwinia
When infected seed is planted, bacteria pass from
cotyledons into stems and xylem vessels. Other Motile rods (usually) with peritrichous flagella;
plants are infected by mechanical injury and per- Gram-negative; producing acid with or without
haps by insects, but there is not much danger from visible gas from a variety of sugars; invading
splashed rain. Plants girdled at nodes may break tissues of living plants producing dry necroses,
over. galls, wilts, and soft rots. The genus is named for
Control Use seed grown in Idaho or California. Erwin F. Smith, pioneer in plant diseases caused
Curtobacterium flaccumfaciens pv. poinset- by bacteria.
tia (formerly Clavibacter poinsettiae). Stem Enterobacter cloacae Bulb Decay on onion.
Canker and Leaf Spot of Poinsettia, a relatively Erwinia amylovora Fire Blight, general on
new disease, first noted in greenhouses in 1941. many species in several tribes of the Rosaceae,
Longitudinal water-soaked streaks appear on one particularly serious on apple, pear, and quince.
side of green stems, sometimes continuing through Other hosts include almond, amelanchier, apri-
leaf petioles to cause spotting or blotching of cot, aronia, blackberry, cherry, chokecherry,
leaves and complete defoliation. The cortex of cotoneaster, crabapple, exochorda, geum, haw-
stems turns yellow, the vascular system brown. thorn, holodiscus, India hawthorn, kerria, Japa-
Stems may crack open and bend down, with glis- nese quince, loquat, medlar, mountain-ash, plum,
tening, golden brown masses of bacteria oozing photinia, pyracantha, raspberry, rose, spirea, and
from stem ruptures and leaf lesions. strawberry.
Control Discard diseased stock plants; place Apparently a native disease, first noticed near
cuttings from healthy mother plants in sterilized the Hudson River in 1780, fire blight spread south
media; avoid overhead watering and syringing; and west with increased cultivation of pears and
rogue suspicious plants promptly. apples. By 1880 it had practically wrecked pear
Enterobacteriaceae 73

Fig. 2 Fire Blight. A hold

over canker developed on
apple limb at base of
blighted twig; B bacteria
swarming through tissue

orchards in Illinois, Iowa, and other states in the The tissue first appears water-soaked, then
Northern Mississippi Valley. Then it devastated reddish, then brown to black as the bacteria
pears on the Texas Gulf. Reaching California by swarm between the dying parenchyma cells.
1910 it played havoc up the coast to Washington. Division may take place every half hour; so they
Symptoms Blossoms and leaves of infected multiply rapidly and are usually well in advance
twigs suddenly wilt, turn dark brown to black, of discolored external tissue. A collar rot may
shrivel and die, but remain attached to twigs (see develop when cankers are formed near the base
Fig. 2). The bark is shrunken, dark brown to of a tree. Water sprouts are common sources of
purplish, sometimes blistered with gum oozing infection.
out. Brown or black blighted branches with dead As spring changes to summer, the bacteria
persistent leaves look as if scorched by fire. The gradually become less active and remain dormant
bacteria survive the winter in living tissue at the at the edge of a woody canker until the next
edge of “holdover cankers” on limbs. These are spring at sap flow. Ordinarily they do not winter
dead, slightly sunken areas with a definite margin on branches smaller than 1/2 inch in diameter.
or slight crack where dead tissue has shrunk away Control Spraying during bloom is now
from living. In moist weather bacteria appear on a standard means of preventing blossom blight.
the surface of cankers in pearly viscid drops of Use bordeaux mixture or a fixed copper or strep-
ooze, which is carried by wind-blown rain or tomycin at 60 to 100 ppm. The latter is very
insects to blossoms. Infection spreads from the effective at relatively high temperatures; at
blighted bloom to the young fruit, then down the 65  F and below, copper is more satisfactory.
pedicel to adjacent leaves, which turn brown, Start spraying when about 10 % of the blossoms
remaining hanging around the blighted blossom are open and repeat at 5-to 7-day intervals until
cluster. Leaf and fruit blight is also possible by late bloom is over. A dormant spray for aphid
direct invasion, a secondary infection via bacteria control helps in preventing fire blight. One or
carried from primary blossom blight by ants, more sprays may be needed for leafhoppers,
aphids, flies, wasps, fruit-tree bark beetles, and starting at petal fall.
honeybees, sometimes tarnished plant bugs, and Inspect trees through the season and cut or
pear psyllids. break out infected twigs 12 inches below the
74 Bacterial Diseases

portion visibly blighted. If lesions appear on large Erwinia carnegieana Bacterial Necrosis of
limbs they may be painted with one of the fol- giant cactus in the entire habitat of Carnegia
lowing mixtures: gigantea. Long present in southern Arizona, this
I. 1 quart denatured alcohol, 1/4 pint distilled disease was not described until 1942, after it had
water, 3/4 ounce muriatic acid, 1 1/2 pounds encroached on cactus parks and private estates.
zinc chloride. Many giant cacti in the Saguaro National Monu-
II. 100 grams cobalt nitrate, 50 cc glycerine, ment have been killed, with heaviest mortality in
100 cc oil of wintergreen, 50 cc acetic acid, trees 150 to 200 years old.
80 cc denatured alcohol. Symptoms start with a small, circular, light
III. 5 parts cadmium sulfate stock solution spot, usually with a water-soaked margin. The
(1 pound stirred into 2 pints warm water), tissues underneath turn nearly black; the spot
2 parts glycerine, 2 parts muriatic acid, enlarges and has a purplish hue with the center
5 parts denatured alcohol. cracking and bleeding a brown liquid. The rotten
Formulas I and II were developed for use on tissues dry, break up into granular or lumpy
the West Coast, III for New York. The paint is pieces, and fall to the ground. Rotting on one
brushed on the unbroken bark over the lesions side means leaning to that side; when the trunk
and for several inches above and below the can- is girdled near the base, the giant is likely to fall in
ker; it may injure if there are wounds or cuts. a wind-storm. If it does not break, it stands as
In cutting out cankered limbs during the dor- a bare, woody skeleton, with all parenchyma
mant season, take the branch off at least 4 tissue disintegrated. An insect, Cactobrosis
inches back from edge of the canker, and fernaldialis, is largely responsible for the rapid
disinfect the cut. The home gardener may want spread of the disease. The larvae tunnel inside the
to use 10 % Chlorox for tools and bordeaux paint stems most of the year, emerging from May to
for cut surfaces. Dry bordeaux powder is stirred August to pupate for a month or so before the
into raw linseed oil until a workable paste is adult, a tan and brown nocturnal moth, lays eggs.
formed. Control A phosphate dust, applied monthly from
Almost all desirable pear varieties are suscep- April to September, has effectively controlled the
tible to fire blight, particularly Bartlett, Flemish insect vector. Incipient infections can be cut out
Beauty, Howell, Clapps Favorite. Varieties Old and the cavity allowed to dry out and cork over.
Home, Orient, and the common Kieffer are more Before the insect vector was known, fallen trees
or less resistant. Jonathon apples are very suscep- were cut into short lengths, dragged to a burial
tible. Less apt to be severely blighted are pit, covered with a disinfectant, and then with
Baldwin, Ben Davis, Delicious, Duchess, McIn- soil.
tosh, Northern Spy, Stayman, and Winter Erwinia carotovora subsp. carotovora (for-
Banana. At the University of California some merly Erwinia carotovora). Soft Rot of calla,
work has been done on susceptibility of ornamen- originally described from common calla, found
tals to fire blight. Pyracantha angustifolia is quite on golden calla, and also on beet, cactus, cab-
susceptible, but P. coccinea and P. crenulata are bage, cauliflower, celery, cucumber, carrot, egg-
rather resistant. Cotoneaster salicifolia is suscep- plant, geranium (Pelargonium), hyacinth, iris,
tible; C. dammeri, C. pannosa, and C. onion, parsnip, pepper, potato, salsify, sansevie-
horizontalis are more resistant; and C. adpressa ria, tobacco, tomato, and turnip.
and C. microphylla show marked resistance. On calla lily the soft rot starts in upper portion
Cultural methods influence the degree of fire of the corm and progresses upward into leaf and
blight, which is worse on fast-growing succulent flower stalks or down into roots, with the corm
tissue. Avoid heavy applications of nitrogen fer- becoming soft, brown, and watery. Sometimes
tilizers; apply such nitrogen as is required in infection starts at edge of petiole, which turns
autumn or in spring in foliar sprays after danger slimy. Leaves with brown spots and margins die
of blossom blight is over. or rot off at the base before losing color. Flowers
Enterobacteriaceae 75

turn brown; stalks fall over; roots are soft and apparently carried in seed; hot-water treatment
slimy inside the epidermis. Corms may rot so is helpful. Drenching delphinium crowns with
fast the plant falls over without other symptoms, bordeaux mixture has been recommended in the
or the diseased portion may dry down to sunken past. Insect larval control is helpful with potato.
dark spots, in which the bacteria stay dormant to Avoid excessive watering or irrigation.
the next season. Erwinia carotovora subsp. carotovora (for-
On tomatoes, infection takes place through merly Erwinia carotovora var. carotovora). Wilt
growth cracks, insect wounds, or sunscald areas. of sunflower, Kalanchoë; zucchini squash, and
The tissue is at first water-soaked, then opaque, draceana. Soft Rot, general on many vegetables,
and in 3 to 10 days the whole fruit is soft, watery, in field, storage, and transit, and many ornamen-
colorless, with an offensive odor. tals, especially iris. The bacteria were first iso-
Control Scrub calla corms, cut out rotted spots, lated from rotten carrots, whence the name, but
and let cork over for a day or two. Plant in fresh or they are equally at home in asparagus, cabbage,
sterilized soil in sterilized containers and keep turnips and other crucifers, celery, cucumber,
pots on clean gravel or wood racks, never on eggplant, endive, garlic, horseradish, melon,
beds where diseased callas have grown previ- parsnip, pepper, spinach, sunflower (stalk rot),
ously. Grow at cool temperatures and avoid sweet potato, and tomato. Besides wide distribu-
overwatering. tion on iris, soft rot has been reported, among
Erwinia carotovora subsp. atroseptica (for- ornamentals, on chrysanthemum, dahlia, Easter
merly Erwinia carotovora). Potato Blackleg, lily, geranium, orchid, sansevieria, poinsettia,
Basal Stem Rot, Tuber Rot, general on potato. and yellow calla.
This is a systemic disease perpetuated by natu- The bacteria enter through wounds, causing
rally infected tubers. Lower leaves turn yellow; a rapid, wet rot with a most offensive odor. The
upper leaves curl upward; stems and leaves tend middle lamella is dissolved, and roots become
to grow up rather than spread out; stem is black- soft and pulpy. Soft rot in iris often follows
spotted, more or less softened at base and up to borer infestation. Tips of leaves are withered,
3 or 4 inches from ground, and may be covered the basal portions wet and practically shredded.
with bacterial slime; shoots may wilt and fall The entire interior of a rhizome may disintegrate
over. Tubers are infected through the stem end. into a vile yellow mess while the epidermis
The disease is most rapid in warm, moist weather, remains firm. The rot is more serious in shaded
and may continue in storage. The bacteria are locations, when iris is too crowded or planted too
spread on the cutting knife, as with ring rot, and deeply.
by seed-corn maggots, and may persist for a time Control Borer control, starting when fans are
in soil. 6 inches high, has greatly reduced the incidence
Control Use certified seed potatoes and plant of rot. If it appears, dig up the clumps, cut away
whole tubers; if cut seed must be used, allow to all rotted portions, cut leaves back to short fans.
cork over to prevent infection from soil. Practice Allow to dry in the sun for a day or two, then
long rotation; disinfest cutting knife. Late varie- replant in well-drained soil, in full sun with upper
ties seem to be more resistant. portion of the rhizome slightly exposed. Many
Erwinia carotovora subsp. atroseptica (for- good iris growers do not agree with this “sitting
merly Erwinia carotovora). Delphinium Black- duck” method, preferring to cover with an inch of
leg, Foot Rot, Bacterial Crown Rot of perennial soil; but the sun is an excellent bactericide, and
Delphinium; Stem and Bud Rot of Rocket Lark- shallow planting is one method of disease con-
spur. In delphinium there is a soft black discolor- trol. Clean off all old leaves in late fall after frost.
ation at the base of the stem, with bacteria oozing Prevent rot on stored vegetables by saving
out from cracks. In larkspur there is a black rot of only sound, dry tubers, in straw or sand, in
buds as well as yellowing of leaves, blackening of a well-ventilated room with temperature not too
stem, stunting of plants. The bacteria are much above freezing. In the garden, rotate
76 Bacterial Diseases

vegetables with fleshy roots with leafy varieties. to August or September, reaching 5 to 30 pounds
Avoid bruising at harvest time. per square inch (as much as 60 pounds in one
Erwinia chrysanthemi Bacterial Blight of record). The bacteria inhabit ray cells mostly and
Chrysanthemum, a florists’ disease, first noted do not cause a general clogging of water-
in 1950. First evidence of blight is a gray water- conducting tissues. This pressure, caused by fer-
soaked area mid-point on the stem, followed by mentation of tissues by bacteria, causes fluxing,
rot and falling over. The diseased tissue is brown a forcing of sap out of trunks through cracks,
or reddish brown; the rot progresses downward to branch crotches, and wounds. The flux flows
the base of the stem or, under unfavorable condi- down the trunk, wetting large areas of bark and
tions, may be checked with axillary buds below drying to a grayish white incrustation. Bacteria
the diseased area producing normal shoots. Cut- and yeasts working in the flux cause an offensive
tings rot at the base. Sometimes affected plants do odor that attracts insects.
not show external symptoms, and cuttings taken Control Bore drain holes through the wood
from them spread the disease. Bacteria can be below the fluxing wound, slightly slanted to facil-
spread via cutting knife, or fingernails in itate drainage. Install 1/2-inch copper pipe to
pinching, and can live several months in soil. carry the dripping sap away from the trunk and
A form of this species causes a leaf blight of buttress roots. Screw the pipe in only far enough
philodendron and may also infect banana, carna- to be firm; if it penetrates the water-soaked wood,
tion, corn, and sorghum and pith/stem rot of it interferes with drainage.
tomato. Erwinia rhapontica Rhubarb Crown Rot, sim-
Control Snap off cuttings; sterilize soil and ilar to soft rot.
tools. Erwinia stewartii (see Pantoea stewartii). Bac-
Corn rot. Corn leaves show light or dark terial Wilt of corn, Stewart’s Disease on sweet
brown rotting at base; husks and leaf blades corn, sometimes field corn, in the middle regions
have dark brown spots; lower portion of stalk is of the United States, from New York to
rotten, soft, brown, with strong odor of decay; California.
plants may break over and die, with little left Erwinia tracheiphila Bacterial Wilt of cucur-
but a mass of shredded remnants of fibrovascular bits, Cucumber Wilt on cucumber, pumpkin,
bundles. Bacteria enter through hydathodes squash, and muskmelon but not watermelon.
(water pores), stomata, and wounds. The disease is generally east of the Rocky Moun-
Erwinia cypripedii Reported from California, tains and is also present in parts of the West; is
causing brown rot of Cypripedium orchids. most serious north of Tennessee. Total loss of
Small, circular to oval, water-soaked, greasy vines is rare, but a 10 to 20 % loss is common.
light brown spots become sunken, dark brown to This is a vascular wound disease transmitted
chestnut. Affected crowns shrivel; leaves drop. by striped and 12-spotted cucumber beetles. Dull
Erwinia herbicola (see Pantoea herbicola). green flabby patches on leaves are followed by
Leaf Spot of dracaena. On Dracaena sanderana, sudden wilting and shriveling of foliage, and
gypsophila and related plants. drying of stems. Bacteria ooze from cut stems in
Erwinia nimipressuralis Wetwood of elm, viscid masses. Partially resistant plants may be
slime flux, due to bacteria pathogenic in elm dwarfed, with excessive blooming and
trunk wood, especially Asiatic elms, but possibly branching, wilting during the day but partially
occurring in many other trees, including maple, recovering at night. The bacteria winter solely
oak, mulberry, poplar, and willow. A water- in the digestive tract of the insects and are depos-
soaked dark discoloration of the heartwood is ited on leaves in spring with excrement, entering
correlated with chronic bleeding at crotches and through wounds or stomata.
wounds and abnormally high sap pressure in Control is directed chiefly at the insects. Start
trunk, with wilting a secondary symptom. The vines under Hotkaps and spray or dust with rote-
pressure in diseased trees increases from April none or other insecticide when the mechanical
Pseudomonadaceae 77

protection is removed. Experimental spraying terracmycin or streptomycin. Use resistant varie-

with antibiotics – streptomycin, terramycin, and ties such as Golden Cross Bantam, Carmel-cross,
neomycin has reduced wilt and increased yield. Ioana, Marcross, and Iochief.
Pantoea ananatis, Leaf Blight and Bulb
Decay of onion.
Pantoea herbicola (formerly Erwinia
herbicola). Leaf Spot of dracaena. On Dracaena Pseudomonadaceae
sanderana, gypsophila and related plants. Galls
are formed at crown and roots of grafted plants Pseudomonas
from 1/4 to 1 inch in diameter, but with a flat
nodular growth rather than the usual globose Motile with polar flagella; straight or curved rods;
crown gall. Gram-negative. Many species produce
Pantoea stewartii (formerly Erwinia stewartii). a greenish, water-soluble pigment. Many species
Bacterial Wilt of corn, Stewart’s Disease on sweet are found in soil and water; many are plant path-
corn, sometimes field corn, in the middle regions ogens causing leaf spots or blights.
of the United States, from New York to Califor- Acidovorax avenae formerly Pseudomonas
nia. This is a vascular disease with yellow slime albopreciptans). Bacterial Spot of cereals,
formed in the water-conducting system, resulting grasses, and corn. Light or dark brown spots or
in browning of nodes, and dwarfing of plants; or streaks on grass blades. Bacteria enter through
long pale green streaks on leaf blades, followed stomata or water pores.
by wilting and death of whole plant. Tassels may Acidovorax avenae (formerly Pseudomonas
be formed prematurely and die before the rest of avenae). Bacterial Leaf Spot of sweet corn. Bac-
the plant. The bacteria are chiefly disseminated terial Leaf Blight of johnsongrass.
by corn flea beetles, and winter either in the Acidovorax avenae subsp. citrulli (formerly
beetles or in seed. Primary infections come from Pseudomonas pseudoalcaligenes). Angular Leaf
flea beetles feeding in spring, from infected seed, Spot of muskmelon and watermelon. Fruit
and occasionally from soil; but secondary spread blotch; on melon, squash, pumpkin, and
is mostly by insects. watermelon.
Corn grown in rich soil is more susceptible to Acidovorax cattleyae (formerly Pseudomonas
wilt, and so are early varieties, especially Golden cattleyae). Brown Spot of orchids, Phalaenopsis
Bantam. Winter temperatures influence the and Cattleya, common in greenhouses. Infection
amount of wilt the following summer. If the win- is through stomata of young plants, wounds of
ter index, which is the sum of mean temperatures older plants. Dark green, circular water-soaked
for December, January, and February, is above spots change to brown and finally black. On Cat-
100, bacterial wilt will be present in destructive tleya the disease is limited to older leaves.
amounts on susceptible varieties. If the index is Burkholderia andropogonis (formerly Pseu-
below 90, the disease will be very sparse in domonas and ropogonis). Bacterial Stripe of sor-
northeastern states; if the index is between 90 ghum and corn. Bacterial Leaf Spot on
and 100, there will be a moderate amount of bougainvillea. Black Spot on clovers and statice.
wilt. Disease surveys over a period of years tes- Also causes blight of chickpea, and bacterial leaf
tify to the reliability of such forecasts (based on spot on white clover. Red streaks and blotches
the amount of cold the flea beetle vectors can appear on leaves and sheaths, with abundant exu-
survive); but with the increasing use of hybrid date drying down to red crusts or scales, readily
sweet corn resistant to wilt, the importance of washed off in rains. Bacteria enter through
winter temperatures is reduced. stomata.
Control Use insecticides to control flea beetles; Bacterial Leaf Spot of velvet bean, clovers.
substitute commercial fertilizer for manure; Translucent angular brown leaf spots have lighter
destroy infected refuse; try treating seed with centers and chlorotic surrounding tissue; there is
78 Bacterial Diseases

no exudate. Bacteria enter through stomata and nearly black, eventually sunken with raised,
fill intercellular spaces of parenchyma. horny, or brittle margins that are scablike and
Burkholderia caryophylli formerly Pseudomo- exude a gummy substance. Bacteria overwinter
nas caryophylli). Bacterial Wilt of carnation, usu- on corms. First symptoms after planting are tiny
ally under glass. Plants wilt, turn dry, colorless reddish raised specks on leaves, mostly near the
with roots disintegrating. Grayish-green foliage base, enlarging to dark sunken spots, which grow
is the first symptom, but leaves rapidly turn together into large areas with a firm or soft rot.
yellow and die. Yellow streaks of frayed tissue Sometimes plants fall over, but the disease is not
in vascular areas extend a foot or two up the stem. ordinarily very damaging in the garden. The chief
It takes a month for disease to show up after loss is to the grower in disfigured, unsalable
inoculation, but it can be transmitted on cuttings corms. Brown streaks in husks sometimes disin-
taken from plants before appearance of symp- tegrate, leaving holes.
toms. The sticky character of diseased tissue dis- Gladiolus scab is increased by bulb mites, may
tinguishes this wilt from Fusarium wilt. Varieties be related to grub and wireworm injury.
Cardinal Sim, Laddie, Mamie, Portrait, and Pseudomonas aceris (see Pseudomonas
others may have severe cankers at base of syringae pv. aceris). Maple Leaf Spot found in
stems, orange-yellow when young, very sticky. California on big leaf maple.
Bacteria are spread by hands, tools, splashing Pseudomonas adzukicola Stem Rot of adzuki
water. Also causes crown and leaf rot of statice. bean.
Control Remove and burn diseased plants and Pseudomonas albopreciptans (see Acidovorax
all within 1 1/2 -foot radius. After handling wash avenae). Bacterial Spot of cereals, grasses, and
with hot water and soap, sterilize tools (10 % corn.
Clorox for 5 min). Obtain rooted cuttings from Pseudomonas and ropogonis (see
propagators of cultured, disease-free material; Burkholderia and ropogonis). Bacterial Stripe
keep in shipping bags until ready for benching of sorghum and corn.
and then place in raised, steam-pasteurized Bacterial Leaf Spot of velvet bean, clovers.
benches. Never place cuttings in water or Translucent angular brown leaf spots have lighter
a liquid fungicide (use dust if a fungicide is centers and chlorotic surrounding tissue; there is
required for other diseases); never place tempo- no exudate. Bacteria enter through stomata and
rarily on an unsterilized table; never cut or trim fill intercellular spaces of parenchyma.
with hands or knives; never plant in outdoor Pseudomonas angulata (see Pseudomonas
“nurse beds”; never use overhead watering. syringae pv. angulata). Blackfire of tobacco.
Burkholderia cepacia (formerly Pseudomonas Pseudomonas asplenii Bacterial Leaf Blight of
cepacia). Sour Skin Rot of onion. Slimy yellow bird’s-next fern, first reported from greenhouses
rot of outer fleshy scales, with a vinegar odor. Let in California. Small translucent spots enlarge to
crop mature well before harvesting, tops dry cover whole frond; bacteria may invade crown
before topping; cure bulbs thoroughly before and kill whole plant. Control depends on strict
storage. sanitation – sterilizing flats, pots, media, and
Burkholderia gladioli (formerly Pseudomonas foreceps used in transplanting. Avoid excessive
gladioli). Leaf Spot and Blight on bird’s nest fern. watering and too high humidity.
Onion Bulb Rot, a storage disease, inner scales Pseudomonas avenae (see Acidovorax avenae).
of bulb water-soaked and soft, sometimes entire Bacterial Leaf Spot of sweet corn. Bacterial Leaf
bulb rotting. Blight of johnsongrass.
Orchid Brown Rot and Leaf Spot. Pseudomonas berberidis Bacterial Leaf Spot
Gladiolus Scab, Stem Rot, Neck Rot, wide- of barberry. Small, irregular, dark green water-
spread on gladiolus, also on iris, bell peppers and soaked areas on leaves turn purple-brown with
tigridia. Lesions on corms are pale yellow, water- age; occasional spotting occurs on leaf stalks and
soaked circular spots deepening to brown or young shoots. If twigs are infected, buds do not
Pseudomonadaceae 79

Pseudomonas melophthora Apple Rot, proba-

bly widespread. This is a decay of ripe apples
following after apple maggots and eventually
rotting whole fruit.
Pseudomonas pseudoalcaligenes (see
Acetovorax avenae subsp. citrulli). Angular
Leaf Spot of muskmelon and watermelon. Fruit
Blotch on melon, squash, pumpkin, and
Pseudomonas ribicola On golden currant in
Fig. 3 Bacterial Black Spot on Chrysanthemum Pseudomonas sesami Bacterial Leaf Spot of
sesame. Brown spots on leaves and stems. Can
develop in the next season; if they are girdled, the be controlled by treating seed with streptomycin.
entire twig is blighted. Cut out infected twigs and Pseudomonas solanacearum (see Rolstonia
spray with bordeaux mixture or an antibiotic. solanacearum). Southern Bacterial Wilt, also
Pseudomonas caryophylli (see Burkholderia called Brown Rot, Bacterial Ring Disease, Slime
caryophylli). Bacterial Wilt of carnation, usually Disease, Granville Wilt (of tobacco), present in
under glass. many states but particularly prevalent in the
Pseudomonas cattleyae (see Acidovorax South, from Maryland around the coast to Texas.
cattleyae). Brown Spot of orchids, Phalaenopsis Pseudomonas syringae Canker on kiwifruit;
and Cattleya, common in greenhouses. also Blight on impatiens and mock orange. Also
Pseudomonas cepacia (see Burkholderia Leaf Spot on English and American elm, moun-
cepacia). Sour Skin Rot of onion. Slimy yellow tain-laurel, arugula and coriander. Stem Dieback
rot of outer fleshy scales, with a vinegar odor. of Centaurea and fennel.
Pseudomonas cichorii Bacterial Leaf Spot on Pseudomonas syringae pv. aceris (formerly
basil. Bacterial Blight on Lobelia. Pseudomonas aceris). Maple Leaf Spot found in
Pseudomonas cichorii Bacterial Rot of chicory, California on big leaf maple. Small, water-soaked
Belgium endive, French endive, iris, Soft Rot of spots, surrounded by yellow zones, turn brown or
potato, and Bacterial Leaf Spot of hibiscus, gera- black; cankers develop on petioles and bracts in
nium, magnolia and rhododendron. May also serious cases; leaves may drop; disease present in
cause a Leaf Spot and Stem Necrosis on chrysan- cool, damp weather of early spring.
themum (see Fig. 3) and Bacterial Leaf Blight on Pseudomonas syringae pv. angulata (formerly
dwarf Schefflera. A yellowish olive center rot, Pseudomonas angulata).
affecting young inner leaves. Blackfire of tobacco.
Pseudomonas corrugata Stem Rot of tomato, Pseudomonas syringae pv. apii Bacterial
also Pith Necrosis. Blight of celery. Small, irregularly circular rusty
Pseudomonas fluorescens (marginalis). Mar- leaf spots, with a yellow halo, are occasionally
ginal Blight of lettuce, Kansas Lettuce Disease, numerous enough to cause death of foliage, but
also on witloof chicory, Soft Rot of potato tubers. commonly are only disfiguring. Spray plants in
Leaf margins are dark brown to almost black, first seedbed with bordeaux mixture, or dust with cop-
soft, then like parchment. Yellowish red spots, per lime dust; clean up old refuse.
turning dark, are scattered over leaves. Infected Pseudomonas syringae pv. apii Bacterial Leaf
tissue disintegrates into an odorous mass. Bacte- Spot of celery.
ria live in the soil, which should not be splashed Pseudomonas syringae pv. aptata Bacterial
on plants by careless watering. Spot on beets, Swiss chard, and nasturtium.
Pseudomonas gladioli (see Burkholderia glad- Spots on nasturtium leaves are water-soaked,
ioli). Leaf Spot and Blight on bird’s nest fern. brownish, 1/8 to 1/4 inch across. On beets they
80 Bacterial Diseases

are dark brown or black, irregular, and in addition muskmelon, summer squash, occasional on
there are narrow streaks on petioles, midribs, and other cucurbits. Leaves or stems have irregular,
larger veins. Petiole tissue may be softened as angular, water-soaked spots with bacteria oozing
with soft rot. Infection is only through wounds. out in tearlike droplets that dry down to a white
Pseudomonas syringae pv. coronafa- residue. Eventually the spots turn gray, die, and
ciens Halo Blight on grasses, such as Poa shrink, leaving holes in foliage. Fruit spots are
spp. and Calamagrostis spp. small, nearly round, with the tissue turning white,
Pseudomonas syringae pv. delphinii - sometimes cracking. The bacteria overwinter in
Delphinium Black Spot on delphinium and aco- diseased plant tissue and in the seed coat. They
nite (monkshood). Irregular tarry black spots on are spread from soil to stems and later to fruit in
leaves, flower buds, petioles, and stems may coa- rainy weather, also transferred from plant to plant
lesce in late stages to form large black areas. The on hands and clothing. Infection is most severe in
bacteria enter through stomata or water pores. plants gone over by pickers early in the morning
Occasionally this bacterial leaf spot results in before dew has dried off.
some distortion, but most abnormal growth and Control Plow under or remove vines immedi-
blackening of buds is due to the cyclamen mite, ately after harvest.
a much more important problem than black spot. Pseudomonas syringae pv. mori Bacterial
Control Remove diseased leaves as noticed; cut Blight of mulberry, general on black and white
and burn all old stalks at end of season; avoid mulberry. Numerous water-soaked leaf spots join
overhead watering. In a wet season spraying with to form brown or black areas with surrounding
bordeaux mixture may have some value. yellow tissue. Young leaves may be distorted,
Pseudomonas syringae pv. glycinea Bacterial with dark sunken spots on midribs and veins.
Blight of soybean. Perhaps the most common Dark stripes with translucent borders on young
and conspicuous disease of soybean, appearing shoots exude white or yellow ooze from lenticels.
in fields when plants are half-grown and Dead twigs and brown leaves resemble fire
remaining active until maturity, with defoliation blight; trees are stunted but seldom killed.
during periods of high humidity or heavy dews. Remove and burn blighted branches; do not
Small, angular, translucent leaf spots, yellow to plant young mulberry trees near infected
light brown, turn dark reddish brown to nearly specimens.
black with age. There is often a white exudate Pseudomonas syringae pv. mors-prunorum -
drying to a glistening film on under leaf surfaces. Bacterial Canker of stone fruits, Citrus Blast, Lilac
Black lesions appear on stems and petioles, and Blight on many unreleated plants, including
on pods water-soaked spots enlarge to cover apple, plum, peach, cherry, pear, almond, avo-
a wide area, darken, and produce an exudate cado, citrus fruits, lilacs, flowering stock, rose,
drying to brownish scales; seeds are often beans, cowpeas, oleander, and leaf spot on peas.
infected. Seedlings from infected seed have On stone fruits all plant parts are subject to
brown spots on cotyledons and often die. Flam- attack, but most destructive are elongated water-
beau and Hawkeye varieties are somewhat less soaked lesions or gummy cankers on trunks and
susceptible. Use seed taken from disease-free branches, usually sour-smelling. Dormant buds
pods. of cherry and apricot are likely to be blighted,
Pseudomonas syringae pv. helianthi Bacterial pear blossoms blasted. Small purple spots appear
Leaf Spot of sunflower. Leaves show brown, on leaves of plum and apricot, black lesions on
necrotic spots, first water-soaked, then dark and fruit of cherry and apricot. All varieties of apricot
oily. are very susceptible to the disease. Plums on
Pseudomonas syringae pv. hibisci Bacterial Myrobalan rootstock are more resistant, and vari-
Leaf Spot on Hibiscus. eties California, Duarte, and President are toler-
Pseudomonas syringae pv. lachrymans Angular ant. On citrus, and particularly lemons, dark
Leaf Spot of cucurbits, general on cucumber, sunken spots, called black pit, are formed on
Pseudomonadaceae 81

fruit rind, but there is no decay. The blast form of Pseudomonas syringae pv. pisi Bacterial Blight
the disease is most often on oranges and of pea, general on field and garden peas, espe-
grapefruit – water-soaked areas in leaves, which cially in East and South, and causing a leaf spot of
may drop or hang on, twigs blackened and shriv- sweet peas. Dark green water-soaked dots on
eled. The disease is most serious in seasons with leaves enlarge and dry to russet brown; stems
cold, driving rainstorms. have dark green to brown streaks. Flowers are
On lilac, brown water-soaked spots on leaves killed or young pods shriveled, with seed covered
and internodes on young shoots blacken and rap- with bacterial slime. Bacteria enter through sto-
idly enlarge. Young leaves are killed; older mata or wounds, and if they reach the vascular
leaves have large portions of the blade affected. system, either leaflets or whole plants wilt. Vines
Infection starts in early spring in rainy weather. infected when young usually die. Alaska and
The bacteria are primarily in the parenchyma, Telephone varieties are particularly susceptible.
spreading through intercellular spaces, blacken- Control Avoid wounding vines during cultiva-
ing and killing cells, forming cavities. The vas- tion. Sow peas in early spring in well-drained
cular system may also be affected, followed by soil. Use disease free seed and plan a 4-year
wilting of upper leaves. rotation.
Control Prune out infected twigs and branches. Pseudomonas syringae pv. porri Bacterial
In California spray fruits in fall with bordeaux Blight of shallot.
mixture, at the time first leaves are dropping. Pseudomonas syringae pv. primulae Bacterial
Grow bushy, compact citrus trees less liable to Leaf Spot of primrose in ornamental and com-
wind injury; use windbreaks for orchards. mercial plantings in California. Infection is con-
Pseudomonas syringae pv. papulans Blister fined to older leaves – irregularly circular brown
Spot of apple. Small, dark brown blisters on lesions surrounded by conspicuous yellow halos.
fruit and rough bark cankers on limbs start at Spots may coalesce to kill all or part of leaf.
lenticels. Bark may have rough scaly patches Spraying with bordeaux mixture has prevented
from a few inches to a yard long, bordered with infection.
a pimpled edge, and with outer bark sloughing off Pseudomonas syringae pv. savastanoi Olive
in spring. Knot, Bacterial Knot of olive. Irregular, spongy,
Pseudomonas syringae pv. phaseolicola Bean more or less hard, knotty galls on roots, trunk,
Halo Blight, halo spot on common, lima, and branches, leaf, or fruit pedicels start as small
scarlet runner beans. The symptoms are those of swellings and increase to several inches with
other bean blights except that there are wide irregular fissures. Terminal shoots are dwarfed
green or yellowish green halos around water- or killed; whole trees may die. Bacteria enter
soaked leaf spots, such spots later turning brown through wounds, often leaf scars or frost cracks.
and dry. Leaves wilt and turn brown; young pods Variety Manzanilla is most susceptible of the
wither and produce no seed; sometimes plants are olives commonly grown in California. Another
dwarfed with top leaves crinkled and mottled. In form of this species causes similar galls on ash.
hot weather, spots are often angular, reddish Control Cut out galls carefully, disinfesting
brown, and without halo. Stem streaks are red- tools; paint larger cuts with bordeaux paste and
dish, with gray ooze; pod spots are red to brown spray trees with bordeaux mixture in early
with silver crusts; seeds are small, wrinkled, with November, repeating in December and March if
cream-colored spots. All snap beans are suscep- infection has been abundant. Do not plant
tible; many dry beans – Pinto, Great Northern, infected nursery trees or bring equipment from
Red Mexican, Michelite – are rather resistant. an infected orchard into a healthy one.
Control Use seed from blight-free areas. Blight Pseudomonas syringae pv. syringae Brown
is rare in California, occasional in Idaho. Plan Spot, Foliar on wild rice (Zizania); leaf spot and
a 3-year rotation. Do not pick beans when foliage stem collapse on urd bean; leaf spot and stem
is wet. canker on Ginkgo.
82 Bacterial Diseases

Oleander Bacterial Gall. Galls or tumors are Control Change location of hotbed starting
formed on branches, herbaceous shoots, leaves, seedlings; use 2-year rotation in field; have seed
and flowers but not on underground parts. Small hot-water treated.
swellings develop on leaf veins, surrounded by Pseudomonas syringae pv. tomato Bacterial
yellow tissue, with bacterial ooze coming from Speck of tomato. Numerous, dark brown raised
veins in large quantity. Young shoots have longi- spots on fruit are very small, less than 1/16 inch;
tudinal swellings with small secondary tubercles; they do not extend into flesh and are more
young leaves and seedpods may be distorted and disfiguring than harmful.
curled. On older branches tumors are soft or Pseudomonas syringae pv. zizaniae Leaf Spot
spongy and roughened with projecting tubercles; and Stem Spot of wild rice.
they slowly turn dark. Prune out infected por- Pseudomonas tabaci (see Pseudomonas
tions, sterilizing shears between cuts; propagate syringae pv. tabaci). Blackfire of tobacco.
only from healthy plants. Pseudomonas viburni Bacterial Leaf Spot of
Pseudomonas syringae pv. tabaci (see Pseudo- viburnum, widespread. Circular water-soaked
monas tabaci). Blackfire of tobacco. spots appear on leaves, and irregular sunken
Pseudomonas syringae pv. tabaci Tobacco brown cankers on young stems, and the bacteria
Wildfire on tobacco, tomato, eggplant, soybean, overwinter in leaves, stems or buds. Remove and
cowpea, pokeberry, and ground-cherry, in all burn infected leaves. Spray with bordeaux mix-
tobacco districts sporadically. Leaf spots have ture or an antibiotic such as Agrimycin two or
tan to brown dead centers with chlorotic halos. three times at weekly intervals.
The disease appears first on lower leaves and Pseudomonas viridiflava Bacterial Leaf Spot
spreads rapidly in wet weather. The bacteria per- on basil. Bacterial Canker on poinsettia.
sist a few months in crop refuse and on seed and Pseudomonas viridilivida Louisiana Lettuce
enter through stomatal cavities. In buried soybean Disease on lettuce, bell pepper, and tomatoes.
leaves the bacteria have lived less than 4 months; Numerous water-soaked leaf spots fuse to infect
so fall plowing may be beneficial. Seed stored for large areas, first with a soft rot, then a dry shriv-
18 months produces plants free from wildfire. eling. Sometimes outer leaves are rotted and the
Pseudomonas syringae pv. tagetis Bacterial heart sound. This bacterium also causes greasy
Leaf Spot. Circular necrotic lesions on leaves and canker of poinsettia.
petioles. The lesions have dark purple margins. This Pseudomonas washingtoniae This bacterium
disease occurs on marigold, sunflower, Jerusalem causes spots on leaves of Washington palm.
artichoke, and common ragweed. Apical chlorosis Pseudomonas woodsii Bacterial Spot and
is also caused by this pathogen on sunflower and Blight of carnation. Leaf lesions are small,
sunflower seed may be a source of inoculum. elongated, brown with water-soaked
Pseudomonas syringae pv. tagetis Bacterial borders, withering to brown sunken areas, with
Leaf Spot on compass plant and sunflower. masses of bacteria oozing out of stomata. They
Pseudomonas syringae pv. tomato Bacterial are spread in greenhouses by syringing, and out-
Leaf Spot of crucifers, Pepper Spot of cabbage, doors by rain. Follow cultural practices suggested
cauliflower, Chinese cabbage, and turnip, mostly under P. caryophylli for carnation wilt.
in northeastern and Middle Atlantic states. Pseudomonas sp. Blueberry Canker reported
Numerous brown or purple spots range from pin- from Oregon. Reddish brown to black cankers
point to 1/8 inch in diameter. If spots are very appear on canes of the previous season; all buds in
numerous, leaves yellow and drop off. Cauli- the cankered areas are killed; stems are sometimes
flower is more commonly affected than cabbage. girdled. Varieties Weymouth, June, and Rancocas
Bacteria, disseminated on seed or in diseased are resistant, but Jersey, Atlantic, Scammel,
plant parts, enter through stomata, and visible Coville, and Evelyn are highly susceptible.
symptoms appear in 3 to 6 days. Disease is most Rhizomonas suberifaciens Corky Rot on
severe in seedbeds. lettuce.
Xanthomonas 83

Rolstonia solanacearum (formerly Pseudomo- Ralstonia solanacearum, Race 3, Biovar 2.

nas solanacearum). Southern Bacterial Wilt, also Bacterial wilting of geranium; also yellowing and
called Brown Rot, Bacterial Ring Disease, Slime stunting.
Disease, Granville Wilt (of tobacco), present in Xanthomonas albilineans Leaf Scald of sugar-
many states but particularly prevalent in the South, cane (FL, TX).
from Maryland around the coast to Texas. South- Xanthomonas axonopodis Leaf Streak (water
ern wilt is common on potatoes in Florida but also soaking) of African lily and Leaf Blight of onion.
appears on many other vegetables – bean, lima Xanthomonas begoniae (see Xanthomonas
bean, castor bean, soybean, velvet bean, beet, car- campestris pv. begoniae). Begonia Bacteriosis,
rot, cowpea, peanut, sweet potato, tomato, egg- leaf spot of fibrous and tuberous begonias.
plant, pepper, and rhubarb. Ornamentals Xanthomonas campestris Black Rot of
sometimes infected include ageratum, anthurium, cruciers, Bacterial Blight, Wilt, Stump Rot of
dwarf banana, garden balsam, geranium, canna, alder, arabidopsis, asparagus tree fern, avocado,
cosmos, croton, chrysanthemum, dahlia, holly- cabbage, cauliflower, broccoli, brussels sprouts,
hock, lead-tree, marigold, nasturtium, Spanish kale, lavender, mustard, radish, rutabaga, sun-
needle, sunflower, and zinnia. The symptoms are flower, stock, turnip, and leaf blight of onion.
those of a vascular disease, with dwarfing or sud- Black rot was first observed in Kentucky and
den wilting, a brown stain in vascular bundles, and Wisconsin about 1890 and is generally distrib-
dark patches or streaks in stems. Often the first uted in the country, with losses often 40 to 50 %
symptom is a slight wilting of leaves at end of of the total crop. It is one of the most serious
branches in the heat of the day, followed by recov- crucifer diseases, present each season but epi-
ery at night, but each day the wilting is more demic in warm, wet seasons.
pronounced and recovery less until the plant dies. The bacteria invade leaves through water
Young plants are more susceptible than older ones. pores or wounds and progress to the vascular
In potatoes and tomatoes there may be a brown system. Veins are blackened, with leaf tissue
mushy decay of stems, with bacterial ooze present. browning in a V-shape. With early infection
Potato tubers often have a browning of vascular plants either die or are dwarfed, with a one-
ring, followed by general decay. sided growth. Late infection results in defolia-
Bacteria live in fallow soil 6 years or more and tion, long bare stalks with a tuft of leaves on
may persist indefinitely in the presence of sus- top. When stems are cut across, they show
ceptible plants. They are spread by irrigation a black ring, result of the vascular invasion, and
water, in crop debris, or soil fragments on tools sometimes yellow bacterial ooze. Black rot is
and tractors, or by farm animals. Optimum tem- a hard odorless rot, but it may be followed by
peratures are high, ranging from 77 to 97  F, soft, odorous decays. Primary infection comes
with inhibition of disease below 55  F. from bacteria carried on seed, or in refuse in
Control Use northern-grown seed potatoes and soil, but drainage water, rain, farm implements,
Sebago and Katahdin varieties, more resistant and animals aid in secondary infection.
than Triumph and Cobbler. Use a long rotation Control Use seed grown in disease-free areas in
for tomatoes. Soil can be acidified with sulfur to the West or treat with hot water, 122  F, 25 min
kill bacteria, followed by liming in the fall before for cabbage, 18 min for broccoli, cauliflower, and
planting. collards. Plan a 3-year rotation with plants other
than crucifers, and clean up all crop refuse.
Xanthomonas campestris Horse-Radish Leaf
Xanthomonas Spot. Leaves are spotted but there is no vascular
infection. Also causes leaf spot of Pilea sp.,
Small rods, motile with a single polar flagellum; Pellionia sp. and leaf spot and blight of bird of
form abundant slimy yellow growth. Most spe- paradise, white butterfly. Also, bacterial leaf and
cies are plant pathogens causing necroses. stem lesions.
84 Bacterial Diseases

Xanthomonas campestris Bacterial Leaf Spot precautions so rigid they included walking the
on cabbage and radish. mules through disinfestant, sterilization of
Xanthomonas campestris pv. asclepiadis - clothes worn by workers – ill saved us from
Bacterial Blight on butterfly weed. untold later losses.
Xanthomonas campestris pv. barbareae Black Symptoms of citrus canker are rough, brown
Rot of winter-cress (Barbarea vulgaris), similar corky eruptions on both sides of leaves and fruit.
to black rot of cabbage; small greenish spots turn On foliage the lesions are surrounded by oily or
black. yellow halos. Old lesions become brown and
Xanthomonas campestris pv. begoniae (for- corky.
merly Xanthomonas begoniae). Begonia Xanthomonas campestris pv. corylina (for-
Bacteriosis, leaf spot of fibrous and tuberous merly Xanthomonas corylina). Filbert Blight,
begonias. Blister-like, roundish dead spots are Bacteriosis, the most serious disease of filberts
scattered over surface of leaves. Spots are in the Pacific Northwest, known since 1913 from
brown with yellow translucent margins. Leaves the Cascade Mountains west in Oregon
fall prematurely, and in severe cases the main and Washington. The disease is similar to walnut
stem is invaded, with gradual softening of all blight (see X. juglandis) with infection on
tissues and death of plants. Bacteria remain via- buds, leaves, and stems of current growth; on
ble at least 3 months in yellow ooze on surface of branches; and on trunks 1 to 4 years old. The
dried leaves. Leaves are infected through upper bacteria are weakly pathogenic to the nuts.
surfaces during watering, with rapid spread of Copper-lime dusts are effective, with four to six
disease when plants are crowded together under weekly applications, starting at the early
conditions of high humidity. prebloom stage.
Control Keep top of leaves dry, avoiding syring- Xanthomonas campestris pv. cucurbitae (for-
ing or overhead watering; keep pots widely merly Xanthomonas cucurbitae). Bacterial Spot
spaced; spray with bordeaux mixture and dip on winter squash and pumpkin. Leaf spots are
cuttings in it. first small and round, then angular between
Xanthomonas campestris pv. carotae (for- veins, with bright yellow halos; sometimes trans-
merly Xanthomonas carotae). Bacterial Blight lucent and thin but not dropping out; often coa-
of carrot. The chief damage is to flower heads lescing to involve whole leaf. Bacterial exudate is
grown for seed, which may be entirely killed. present.
Symptoms include irregular dead spots on leaves, Xanthomonas campestris pv. cyamopsidis Rot
dark brown lines on petioles and stems, blighting of Lithops spp.
of floral parts, which may be one-sided. Use clean Xanthomonas campestris pv. dieffenba-
seed, or treat with hot water; rotate crops. chiae Blight of Anthurium; also Leaf Spot of
Xanthomonas campestris pv. citri Citrus Can- cocoyam.
ker on all citrus fruits, but not apparently eradi- Xanthomonas campestris pv. dieffenbachiae
cated from the United States. It came from the (formerly Xanthomonas dieffenbachiae). Dief-
Orient and appeared in Texas in 1910, becoming fenbachia Leaf Spot. Spots are formed on all
of major importance in Florida and the Gulf parts of leaf blade except midrib, but not on
States by 1914, ranking with chestnut blight and petioles and stems. They range from minute,
white pine blister rust as a national calamity. But translucent specks to lesions 3/8 inch in diameter,
here is one of the few cases on record where man circular to elongated, yellow to orange-yellow
has won the fight, where a disease has been nearly with a dull green center. Spots may grow together
eradicated by spending enough money and hav- to cover large areas, which turn yellow, wilt, and
ing enough cooperation early in the game. Sev- dry. Dead leaves are dull tan to light brown, thin
eral million dollars, together with concerted and tough but not brittle. The exudate on lower
intelligent effort by growers, quarantine mea- surface of spots dries to a waxy, silver-white
sures, destruction of every infected tree, sanitary layer.
Xanthomonas 85

Control Separate infected from healthy plants; The disease was first noted in Holland in 1881
keep temperature low; avoid syringing; try pro- and named for the yellow slime or bacterial ooze
tective spraying with streptomycin. seen when a bulb is cut. The bulbs rot either
Xanthomonas campestris pv. before or after planting, producing no plants
fragariae Angular Leaf Spot on strawberry; above ground or badly infected specimens,
also Blossom Blight on strawberry. which do not flower and have yellow to brown
Xanthomonas campestris pv. glycines (for- stripes on leaves or flower stalks. Bacteria are
merly Xanthomonas glycines (phaseoli var. transmitted by wind, rain, tools, and clothes,
sojense). Bacterial Pustule of soybean, similar with rapid infection in wet or humid weather,
to regular bean blight but chiefly a foliage dis- particularly among luxuriantly growing plants.
ease, present in most soybean areas, more severe The disease is usually minor in our Pacific North-
in the South. Small, yellow-green spots with red- west but worse in warm, wet weather on rapidly
dish brown centers appear on upper surface of growing plants. Innocence is more susceptible
leaves with a small raised pustule at the center of than King of the Blues.
the spot on the under leaf surface. Spots run Control Cover infected plants with a jar or can
together to large irregular brown areas, portions until the end of the season; then dig after the
of which drop out, giving a ragged appearance. others. Never work or walk in fields when plants
Bacteria overwinter in diseased leaves and on are wet; avoid bruising; discard rotten bulbs;
seed. Variety CNS is highly resistant; Ogden has rotate plantings; avoid fertilizer high in nitrogen.
some resistance. Xanthomonas campestris pv. incanae (for-
Xanthomonas campestris pv. gummisudans merly Xanthomonas incanae). Bacterial Blight
(formerly Xanthomonas gummisudans). Bacterial of garden stocks causing, since 1933, serious
Blight of Gladiolus. Narrow, horizontal, water- losses on flower-seed ranches in California; also
soaked, dark green spots turn into brown squares present in home gardens. This is a vascular dis-
or rectangles between veins, covering entire leaf, ease of main stem and lateral branches, often
particularly a young leaf, or middle section of the extending into leaf petioles and seed peduncles.
blade. Bacteria ooze out in slender, twisted, white Seedlings suddenly wilt when 2 to 4 inches high,
columns or in a gummy film, in which soil and with stem tissues yellowish, soft and mushy, and
insects get stuck. Disease is spread by planting sometimes a yellow exudate along stem. On older
infected corms or by bacteria splashed in rain plants, dark water-soaked areas appear around
from infected to healthy leaves. The small dark leaf scars near ground, stem is girdled, and
brown corm lesions are almost unnoticeable. lower leaves turn yellow and drop; or entire
Soak corms unhusked for 2 h before planting. plants wilt or are broken by wind at ground
Xanthomonas campestris pv. hederae (for- level. Bacteria persist in soil and on or in seed;
merly Xanthomonas hederae). Bacterial Leaf they are also spread in irrigation water.
Spot of English ivy. Small water-soaked area on Control Use a 2 to 3-year rotation. Treat seed
leaves develop dark brown to black centers as with hot water, 127.5 to 131  F for 10 min,
they increase in size, sometimes cracking, with followed by rapid cooling.
reddish purple margins. Spots are sometimes Xanthomonas campestris pv. juglandis -
formed on petioles and stems, with plants formerly Xanthomonas juglandis). Walnut Blight
dwarfed and foliage yellow-green. Spray with on English or Persian walnut, black walnut, but-
bordeaux mixture or an antibiotic. Keep plants ternut, Siebold walnut. Black, dead spots appear
well spaced; avoid overhead watering and high on young nuts, green shoots, and leaves. Many
humidity. nuts fall prematurely, but others reach full size
Xanthomonas campestris pv. hyacinthi (for- with husk, shell, and kernel more or less black-
merly Xanthomonas hyacinthi). Hyacinth Yel- ened and destroyed. Bacteria winter in old nuts
lows, yellow rot of Dutch hyacinth, occasionally or in buds, and may be carried by the walnut
entering the country in imported bulbs. erinose mite.
86 Bacterial Diseases

Control Spray with a fixed copper, as copper purchase culture-indexed cuttings. Be sure to
oxalate, or with streptomycin. Apply when 10 % sterilize cutting knives. Use 1-year rotation. Try
of the blossoms are open, repeat when 20 % are Agrimycin as a preventive spray, or copper.
open, and again after bloom. Xanthomonas campestris pv. pelargonii (for-
Xanthomonas campestris pv. merly Xanthomonas pelargonii) Geranium Leaf
malvacearum Leaf Spot on Hibiscus. Spot on Pelargonium spp. Leaf spots are small,
Xanthomonas campestris pv. oryzae (for- brown, necrotic, sometimes with reddish tinge on
merly Xanthomonas oryzae). Carnation Pimple upper surface and a slightly water-soaked condi-
reported from Colorado as caused by a new form tion on underside. Young leaves may die and
of the rice blight organism. Very small, 1 mm, drop. Petioles are occasionally spotted. Bacteria
pimples are formed near base and tips of leaves, winter in old leaves or under mulch.
which may shrivel. Xanthomonas campestris pv. phaseoli (for-
Xanthomonas campestris pv. papavericola merly Xanthomonas phaseoli). Bacterial Bean
(formerly Xanthomonas papavericola). Bacterial Blight, general and serious on beans but rare in
Blight of poppy on corn poppy and on Oriental, some western states. Leaf spots are at first very
opium, and California poppies. Minute, water- small, water-soaked or light green wilted areas,
soaked areas darken to intense black spots which enlarge, turn brown, are dry and brittle,
bounded by a colorless ring. Spots are scattered, and have a yellow border around edge of lesions
circular, small, often zonate, with tissue between and often a narrow, pale green zone outside that.
yellow and then brown. There is a noticeable, Leaves become ragged in wind and rainstorms.
slimy exudate. Infection is through stomata and Reddish brown horizontal streaks appear in stem,
often into veins. Stem lesions are long, very which may be girdled and break over at cotyle-
black, sometimes girdling and causing young dons or first leaf node.
plants to fall over. Flower sepals are blackened, Pod lesions are first dark green and water-
petals stop developing; pods show conspicuous soaked, then dry, sunken and brick red, some-
black spots. times with a yellowish encrustation of bacterial
Control Remove and destroy infected plants; do ooze. White seeds turn yellow, are wrinkled with
not replant poppies in the same location. Try a varnished look.
Agrimycin as a preventive spray. Control Use disease-free western-grown seed.
Xanthomonas campestris pv. pelargonii (for- Keep away from beans when plants are wet.
merly Xanthomonas pelargonii). Bacterial Leaf Xanthomonas campestris pv. pruni (formerly
Spot of geranium (Pelargonium). Irregular to Xanthomonas pruni). Bacterial Spot of stone
circular brown leaf spots start as water-soaked fruits, also called canker, shot hole, black spot;
dots on undersurface, becoming sunken as they general on plum, Japanese plum, prune, peach,
enlarge and with tissue collapsing. If spots are and nectarine east of the Rocky Mountains; one
numerous, the entire leaf turns yellow, brown, of the more destructive stone fruit diseases, caus-
and shriveled, then drops. The leaves sometimes ing heavy losses in some states.
wilt and droop but hang on the plant for a week or Symptoms on leaves are numerous, round or
so. Exterior of stem is gray and dull, the pith and angular, small reddish spots with centers turning
cortex black, later disintegrating into a dry rot. brown and dead, dropping out to leave shot holes.
The roots are blackened but not decayed. Cut- Spots may run together to give a burned, blighted,
tings fail to root, and rot from the base upward. or ragged appearance, followed by defoliation,
Bacteria can live 3 months in moist soil; are with losses running high in devitalized trees. On
spread by handling, splashing water, cutting twigs dark blisters dry out to sunken cankers.
knives, and whiteflies. Fruit spots turn into brown to black, saucer-
Control Remove diseased plants. Take cuttings shaped depressions with small masses of
from plants known to be healthy; place in steril- gummy, yellow exudate, often with cracking
ized media and pots. Commercial growers should through the spot.
Xanthomonas 87

Control Plant new orchards from nurseries free Xanthomonas carotae (see Xanthomonas
from the disease. Prune to allow air in the interior campestris pv. carotae). Bacterial Blight of carrot.
of trees. Feed properly; trees with sufficient nitro- Xanthomonas corylina (see Xanthomonas
gen do not defoliate so readily. Zinc sulfate-lime campestris pv. corylina). Filbert Blight,
sprays have been somewhat effective. Bacteriosis, the most serious disease of filberts
Xanthomonas campestris pv. raphani (for- in the Pacific Northwest, known since 1913 from
merly Xanthomonas vesicatoria var. raphani). the Cascade Mountains west in Oregon and
Leaf Spot of radish, turnip, and other crucifers, Washington.
similar to bacterial spot on tomato. Xanthomonas cucurbitae (see Xanthomonas
Xanthomonas campestris pv. vesicatoria (for- campestriis pv. cucurbitae). Bacterial Spot on
merly Xanthomonas vesicatoria). Bacterial Spot winter squash and pumpkin
of tomato and pepper, common in wet seasons. Xanthomonas dieffenbachiae (see
Small, black, scabby fruit spots, sometimes with Xanthomonas campestriis pv. dieffenbachiae).
a translucent border, provide entrance points for Dieffenbachia Leaf Spot. Spots are formed on
secondary decay organisms. Small, dark greasy all parts of leaf blade except midrib, but not on
spots appear on leaflets and elongated black petioles and stems.
spots on stems and petioles. Bacteria are carried Xanthomonas glycines (phaseoli var. sojense)
on seed. (see Xanthomonas campestris pv. glycines). Bac-
Control Rotate crops; destroy diseased vines. terial Pustule of soybean, similar to regular bean
Spraying or dusting with copper may reduce blight but chiefly a foliage disease, present in
infection. These may be combined with most soybean areas, more severe in the South
streptomycin. Xanthomonas gummisudans (see
Xanthomonas campestris pv. vignicola (for- Xanthomonas campestris pv. gummisudans).
merly Xanthomonas vignicola). Cowpea Canker Bacterial Blight of Gladiolus.
on cowpeas and red kidney beans, a destructive Xanthomonas hederae (see Xanthomonas
disease, first described in 1944. Beans are campestris pv. hederae). Bacterial Leaf Spot of
blighted; cowpea stems have swollen, cankerlike English ivy.
lesions, with the cortex cracked open and a white Xanthomonas hyacinthi (see Xanthomonas
bacterial exudate. The plants tend to break over. campestris pv. hyacinthi). Hyacinth Yellows, yel-
Leaves, stems, pods, and seeds are liable to infec- low rot of Dutch hyacinth, occasionally entering
tion. Chinese Red cowpeas seem particularly sus- the country in imported bulbs.
ceptible, but the disease appears on other Xanthomonas incanae (see Xanthomonas
varieties. campestris pv. incanae). Bacterial Blight of gar-
Xanthomonas campestris pv. vitians (for- den stocks causing, since 1933, serious losses on
merly Xanthomonas vitians). Bacterial Wilt and flower-seed ranches in California; also present in
Leaf Spot of lettuce, South Carolina Lettuce home gardens.
Disease, wilting and rotting of lettuce leaves Xanthomonas juglandis (see Xanthomonas
and stems. In early stages plants are lighter campestris pv. juglandis). Walnut Blight on
green than normal. Leaves may have definite English or Persian walnut, black walnut, butter-
brown spots coalescing to large areas or may nut, Siebold walnut.
wilt following stem infection. Use windbreaks Xanthomonas oryzae (see Xanthomonas
to prevent injuries affording entrance to bacteria; campestris pv. oryzae). Carnation Pimple
also causes leaf spot of pepper and tomato. reported from Colorado as caused by a new
Xanthomonas campestris pv. zinniae Leaf and form of the rice blight organism.
Flower Spot of zinnia. Xanthomonas papavericola (see Xanthomonas
Xanthomonas campestris pv. zinniae (for- campestris pv. papavericola). Bacterial Blight of
merly Xanthomonas nigromaculans). Leaf Spot poppy on corn poppy and on Oriental, opium, and
on zinnia. California poppies.
88 Bacterial Diseases

Xanthomonas pelargonii (see Xanthomonas and other ornamentals that are symptomless car-
campestris pv. pelargonii). Bacterial Leaf Spot riers. There is no adequate control; roguing of
of geranium (Pelargonium). diseased vines and spraying for leafhoppers has
Xanthomonas pelargonii (see Xanthomonas proved ineffective. Propagate by cuttings from
campestris pv. pelargonii). Geranium Leaf Spot disease-free vineyards.
on Pelargonium spp.
Xanthomonas phaseoli (see Xanthomonas
campestris pv. phaseoli). Bacterial Bean Blight, Mycoplasmataceae
general and serious on beans but rare in some
western states. Phytoplasma
Xanthomonas pruni (see Xanthomonas
campestris pv. pruni. Bacterial Spot of stone Ash Yellows and Witches’ Broom On ash in
fruit, also called canker, shot hole, black spot; Michigan, Montana, Nebraska, North Carolina,
general on plum, Japanese plum prune, peach, and South Dakota and peanut in Oklahoma.
and nectarine east of the Rocky Mountains. Aster Yellows Throughout the United States,
Xanthomonas vesicatoria (see Xanthomonas also called Lettuce Rio Grande Disease, Lettuce
campestris pv. vesicatoria. Bacterial Spot of White Heart, Potato Purple Top.
tomato and pepper, common in wet seasons. Bean Phyllody Perhaps caused by a strain of
Xanthomonas vesicatoria var. raphani (see aster-yellows MLO.
Xanthomonas campestris pv. raphani). Leaf California Aster Yellows In the West, also
Spot of radish, turnip, and other crucifers, similar known as Celery Yellows, Western.
to bacterial spot on tomato. Aster Yellows, Potato Late Break, Strawberry
Xanthomonas vignicola (see Xanthomonas Green Petal. Aster yellows may appear in more
campestris pv. vignicola). Cowpea Canker on than 170 species of 38 families of dicotyledons.
cowpeas and red kidney beans. It is serious on China aster, may also affect
Xanthomonas vitians (see Xanthomonas anemone, calendula, coreopsis, cosmos, purple
campestris pv. vitians). Bacterial Wilt and Leaf coneflower (Echinacea), delphinium, daisies,
Spot of lettuce, South Carolina Lettuce Disease, golden-glow, hydrangea, marigold, petunia,
wilting and rotting of lettuce leaves and stems. phlox, scabiosa, strawflower, and other flowers.
Xanthomonas nigromaculans (see It is serious on lettuce, alfalfa, endive, carrot,
Xanthomonas campestris pv. zinniae). Leaf Spot parsley, New Zealand spinach, radish, and some
on zinnia. other vegetables, but not on peas, beans, or other
Xylella fastidiosa Bacterial Leaf Scorch on legumes. This disease is now known to be caused
maple, pecan, mulberry, northern red oak and by a phytoplasma organism.
sweet gum. In most plants vein clearing is followed by
Pierce’s Grape Disease First described as Cal- chlorosis of newly formed tissues, adventitious
ifornia vine disease by Pierce in 1892, now growth, erect habit, virescence of flowers. Asters
known as cause of grape degeneration in Gulf have a stiff yellow growth with many secondary
states; reported from Rhode Island. First symp- shoots; are stunted, with short internodes; flowers
toms are scalding and browning of leaf tissues, are greenish, dwarfed, or none. The chief vector
often with veins remaining green; canes die back is the six-spotted leafhopper (Macrosteles
from tips in late summer; growth is dwarfed, fruit fascifrons). The virus multiplies in the insect,
shriveled; roots die. The bacterium invades the and there is a delay of 10 days or more after the
xylem and turns it brown. Alfalfa plants are insect feeds on a diseased plant before it can
stunted with short stems and small leaves. Many infect a healthy specimen. There is no transmis-
species of sharpshooter leafhoppers transmit the sion through insect eggs or aster seeds.
bacterium to grape from alfalfa, clovers, grasses, Celery petioles are upright, somewhat elon-
also from ivy, acacia, fuchsia, rosemary, zinnia, gated, with inner petioles short, chlorotic,
Mycoplasmataceae 89

twisted, brittle, often cracked, yellow. The celery about May 1 from eggs wintered on elm bark and
strain of the virus causes yellowing and stunting feed on leaf veins. Adults move from diseased to
of cucumber, squash, pumpkin; infects gladiolus healthy trees.
and zinnia. There is hope of propagating elms resistant to
Control of aster yellows is directed against the phloem necrosis. Communities should interplant
leafhoppers. Asters are grown commercially existing elms with Asiatic or European varieties
under frames of cheesecloth, 22 threads to the or with some other type of tree to provide shade if
inch, or wire screening, 18 threads to the inch. and when present elms die.
In home gardens all diseased plants should be Peach Western X-Disease Perhaps same as
rogued immediately and overwintering weeds, X-disease but usually treated separately; also
which harbor leafhopper eggs, destroyed. known as cherry buckskin and western-X little
Spraying or dusting ornamentals and vegetables cherry. The pathogen is transmitted by leafhop-
with pyrethrum will reduce the number of vectors pers (Colladonus germinatus, Fieberella florii,
but will not entirely eliminate the disease. Osbornellus borealis, and others) to peach, nec-
Recent work raises the probability that the tarine and cherry in western states. Symptoms
etiological agent of aster yellows is vary according to rootstock, but cherry fruit is
a mycoplasma rather than a virus. Therefore, smaller than normal. Sour cherries are puttylike,
treatment with antibiotics, such as chlortetracy- pinkish; sweet cherries are small, conical, hang
cline, has suppressed the development of yellows on trees late, fail to develop normal color. Symp-
symptoms. Mycoplasma-like bodies have been toms on peach are similar to those of X-disease.
seen in microscopic study of diseased plants and Peach X-Disease On peach and chokecherry,
in transmitting leafhopper vectors, but not in sometimes cherry in the northern United States
healthy plants or nontransmitting vectors. and of major importance in Connecticut, Massa-
Clover Proliferation On strawberry and onion. chusetts, and New York. Peach trees appear nor-
Corn Stunt A dwarfing disease present primar- mal in spring for 6 or 7 weeks after growth starts,
ily in the South; transmitted by leafhoppers. then foliage shows a diffused yellow and red
Mycoplasma-like bodies present; See discoloration with a longitudinal upward curling
Spiroplasma citri. of leaf edges; spots may drop out, leaving
Elm Phloem Necrosis On American elm from a tattered effect. Defoliation starts by mid-sum-
West Virginia and Georgia to northern Missis- mer. Fruits shrivel and drop or ripen prematurely.
sippi, eastern Oklahoma, Kansas, and Nebraska. Seed do not develop. Weakened trees are killed
Origin unknown but apparently present since by low temperatures or remain unproductive.
1882; the disease reached epidemic proportions Chokecherry has conspicuous premature red-
in Ohio in 1944, killing 20,000 trees that year dening of foliage, dead embryos in fruit. The
near Dayton and 10,000 at Columbus. The most second and third seasons after infection foliage
reliable diagnostic character is a buttercup yellow colors are duller, there are rosettes of small leaves
discoloration of the phloem, often flecked with on terminals, and death may follow. Natural
brown or black and an odor of wintergreen. infection is apparently from chokecherry to
Destruction of phloem causes the bark to loosen peach (not peach to peach or peach to choke-
and fall away. Roots die first, then the phloem in cherry) by a leafhopper (Colladonus clitellarius).
lower portions of tree, followed by wilting and Elimination of chokecherries within 500 feet of
defoliation. American elms may be attacked at peach trees provides the best control.
any age; they wilt and die suddenly within 3 or Peach Yellow Leaf Roll; a form of Western
4 weeks or gradually decline for 12 to 18 months. X-Disease; perhaps caused by a more severe
This is now thought to be caused by strain of the MLO.
a mycoplasma-like agent. Transmission is by Peach Yellows; Little Peach. First noted near
the white-banded elm leafhopper (Scaphoideus Philadelphia in 1791 and so serious that in 1796
luteolus) and possibly other species. Nymphs hatch the American Philosophical Society offered
90 Bacterial Diseases

a $60 prize for the best method of preventing pre- Strawberry Green Petal Perhaps due to a strain
mature decay of peach trees. Present in eastern of aster yellows MLO, as is chlorotic phyllody
states on peach, almond, nectarine, apricot and reported from Louisiana. Flowers have enlarged
plum. Not found west of the Mississippi or in the sepals, small green petals.
South. In peach, clearing of veins, production of Bud Proliferation and Delayed Maturity, on
thin erect shoots with small chlorotic leaves, pre- soybean.
mature ripening of fruit (with reddish streaks in Decline of ash.
flesh and insipid taste) is followed by death of the Lethal Yellowing on palms.
tree in a year or so. The little peach strain of the Phloem Necrosis of chrysanthemum.
MLO causes distortion of young leaves at tips of Spiroplasma citri Corn Stunt. Has been
branches, small fruit, delayed ripening. Plum is reported on corn, onions, horseradish, shepherd’s
systemically infected, with few obvious symptoms. purse, yellow rocket, and wild mustard.
Transmission is by the plum leafhopper or budding. Stunt of blueberry.
Control Budsticks and dormant nursery trees Virescence on horseradish.
can be safely treated with heat sufficient to kill Witches’ Broom on pigeon pea (Cajanus
the MLO (122  F for 5 to 10 min), but cured trees cajan), and black raspberry.
are susceptible to reinfection. Most effective con- Witches’ Broom on Japanese persimmon,
trol is removal of wild plum trees around peach and lilac.
orchard and spraying to control leafhoppers. Witches’ Broom and Yellowing on annual
Potato Apical Leaf Roll and Arizona Purple statice.
Top Wilt Caused by aster yellows. Yellows of elm.
Black Knot

The term black knot is used to designate a disease with an olive green, velvety layer made up of
with black knotty excrescences. brownish conidiophores and one-celled hyaline
conidia of the anamorph Hormodendron state.
Conidia are spread by wind.
Apiosporina In late summer black stromata cover the
affected tissues, and the galls become hard. Asci
Ascomycetes, Pleosporales are formed during the winter in cavities in the
stroma; ascospores are discharged and germinate
Asci are in locules, without well-marked perithe- in early spring, completing the 2-year cycle.
cial walls, immersed in a massive, carbonaceous Knots are produced from primary infection by
stroma, erumpent and superficial at maturity. ascospores or from secondary infection from
Spores are hyaline, unequally two-celled. mycelium formed in old knots and growing out
Apiosporina morbosa (formerly Dibotryon to invade new tissue. Limbs may be girdled and
morbosum). Black Knot of plum and cherry, killed; trees are stunted and dwarfed, nearly
Prunus Black Knot, Plum Wart, widespread and worthless after a few years. Old knots may be
serious on garden plums, also present on sweet riddled with insects or covered with a pink fungus
and sour cherries, chokecherry, and apricot. growing on the Apiosporina mycelium.
Apparently a native disease, destructive in Mas- Control Cut out infected twigs and branches, 3
sachusetts by 1811 and the pathogen described or 4 inches beyond the knot, to include advancing
from Pennsylvania in 1821, black knot has been perennial mycelium. Do this in winter or before
reported on peach, long thought to be immune. April 1. Eradicate or thoroughly clean up wild
The chief symptoms are black, rough, cylin- plums and cherries in the vicinity. Spray at
drical or spindle-shaped enlargements of twigs delayed dormant stage in spring (just as buds
into knots two to four times their thickness and break) with bordeaux mixture or with liquid
several inches long (see Fig. 1). Infection takes lime sulfur. The latter is preferable unless oil
place in spring, but swelling is not evident until is combined in the spray as an insecticide.
growth starts the following spring, at which time Spray with lime sulfur at full bloom. Dibotryon
the bark ruptures, and a light yellowish growth morbosum (see Apiosporina morbosa). Black
fills the crevices. In late spring this is covered Knot of plum and cherry, Prunus Black Knot,

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_16, 91

# Springer Science+Business Media Dordrecht 2013
92 Black Knot

Plum Wart, widespread and serious on garden

plums, also present on sweet and sour cherries,
chokecherry, and apricot.


Ascomycetes, Dothideales

Perithecia in clusters on wood; spores dark, with

several cells.
Gibberidea heliopsidis (see Leptosphaeria
heliopsidis). Black Knot, Black Patch on
goldenrod and sunflower.
Leptosphaeria heliopsidis (formerly
Gibberidea heliopsidis). Black Knot, Black
Patch on goldenrod and sunflower.

Fig. 1 Black Knot on Prunus sp.


The term blackleg is used to describe darkening reported in France in 1849, and in the United
at the base of a stem or plant. Blackleg of potatoes States in 1910. It is generally distributed east of
and delphinium are described under Bacterial the Rocky Mountains and formerly caused from
Diseases; blackleg of geraniums is under Rots. 50 to 90 % loss. With improved seed and seed
treatment it has become less important.
The first symptom is a sunken area in the stem
Cylindrocarpon near the ground, which extends until the stem is
girdled and the area turns black. Leaves, seed
▶ Rots. stalks, and seed pods have circular, light brown
Cylindrocarpon obtusisporum Blackleg; on spots. Small black pycnidia appearing on the
grape. lesions distinguish blackleg from other cabbage
diseases. The leaves sometimes turn purple and
wilt, but there is no defoliation, as in black rot.
Phoma The fungus reaches the soil via infected plant
debris, remaining alive 2 or more years. Spores
Deuteromycetes, Coelomycetes are spread by splashing rain, or manure, on tools,
and perhaps by insects, with new lesions resulting
Pycnidia dark, ostiolate, lenticular to globose, in 10 to 14 days. But the chief spread is by
immersed in host tissue, erumpent or with short mycelium wintering in infected seed. When
beak piercing the epidermis; conidiophores short such seed is planted, fruiting bodies are formed
or obsolete, conidia small, one-celled, hyaline, on cotyledons as they are pushed above ground,
ovate to elongate; parasitic on seed plants, chiefly and these serve as a source of inoculum for
on stems and fruits, rarely on leaves. nearby plants. A few diseased seed can start an
Phoma lingam Blackleg of crucifers, Foot Rot, epiphytotic in wet weather.
Phoma Wilt of plants of the mustard family, Control Use seed grown on the Pacific Coast,
including cabbage, cauliflower, Chinese cabbage, which is usually, although not always, disease-
brussels sprouts, charlock, garden cress, pepper free. If the seed is infected, tie loosely in cheese-
grass, kale, kohlrabi, mustard, rape, radish, ruta- cloth bags and immerse in hot water, held at
baga, turnip, stock, and sweet alyssum. The 122  F for 30 min. It is sometimes possible to
teleomorph state, Lystosphaeria maculans has buy seed already treated. Sterilize soil for the
been found on cabbage. The fungus was first seedbed; use a 3-year rotation; do not splash
noticed in Germany in 1791; the disease was seedlings when watering; do not transplant any

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_17, 93

# Springer Science+Business Media Dordrecht 2013
94 Blackleg

seedlings if the disease shows up in the seedbed;

do not feed cabbage refuse to cattle; do not trans- Leptosphaeria
fer cultivators and other tools from a diseased to
a healthy field without using a disinfestant. Leptosphaeria maculans Blackleg on canola.
Black Mildew

The terms black mildew, sooty mold, and black Asterina

spot have been used to some extent inter-
changeably. In this text the term sooty mold Ascomycetes, Asterinales
is restricted to those fungi living on insect
exudate and hence not true parasites. Included Asterina species are parasites on the surface of
here under Black Mildew are parasitic fungi leaves and are usually found in warm climates. In
that have a superficial dark mycelium. They some cases the disease is called black mildew, in
are members of the Erysiphales (Meliolales others, black spot. The perithecia are dimidiate,
according to some classifications) and hence having the top half covered with a shield, a small,
similar to powdery mildews except for the dark round stroma composed of radially arranged dark
color, or they belong to the Hemisphaeriales, hyphae. Underneath this stromatic cover, called
characterized by a dark stroma simulating the scutellum, there is a single layer of fruiting cells;
upper portion of a perithecium. In a few cases paraphyses are lacking; spores are dark, two-
the diseases are called black spot rather than celled. The mycelium, which is free over the
mildew. surface, has lobed appendages, hyphopodia,
which act as haustoria in penetrating the cuticle
and obtaining nourishment from the host.
Apiosporina Asteridium lepidigenum (formerly Asterina
lepidigena). Black Mildew on lyonia, Florida.
Ascomycetes, Pleosporales Asterina anomala (see Limacinula anomala).
Black Mildew on California-laurel, California.
Perithecia and mycelium superficial; mycelium Asterina delitescens Black Spot on redbay.
with setae and perithecia usually hairy; Asterina diplopoides Black Spot on leucothoë.
paraphysoids present; spores two-celled; dark. Asterina gaultheriae (see Schizothyrium
Apiosporina collinsii Witches’ Broom of ser- pomi). Black Mildew on bearberry, Wisconsin.
viceberry (Amelanchier) widespread. Perennial Asterina lepidigena (see Asteridium
mycelium stimulates the development of numer- lepidigenum). Black Mildew on lyonia, Florida.
ous stout branches into a broom. A sooty growth Asterina orbicularis Black Spot on American
on underside of leaves is first olive brown, then holly and Ilex spp.
black. Numerous globose, beadlike, black peri- Limacinula anomala (formerly Asterina
thecia appear in late summer. The damage to the anomala). Black Mildew on California-laurel,
host is not serious. California.

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# Springer Science+Business Media Dordrecht 2013
96 Black Mildew

Schizothyrium pomi (formerly Asterina Appendiculella calostroma (formerly Irene

gaultheriae). Black Mildew on bearberry, calostroma). Black Mildew on wax-myrtle, Gulf
Wisconsin. States.
Appendiculella perseae (formerly Irene
perseae). Black Mildew on avocado, Florida.
Asterinella Like Irene except that perithecia have no
Ascomycetes, Asterinales Asteridiella manca (formerly Irenina
Like Asterina but lacking hyphopodia; with or manca). Black Mildew on wax-myrtle,
without paraphyses; spores dark, two-celled. Mississippi.
Asterinella puiggarii Black Spot on eugenia. Irene araliae (see Appendiculella araliae).
Black Mildew on magnolia, Mississippi.
Irene calostroma (see Appendiculella
Dimerosporium calostroma). Black Mildew on wax-myrtle,
Gulf States.
According to some authorities this is the same as Irene perseae (see Appendiculella perseae).
Asterina but the name Dimerosporium is in com- Black Mildew on avocado, Florida.
mon use. Irenina manca (see Asteridiella manca).
Dimerosporium abietis (see Rasutoria Black Mildew on wax-myrtle, Mississippi.
abietiis). Black Mildew on Pacific silver and low-
land white firs.
Dimerosporium hispidulum Black Mildew Lembosia (Morenoella)
on boxelder.
Dimerosporium pulchrum Black Mildew on Ascomycetes, Dothideales
Dimerosporium robiniae Black Mildew on Brown vegetative mycelium with hyphopodia on
ailanthus. surface of host; linear stroma, scutellum, over
Dimerosporium tropicale Black Mildew on single layer of fruiting cells; paraphyses present;
bignonia, Mississippi. spores dark, two-celled.
Rasutoria abietiis formerly Dimerosporium Echidnodella angustiformis (formerly
abietis). Black Mildew on Pacific silver and Morenoella angustiformis). Black Mildew on
lowland white firs. Black patches are formed on holly (Ilex spp.), Mississippi.
older needles, usually on under surface. There is Echidnodella rugispora (formerly Lembosia
no apparent injury to trees. rugispora). Black Mildew on redbay, swampbay,
Mississippi, North Carolina.
Lembosia cactorum Black Mildew on cactus,
(Irene) Asteridiella Florida.
Lembosia coccolobae Black Mildew on
Ascomycetes, Meliolales sea-grape, Florida; also L. portoricensis and
L. tenella.
Mycelium with capitate hyphopodia but no Lembosia illiciicola Black Mildew on
bristles; perithecia with larviform appendages; anise-trees, Alabama, Mississippi.
spores dark, with several cells. Lembosia illiciicola (see Echidnodella
Appendiculella araliae (formerly Irene rugispora). Black Mildew on redbay, swampbay,
araliae). Black Mildew on magnolia, Mississippi. Mississippi, North Carolina.
Sthughesia 97

Morenoella angustiformis (see Echidrodella Meliola cookeana Black Mildew on

angustiformis). Black Mildew on holly (Ilex spp.), callicarpa, lantana.
Mississippi. Meliola cryptocarpa (see Irenopsis
Schiffnerula pulchra On dogwood. cryptocarpa). Black Mildew on gordonia.
Meliola lippiae Black Mildew on lippia.
Meliola magnoliae Black Mildew on
Meliola magnolia.
Like Irene except that mycelium has setae (stiff
Ascomycetes, Erysiphales (or bristles) and perithecia lack larviform appendages.
Meliolales), Meliolaceae Meliola martiana (formerly Irenopsis
martiniana). Black Mildew on redbay,
Most abundant in tropics. Superficial dark swampbay, Alabama, Florida, Mississippi.
mycelium with hyphopodia and setae; perithecia Meliola nidulans Black Mildew on blueberry,
globose, coal black without ostiole or append- wintergreen.
ages but often with setae; spores several-celled, Meliola palmicola Black Mildew on palmetto.
dark; paraphyses lacking. Conidia are lacking Meliola tenuis Black Mildew on bamboo.
in most species, of Helminthosporium type in Meliola wrightii (see Diplotheca tunaei).
others. Black Mildew on chinaberry.
Diplotheca tunae (formerly Meliola
wrightii). Black Mildew on chinaberry.
Irenopsis cryptocarpa formerly Meliola Sthughesia
cryptocarpa). Black Mildew on gordonia.
Irenopsis martiniana (see Meliola Ascomycetes, Dothideales
martiana). Black Mildew on redbay, swampbay,
Alabama, Florida, Mississippi. Perithecia smooth; spores two-celled, dark;
Meliola amphitricha Black Mildew on paraphyses lacking.
boxelder, magnolia, redbay, swampbay. Dimerium juniperi (see Sthughesia juniperi).
Meliola bidentata Black Mildew on bignonia. Black Mildew on Rocky Mountain juniper,
Meliola camelliae Black Mildew of camellia. California.
Abundant black growth may cover camellia Sthughesia juniperi (formerly Dimerium
leaves and twigs. Spraying with a light summer juniperi). Black Mildew on Rocky Mountain
oil is sometimes effective. juniper, California.

In common usage the term black spot without transferred to Marssonina. The blackspot fungus
qualifying adjectives has come to mean but one was first reported in the United States in 1831,
disease, rose black spot, with the two words cur- from Philadelphia, and in 1912 Wolf made the
rently written as one, blackspot. This section is connection with the teleomorph state, so that the
limited to the rose disease. Delphinium black spot correct name became Diplocarpon rosae.
will be found under Bacterial Diseases, elm black Blackspot is probably the most widely distrib-
spot under Leaf Spots, other black spots under uted and best known rose disease. It is confined to
Black Mildew. roses, garden and greenhouse, and may affect prac-
tically all varieties, although not all are equally
susceptible. There has been some progress made
Diplocarpon in breeding resistant varieties, but recent investiga-
tion disclosing many physiological races of the
Ascomycetes, Helotiales, fungus explains why roses that are almost immune
Dermateaceae (Mollisiaceae) to blackspot in one location may succumb in
another. Rosa bracteata is the only species thus
Apothecia innate, formed in dead leaves, but far shown to be reasonably resistant to all the
at maturity rupturing overlying tissues; horny to different races tested. Roses with the Pernetiana
leathery with a thick margin or outer wall parentage, which has given us the lovely yellows,
(excipulum) of dark, thick-walled cells; spores coppers, and blends, are especially prone to
two-celled, hyaline; paraphyses present. Anamorph blackspot. Some roses, like Radiance, are tolerant
state is a Marssonina with two-celled hyaline spores of blackspot, usually holding their leaves, even
in an acervulus. though they cannot be considered resistant.
Diplocarpon rosae Rose Blackspot, general Symptoms are primarily more or less circular
on rose but less serious in the semi-arid black spots, up to 1/2 inch in diameter, with
Southwest; reported from all states except radiating fimbriate or fringed margins (see
Arizona, Nevada, and Wyoming. Fig. 1). This fimbriate margin is a special diag-
For nearly 100 years the fungus was known nostic character, differentiating blackspot from
only by its anamorph state, which has had about other leaf spots and from discolorations due to
25 different names. The first definite record is by cold or chemicals. The spots vary from one or two
Fries in Sweden in 1815, under the name to a dozen or more on a leaf, usually on the upper
Erysiphe radiosum, but the first valid description surface. With close examination you can see
was by Libert in 1827 as Asteroma rosae. Later small black dots or pimples in the center of the
Fries called it Actinonema rosae, and that term spots. These are the acervuli, bearing conidia, and
was widely used until Actinonema species were they glisten when wet (see Fig. 2).

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100 Blackspot

In susceptible varieties the appearance of black areas, slightly blistered, without fimbriate
black spots is soon followed by yellowing of margins.
a portion or all of leaflets and then by defoliation. Infection occurs through either leaf surface,
The leaf fall is apparently correlated with the fungus sending its germ tube directly through
increased production of ethylene gas in diseased the cuticle by mechanical pressure. The hyphae
tissue and perhaps by a difference in auxin gradi- form a network under the cuticle, joining together
ent between leaf and stem. Some roses lose into several parallel filaments radiating from the
almost all their leaves, put out another set and point of infection. The hyphae are actually color-
lose those, and often are trying to leaf out for the less, the black color of the spot coming from the
third time by late summer. The process is so death and disorganization of host cells. The
devitalizing that some bushes may die during mycelial growth is between cells, with haustoria
the following winter. On tolerant varieties leaf (suckers) invading epidermal and palisade cells
spots are present, though usually in smaller for nourishment.
numbers, but there is much less yellowing and Acervuli, summer fruiting bodies, formed just
defoliation. Cane lesions are small indistinct under the cuticle, bear two-celled hyaline conidia
on short conidiophores on a thin, basal stroma.
Splashed by rain or overhead watering, or spread
by gardeners working among wet plants, the
conidia germinate and enter a leaf if there is
continued moisture for at least 6 h. Rain, heavy
dew, fog, and sprinklers used late in the day so
foliage does not dry off before night provide the
requisite moisture. New spots show up within
a week and new spores within 10 days. Secondary
cycles are repeated all summer – from late May to
late October around New York City.
In my personal experience, the spread of disease
is most rapid where large numbers of susceptible
varieties are massed together. If all the yellows, for
instance, are planted together, the disease gets such
a head start, and builds up so much inoculum to
spread to the more tolerant red and pink varieties
nearby, that these varieties also are more heavily
infected than usual. When roses are mixed in beds
so that one or two particularly susceptible bushes
are surrounded by more resistant types, the infec-
Fig. 1 Rose Blackspot. Note fimbriate margin to spot tive material cannot increase so rapidly, and the net

Fig. 2 Rose Blackspot.

Two-celled conidia formed
in Acervulus under cuticle
Diplocarpon 101

result is less disease in the garden as a whole. one bush to another during the season. Drastic
Protected corners in the garden where air circula- spring pruning, far lower than normal, reduces the
tion is poor also increase the disease potentiality. amount of inoculum from infected canes.
Spores are apt to be splashed farther when water The importance of a dormant spray is debat-
hits hard-packed soil without a mulch. able. Experiments have shown that as a true
When old leaves drop to the ground, the myce- eradicant, applied in winter, it has little value in
lium continues a saprophytic existence, growing reducing the amount of blackspot the next sum-
through dead tissue with hyphae that are now mer. Use liquid lime sulfur after pruning, pro-
dark in color. In spring three types of fruiting vided the buds have not broken far enough to
bodies may be formed: microacervuli or sperma- show the leaflets.
gonia containing very small cells that perhaps act Summer spraying or dusting, weekly through-
as male cells; apothecia, the sexual fruiting bod- out the season (from late April to early November
ies formed on a stroma between the epidermis in New Jersey) is essential if you want to keep
and palisade cells and covered with a circular enough foliage on bushes for continuous produc-
shield of radiating strands; and winter acervuli, tion of fine flowers (it takes food manufactured in
formed internally and producing new conidia several leaves to produce one bloom) and for
in spring. The Diplocarpon or apothecial stage winter survival. Some strong varieties will, how-
is apparently not essential; it is known only ever, live for years without chemical treatment;
in northeastern United States and south-central they are usually scraggly bushes with erratic
Canada. The shield over the apothecium ruptures, bloom. The idea that floribunda varieties do not
and the two-celled ascospores are forcibly require as much spraying as hybrid teas is
discharged into the air to infect lowest leaves. a misconception. Some floribundas are quite
Where the sexual stage is not formed, primary resistant; others are very susceptible. The same
spring infection comes from conidia splashed by holds true for old-fashioned shrub roses. All too
rain to foliage overhead, from acervuli either in often blackspot gets a head start in a garden from
overwintered leaves on the ground or in cane shrub roses we thought it unnecessary to spray.
lesions. New roses from a nursery sometimes Roses can be defoliated as readily by
bring blackspot via these cane lesions to chemicals as by the blackspot fungus; so the
a garden previously free of disease. fungicide chosen must be safe under the condi-
Control The importance of sanitation may have tions of applications as well as effective. There
been somewhat overstressed; it cannot replace rou- are many chemicals that will control blackspot if
tine spraying or dusting. It is certainly a good idea they are applied regularly and thoroughly. Choice
to pick off for burning the first spotted leaves, if this depends somewhat on climate. Some copper
is done when bushes are dry so that the act of sprays and dusts cause red spotting and defolia-
removal does not further spread the fungus. Raking tion in cool, cloudy weather. Bordeaux mixture is
up old leaves from the ground at the end of the both unsightly and harmful, unless used in very
season makes the garden neater and may reduce weak dilution. At strengths recommended for
the amount of inoculum in spring, but, because the vegetables it will quickly turn rose leaves yellow
fungus winters also on canes in most sections of and make them drop off. Dusts containing more
the country, removal of leaves cannot be expected than 3 to 4 % metallic copper are injurious under
to provide a disease-free garden the next season. some weather conditions. Dusting sulfur fine
Comparative tests have shown that fall cleanup enough to pass through a 325-mesh screen has
is ineffectual. A good mulch, applied after been successfully used for years for blackspot
uncovering and the first feeding in spring, serves control, but in hot weather it burns margins of
as a mechanical barrier between inoculum from leaves. Copper and sulfur have a synergistic
overwintered leaves on the ground and developing effect; a mixture of the two is more effective
leaves overhead. A mulch also reduces disease by than either used alone, but such a mixture also
reducing the distance spores can be splashed from combines injurious effects.
102 Blackspot

There are literally hundreds of combination surfaces, and applications must be repeated at
rose sprays and dusts on the market under brand approximately weekly intervals. This may mean
names, and it seems to me easier, and even every 5 or 6 days when plants are growing rapidly
cheaper, considering the time saved, for home in a rainy spring and perhaps every 7 to 9 days in
gardeners to make use of them to control dry weather, when growth is slow. Intervals of
blackspot and other rose diseases as well as 10 to 14 days between sprays seldom give
insects in one operation. You will have to deter- adequate control. Most directions call for appli-
mine by trial and error the best combination for cation ahead of rain so that the foliage will be
your area, and you may not find one that com- protected when spores germinate during the rain;
bines remedies for all the pests you may have to but if sprays are applied every 7 days, there will
fight through the season. Choose one that con- always be enough residue left on the foliage to
tains ingredients required every week all sum- give protection during the next rain. It is not
mer, and then add other chemicals if and when necessary to make an additional application
necessary. Whatever mixture is chosen, coverage immediately after a rain if your spraying is on
should be complete on upper and lower leaf a regular basis.

According to Webster, blight is “any disease or hayfever. There are also parasitic forms causing
injury of plants resulting in withering, cessation blights and leaf spots. Sometimes the disease
of growth and death of parts, as in leaves, without starts as a leaf spot, but the lesions, typically
rotting.” The term is somewhat loosely used by formed in concentric circles, run together to
pathologists and gardeners to cover a wide vari- form a blight, the dark conidia making the surface
ety of diseases, some of which may have rotting appear dark and velvety.
as a secondary symptom. In general, the chief Alternaria alternata Blight, Foliage and Pod
characteristic of a blight is sudden and conspicu- of pea.
ous leaf and fruit damage, in contradistinction to Alternaria cassiae Seedling Blight of Cassia
leaf spotting, where dead areas are definitely (sicklepod, and coffee senna), and showy
delimited, or to wilt due to a toxin or other dis- crotalaria.
turbance in the vascular system. Fire blight, Alternaria cucumerina Alternaria Blight of
discussed under Bacterial Diseases, is a typical Cucurbits, Cucumber Blight, Black Mold, gen-
blight, with twigs and branches dying back but eral on cucumbers, muskmelon, watermelon, and
holding withered, dead foliage. winter and summer squash. Symptoms appear in
the middle of the season, first nearest the center of
the hill. Circular brown spots with concentric
Alternaria rings are visible only on upper surface of leaves,
but a black, moldy growth, made up of conidio-
Deuteromycetes, Hyphomycetes phores and large brown spores, can be seen on
both leaf surfaces. Leaves curl and dry up, canta-
Dark, muriform conidia formed in chains, simple loupe foliage being more sensitive than that of
or branches, or sometimes singly, on dark, simple other cucurbits. The disease spreads rapidly in
conidiophores growing from dark hyphae (see warm, humid weather, and, with the vines drying,
Fig. 1). The apical portion of each conidium is the fruit is exposed to sunburn. Sunken spots
narrowed and often elongated, bearing at its tip develop on the fruit, covered with an olive
the next ovoid, tapering conidium. Species with green mass of conidia. Other species of
this characteristic formerly placed in Alternaria cause a decay of melons in transit
Macrosporium are now in Alternaria; those with and storage.
spores rounded at both ends have been transferred Control Purdue 44 and some other varieties of
to Stemphylium. muskmelon are rather resistant.
There are many saprophytic species in Alternaria dauci Alternaria Blight of carrot,
Alternaria, the spores of which are wind-borne Carrot Leaf Blight, general on carrot and parsley.
for many miles and are a common cause of Affected leaves and petioles are spotted, then turn

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_20, 103

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104 Blights

Alternaria Botrytis Cercospora Entomosporium Ovulinia

Pestalotia Phomopsis Septoria Volutella

Fig. 1 Conidial Production Among Some Fungi Causing cell, borne free on mycelium; Pestalotia, in acervulus,
Blights. Alternaria, dark muriform spores in chains; median cells colored, end cells hyaline, apical cell with
Botrytis, hyaline spores in clusters; Cercospora, pale to appendages; Phomopsis, oval and filiform hyaline spores
dark septate spores on dark conidia protruding from sto- in pycnidium; Septoria, septate hyaline spores in Pycnid-
mata; Entomosporium, peculiarly appendaged spores in ium; Volutella, hyaline spores formed on a hairy
acervulus; Ovulina, hyaline spore with basal disjunctor sporodochium

yellow and brown; entire tops are killed in severe Control Commercial growers can often avoid
infections. In California the disease is known as Alternaria blight by keeping plants growing con-
late blight, with the peak coming in November. tinuously in the greenhouse. Cuttings should be
The fungus apparently winters in discarded tops disease-free, taken from midway up the stem,
and on seed. broken at the joint rather than cut, and started in
Control Clean up refuse. Spray with a fixed cop- sterilized soil. Ordinarily the foliage should be
per spray or dust, starting soon after seedlings kept dry, but under mist propagation chemicals
emerge and repeating at 7-to 10-day intervals. introduced into the mist system have reduced
Alternaria dianthicola Carnation Collar blight.
Blight, Leaf Spot, Stem and Branch Rot, general Alternaria helianthi Blight and Stem Lesion of
on carnation, widespread on garden pinks and sunflower.
sweet william. The chief symptom is a blight or Alternaria panax Alternaria Blight, Root Rot,
rot at leaf bases and around nodes, which are Leaf Spot of ginseng, ming aralia, and golden-
girdled. Spots on leaves are ashy white but cen- seal, generally distributed. In Ohio the disease
ters of old spots are covered with dark brown to appears each year in semiepidemic form and has
black fungus growth. Leaves may be constricted been controlled with bordeaux mixture or a fixed
and twisted, the tip killed. Branches die back to copper spray plus a wetting agent, starting when
the girdled area, and black crusts of spores are plants emerge in early May and repeating every
formed on the cankers. Conidia are spread during 2 weeks until 3 weeks after bloom.
watering in the greenhouse or in rains, outdoors. Alternaria solani Early Blight of potato and
Entrance is through wounds, stomata, or directly tomato, general on these hosts, occasional on
through the cuticle. The spores are carried on eggplant and pepper. The pathogen was first
cuttings. described from New Jersey, in 1882.
Ascochyta 105

Leaf symptoms are dark brown, circular to

oval spots, marked with concentric rings in Ascochyta
a target effect, appearing first on lower, shaded
foliage, with the spots growing together to blight Deuteromycetes, Coelomycetes
large portions or all of leaves, exposing fruits.
There may be a collar rot of young tomato seed- Pycnidia dark, globose, separate, immersed in
lings, sunken spots or cankers on older stems, host tissue, ostiolate; spores two-celled, hyaline
blossom-drop with loss of young fruits, or dark ovoid to oblong.
leathery spots near the stem end of older fruits. Ascochyta asparagina Stem Blight of asparagus
Alternaria blight is the most common leaf spot fern. Small branchlets dry and drop prematurely;
disease of tomatoes in the Central and Atlantic small branches are killed if attacked at crown.
States but is somewhat less important elsewhere. Ascochyta chrysanthemi (Mycosphaerella
Foliage symptoms on potato are similar to ligulicola). (see ▶Phoma chrysanthemi
those on tomato. Small round spots on tubers (Didymella ligulica)). Ascochyta Ray Blight of
afford entrance to secondary rot organisms. chrysanthemum, a conspicuous and rapid disease
Each leaf spot may produce three or four crops of ray flowers.
of dark spores, which remain viable more than Ascochyta fabae f. sp. spiricia Leaf Blight of
a year. They are blown by wind, splashed by rain, vetch.
sometimes transmitted by flea beetles. The fun- Ascochyta piniperda Spruce Twig Blight on
gus is a weak parasite, entering through wounds young shoots of red, Norway, and blue spruce;
and thriving in warm, moist weather, with 85  F apparently a minor disease.
as optimum temperature. It can survive in soil as Ascochyta pisi, A. pinodes, A.
long as the host refuse is not completely rotted; it pinodella Ascochyta Blight or Mycosphaerella
also winters on seed and on weed hosts. Blight of peas. All three fungi may be connected
Control Plan, if possible, a 3-year rotation with with the disease complex known as Ascochyta
crops not in the potato family; dig under diseased blight, are carried in infected seed and overwinter
refuse immediately after harvest. Use seed from in plant debris. A. pinodes has Mycosphaerella
healthy tomatoes, or purchase plants free from pinodes as its ascospore stage so that the life
collar rot. cycle can start from either pycnidia or perithecia
Alternaria tagetica Blight of marigold. produced on plants or stubble. Lesions begin as
Alternaria tenuissima Alternaria Blight, Leaf small purplish specks on leaves and pods. When
Spot of violet and pansy. Spots vary from greenish infection is caused by M. pinodes or A. pinodella,
yellow to light buff with burnt amber margins. the specks enlarge to round, targetlike spots,
Brown patches run together to form large, blighted which join together to form irregular, brownish
areas. Clean up and burn old leaves in fall. purple blotches. M. pinodes often withers and
Alternaria zinniae Zinnia Blight, Alternariosis distorts young pods; A. pinodella causes
on zinnia. Small reddish brown spots with grayish a severe foot rot, a dark region at the soil line.
white centers increase to irregular, large, brown, Elongated, purplish black stem lesions are com-
dry areas. Similar spots on stem internodes or at mon. A. pisi causes leaf spots with dark brown
nodes may girdle the stem, with dying back of margins, stem and pod spots, but no foot rot.
upper portions. Dark brown to black basal can- Control Use western-grown seed, usually free
kers with sunken lesions are common. Roots may from the disease; clean up all pea refuse and use
turn dark gray, rot, and slough off. Small brown a 3-or 4-year rotation. The host range now
flower spots enlarge to include whole petals, includes many plants such as carrot, banana, and
causing conspicuous blighting. The fungus foliage plants.
apparently winters on seed and in soil. Phoma chrysanthemi (Telemorph, Didymella
Control Clean up refuse; use a long rotation if ligulica) (formerly Ascochyta chrysanthemi
growing plants commercially. (telemorph, Mycosphaerella ligulicola)).
106 Blights

Ascochyta Ray Blight of chrysanthemum, scattered throughout the stromatic tissue, or

a conspicuous and rapid disease of ray flowers. seated on the surface, or like perithecia, as in
If young buds are infected, the head does not Botryosphaeria ribis. In B. ribis there are two
open; if the attack is later, there may be one- pycnidial forms, a Dothiorella stage containing
sided development of flowers. A tan or brown very small spores that may function as male cells
discoloration proceeds from the base toward the and a Macrophoma stage containing larger
tip of each individual flower, followed by with- spores, one-celled, hyaline, functioning as other
ering. Upper portions of stems and receptacles conidia.
may turn black. Keep plants well spaced; avoid Botryosphaeria ribis var. chromogena Current
overhead watering and excessive humidity. Cane Blight, Canker, Dieback of currant,
flowering currant, gooseberry, apple, rose, and
many other plants (also ▶ Cankers and Die-
Balansia backs). There are two forms of this species, one
being a saprophyte developing on already dying
Ascomycetes, Hypocreales, tissue. The parasitic form chromogena is so
Clavicipitaccae. named from its developing a purple-pink color
when grown on starch paste. There are also
Balansia cyperi Diseased Inflorescence, Blight a number of pathogenic strains, varying from
of purple nutsedge; fungus is systemic and trans- high to low in virulence. Some currant varieties
mitted through tubers. are quite resistant, but the widely grown Wilder
and Red Lake are rather susceptible.
Dieback and death of fruiting branches occur
Beniowskia as the berries are coloring, with leaves wilting
and fruit shriveling. Later in the season small,
Deuteromycetes, Hyphomycetes dark, wartlike fruiting bodies appear in rather
definite parallel rows on the diseased canes.
Hyphae are coiled at the periphery of mature
Rose canes show a similar dying back and wilting
sporodochia; spherical spores are borne on short above a canker. The fungus winters in the canes;
denticles. ascospores infect new shoots; secondary infec-
Beniowskia sphaeroidea Blight of knotroot
tion is by spores oozing from pycnidia. The
bristlegrass. mycelium grows downward through bark and
wood to the main stem, which it encircles and
Control Cut out and burn diseased canes as soon
as noticed. Take cuttings from healthy bushes.
Deuteromycetes, Coelomycetes

Pycnidia black, ostiolate, erumpent, stromatic,

confluent; conidiosphores simple, short; conidia
dark and 2-celled, ovoid to elongate
Ascomycetes, Helotiales,
Stroma a typical black sclerotium, loaf-shaped or
Ascomycetes, Dothideales hemispherical, just on or beneath cuticle or epi-
dermis of plant and firmly attached to it;
Asci in locules in a stroma; spores one-celled, apothecia cupulate, stalked, brown; ascospores
hyaline, eight in an ascus. There is a good deal hyaline, one-celled; conidiophores and conidia
of variation in the genus. The locules may be of the Botrytis cinerea type.
Botrytis 107

Botryotinia fuckeliana The apothecial stage of

Botrytis cinerea, the connection having been
made with isolates from grape, apple, celery,
and potato. The name of the conidial stage is
still widely used for the pathogen causing gray
mold blights.
Botryotinia ricini Gray Mold Blight of castor
bean, Soft Rot of caladium. A pale to olive gray
mold develops on castor-bean inflorescence, and
when fading flowers drop onto stem and leaves,
they are infected in turn.


Deuteromycetes, Hyphomycetes
Fig. 2 Botrytis Petal Spot on Magnolia
Egglike conidia hyaline, one-celled, are formed on
branched conidiophores over the surface, not in throwing out half a box of strawberries or rasp-
special fruiting bodies (see Fig. 1). The arrange- berries. But in continued humid weather the
ment of the spores gives the genus its name, from blight appears on fruits before harvest.
the Greek botrys, meaning a cluster of grapes. Blackberries in the Northwest are subject to
Flattened, loaf-shaped, or hemispherical black gray mold. The fungus winters in blighted blue-
sclerotia are formed on or just underneath cuticle berry twigs, and spores infect blossom clusters.
or epidermis of the host and are firmly attached to Vegetables are commonly afflicted as seed-
it. These sclerotia, with a dark rind and light inte- lings grown in greenhouses and in storage after
rior made up of firmly interwoven hyphae, serve as harvest. If lettuce plants are set in the garden too
resting bodies to carry the fungus over winter. close together, they may blight at the base in
Microconidia, very minute spores that are moist weather, as will endive and escarole. Gray
spermatia or male cells, function in the formation mold is common on lima beans, is sometimes
of apothecia in the few cases where a definite found on snap and kidney beans. In rainy or
connection has been made between the Botrytis foggy periods globe artichoke may be covered
stage and the ascospore form, Botryotinia. with a brownish gray, dusty mold, with bud scales
Botrytis species are the common gray molds, rotten. Asparagus shoots are sometimes blighted,
only too familiar to every gardener. Some are tomato stems rotted.
saprophytic or weakly parasitic on senescent Some of the ornamentals on which Botrytis
plant parts on a wide variety of hosts; others are cinerea is troublesome are given in the following
true parasites and cause such important diseases annotated list:
as peony blight, lily blight, tulip fire. African violet –leaf and stem rot, cosmopoli-
Botrytis cinerea Gray Mold Blight, Bud and tan in greenhouses.
Flower Blight (see Fig. 2), Blossom Blight, Gray Amaryllis –gray mold, mostly in the South, on
Mold Rot, Botrytis Blight of general distribution outdoor plants after chilling.
on a great many flowers, fruits and vegetables. Anemone – occasional severe rotting of
There are undoubtedly many strains of this fun- crowns.
gus and perhaps more than one species involved, Arborvitae –twig blight.
but they have not been definitely separated. Aster – brown patches in flower heads of
This gray-mold disease is common on soft ripe perennial aster; gray mold on flowers of China
fruits after picking, as any cook knows after aster grown for seed in California.
108 Blights

Begonia – dead areas in leaves and flowers Rose – bud or flower blight, cane canker.
rapidly enlarging and turning black in a moist When half-open buds ball, the cause is often an
atmosphere; profuse brownish gray mold. infestation of thrips; but if gray mold is present,
Calendula –gray-mold blight. Botrytis is indicated. Canes kept too wet by
Camellia – flower and bud blight, common a manure mulch, or wet leaves, or injured in
after frost. some way, are often moldy.
Carnation – flower rot or brown spotting, Snapdragon – flower spikes wilt; tan cankers
worse in a cool greenhouse. girdle stems.
Century plant –gray mold after overwatering Sunflower –bud rot and mold.
and chilling. Sweet pea –blossom blight.
Chrysanthemum – cosmopolitan on flowers, Viola spp. –gray mold and basal rot of violet
buds, leaf tips, and cuttings. Ray blight on flowers and pansy.
starts as small, water-soaked spots, which rapidly Zinnia –petal blight, head blight, moldy seed.
enlarge with characteristic gray mold. Botrytis cinerea may also infect arabis, ciner-
Dahlia –bud and flower blight. aria, eucharis, euphorbia, fuchsia, gerbera, gyp-
Dogwood – flower and leaf blight. In wet sophila, heliotrope, hydrangea, iris, lilac, lupine,
springs anthers and bracts of aging flowers are May-apple, pyrethrum, periwinkle, rose-of-Sha-
covered with gray mold, and when these rot down ron, stokesia, viburnum, and wallflower.
on top of young leaves, there is a striking leaf Control Sanitation is more important than any-
blight. thing else. Carry around a paper bag as you
Eupatorium – stem blight, common in inspect the garden; put into it all fading flowers
crowded plantings. A tan area girdles stem near and blighted foliage; if infection is near the base,
ground with tops wilting or drying to that point. take the whole plant up for burning. Keep green-
Geranium (Pelargonium) – blossom blight house plants widely spaced, with good ventila-
and leaf spot, most common in cool, moist green- tion; avoid syringing, overhead watering, and too
houses where plants are syringed frequently. cool temperature. Propagate cuttings from
Petals are discolored, flowers drop, gray mold healthy plants in a sterilized medium.
forms on leaves. Botrytis douglasii Seedling Blight of giant
Lily – Botrytis cinerea is common on lilies, but sequoia and redwood, perhaps a form of B.
see also ▶ B. elliptica. cinerea.
Marigold –gray mold prevalent on fading Botrytis elliptica Lily Botrytis Blight, general on
flowers. lilies, also reported on tuberose and stephanotis in
Peony – late blight, distinguished from early California. Lily species vary in susceptibility to
blight (see ▶ B. paeoniae) by the sparse mold, the disease, but there are several strains of the
usually standing far out from affected tissues, fungus, and few lilies are resistant to all strains.
rather than a thick, short velvety mold, and by Madonna lily, L. candidum, is particularly sus-
much larger, flatter sclerotia formed near base of ceptible, with infection starting in autumn on the
the stalk. Late flowers are infected, and when rosette of leaves developed at that time.
they drop down onto wet foliage, irregular If the blight strikes early, the entire apical
brown areas are formed in leaves. growth may be killed with no further develop-
Pine – seedling blight. ment. More often the disease starts as a leaf spot
Pistachio – shoot blight. when stems are a good height. Spots are orange to
Poinsettia – tip blight and stem canker. reddish brown, usually oval. In some species
Primrose – crown rot and decay of basal there is a definite red to purple margin around
leaves, with prominent gray mold, very common a light center; in others the dark margin is
in greenhouses where plants are heavily watered. replaced by an indefinite water-soaked zone. If
Rhododendron – flower, twig, and seedling spots are numerous, they grow together to blight
blight. the whole leaf. Infection often starts with the
Botrytis 109

lowest leaves and works up the stem until all moist weather the whole flower turns brown and
leaves are blackened and hanging limp. This is slimy. Flowers with no visible spotting when
the result of many spot infections and not from an packed often arrive ruined. After the flowers are
invasion of the vascular system. cut, infection spreads down the stalk and into the
Buds rot or open to distorted flowers with corm, producing dark brown spots, irregular in
irregular brown flecks. There are sometimes shape and size, most numerous on the upper sur-
severe stem lesions, but the rot rarely progresses face. Corms may become soft and spongy with
into the bulbs. Spores formed in the usual gray- a whitish mold. Oval, flat, black sclerotia, 1/8 to
mold masses in blighted portions are spread by 1/4 inch long, are formed on corms in storage and
rain, air currents, and gardeners. Optimum spore in rotting tissue in the field or in refuse piles. They
germination is in cool weather, around 60  F, but may persist in the soil several years.
once infection has started 70  F promotes most Control Cure corms rapidly after digging; bury
rapid blighting. With sufficient moisture the or burn all plant refuse.
cycle may repeat every few days through the Botrytis hyacinthi Hyacinth Botrytis Blight
season. The fungus winters as very small black recently found in Washington on plants grown
sclerotia, irregular or elliptical in shape, in fallen from imported bulbs. Leaves have brown tips
flowers or blighted dead stems and leaves, or as with gray mold or brown spots on lower surface.
mycelium in the basal rosette of Madonna lilies. Leaves may be killed, with small black sclerotia
Control Avoid too dense planting, and shady or formed in rotting tissue. Flowers rot and are cov-
low spots with little air circulation and subject to ered with powdery gray spores. Do not work with
heavy dews. Clean up infected plant parts before plants when they are wet; remove infected parts
sclerotia can be formed. Copper sprays are more or whole plants.
effective for the lily Botrytis than the newer Botrytis narcissicola ▶Sclerotinia
organics. Spray with bordeaux mixture; start narcissicola, under Rots.
when lilies are 5 or 6 inches high and continue Botrytis paeoniae Peony Botrytis Blight, Early
at 10-to 14-day intervals until flowering. Blight, Bud Blast, Gray Mold, probably present
Botrytis galanthina Botrytis Blight of snow- wherever peonies are grown. It is also recorded
drop, sometimes found in the sclerotial state on on lily-of-the-valley, but that may be a form of
imported bulbs. If the black dots of sclerotia are Botrytis cinerea. Peony blight was first noticed in
present only on outer scales, remove scales epiphytotic form in this country in 1897 and has
before planting; otherwise discard bulbs. been important in wet springs ever since.
Botrytis gladiolorum Gladiolus Botrytis Blight, Young shoots may rot off at the base as they
Corm Rot, first reported in Oregon in 1939 and come through the ground or when a few inches
now serious in all important gladiolus-growing high, with a dense velvety gray mold on the
areas – the Pacific Coast, the Midwest, rotting portions. This early shoot blight is far
Florida – in cool, rainy weather. In northern more common when the young stems are kept
areas the disease is a corm-rotting problem, in moist by having to emerge through a mulch of
the South a flower blight, damaging in transit, and manure or wet leaves. Flowers are attacked at any
in all areas it is a leaf spot or blight. stage. Buds turn black when they are very tiny,
In dry weather and in more resistant varieties never developing, or they may be blasted when
the leaf spots are very small, rusty brown, they are half open. If it is dry in early spring,
appearing only on the exposed side of the leaf. infection may be delayed until flowers are in
In more humid weather the spots are large, full bloom, at which time they turn brown. Infec-
brown, round to oval or smaller, pale brown tion proceeds from the flower down the stem for
with reddish margins. Flower stems have pale a few inches, giving it a brown and tan zoned
brown spots that turn dark. There may be a soft appearance. Leaf spots develop when infected
rotting at the base of florets. The disease starts on petals fall on foliage. Continued blighting of
petals as pinpoint, water-soaked spots, but in leaves through the summer and late blasting of
110 Blights

flowers may be due to Botrytis cinerea, which

produces a sparser mold and conidiophores
projecting farther from the petal or leaf surface.
Conidia are blown by wind, splashed by rain,
carried on gardeners’ tools, and sometimes
transported by ants. Secondary infection is abundant
in cool moist weather. In late summer small, shiny
black, slightly loaf-shaped sclerotia are formed near
the base of stalks, just under the epidermis. They are
quite different from the large, flat, black sclerotia
often formed by B. cinerea on the same stalks.
Control Sanitation is the most important step. Fig. 3 Botrytis Blight on Tulip
Cut down all tops in autumn at ground level, or
just below, to get rid of sclerotia wintering near flower stems or bulbs. Sometimes the latter have
base of stems. Burn this debris; never use it for yellow to brown, slightly sunken, circular lesions
a mulch. Avoid any moisture-retentive covering. on outermost fleshy scales without the formation
If you insist on manure, apply it in a wide ring of sclerotia. Spring infection comes from spores
around the plant, well outside the area of emerg- produced on such bulbs or from sclerotia on bulbs
ing shoots. Go around with a paper bag periodi- or sclerotia left loose in the soil after infected
cally, cutting off for burning all blighted parts; tissues have rotted.
never carry these parts loose through the garden Control Inspect all bulbs carefully before plant-
for fear of shedding spores to healthy plants. ing; discard those harboring sclerotia or suspi-
Botrytis polyblastis ▶Sclerotinia polyblastis. cious brown lesions. It is wise, though seldom
Botrytis streptothrix (see ▶Streptobotrys possible in a small garden, to plant new bulbs
arisaemae) (teleomorph state Streptotinia where tulips have not grown for 3 years. Plant
arisaemae). Leaf and Stalk Blight of Jack-in- where there is good air circulation. Make periodic
the-pulpit and golden club. inspections, starting early, removing into a paper
Botrytis tulipae Tulip Fire, Botrytis Blight of bag plants with serious primary infection and
tulips, general wherever tulips are grown, causing blighted leaves. Cut off all fading flowers before
much damage in rainy springs. The first indica- petals fall; cut off all foliage at ground level when
tion of disease is the appearance of a few mal- it turns yellow. Burn all debris.
formed leaves and shoots among healthy tulips or Streptobotrys arisaemae (formerly Botrytis
large light patches resembling frost injury on streptothrix) (teleomorph state Streptotinia
leaves. Gray mold forming on such blighted arisaemae). Leaf and Stalk Blight of Jack-in-the-
areas of plants grown from infected bulbs pro- pulpit and golden club. This species has conidio-
vides an enormous number of conidia to be phores with strikingly twisted branches, produc-
splashed by rain to nearby tulips. Secondary ing a reddish brown mat of conidia. Sclerotia are
infection appears as minute, slightly sunken, yel- very small, seldom over 1/32 inch, black, shiny,
lowish leaf spots, surrounded with a water- and somewhat hemispherical.
soaked area, and gray to brown spots on stems,
often zonate, and resulting in collapse. Small
white spots appear on colored flowers, brown Briosia
spots on white petals (see Fig. 3); but with con-
tinued moisture the spots grow together, and in Deuteromycetes, Hyphomycetes
a day or so the fuzzy gray mold has covered rotten
blooms and large portions of blighted leaves. Conidia on synnemata or coremia, erect fascicles
Very small, shiny black sclerotia are formed in of hyphae ending in a small head; spores globose,
leaves and petals rotting into the ground, or on old dark, one-celled, catenulate (formed in chains).
Cercospora 111

Briosia azaleae (see ▶Pycnosystanus azalea) Infection starts near a terminal bud in late
(Pycnostysanus azaleae). Bud and Twig Blight summer and progresses down a twig into
of azalea and rhododendron, widespread but a node, sometimes beyond into 2-year wood.
occasional. The needles redden and die; they are conspicuous
Pycnosystanus azalea (formerly Briosia in spring but drop in late summer. Then brown to
azaleae) (Pycnostysanus azaleae). Bud and black apothecia with a greenish surface to the cup
Twig Blight of azalea and rhododendron, wide- appear on twigs. Cut off and destroy infected
spread but occasional. The disease was reported twigs.
from New York in 1874 and, as a rhododendron
bud rot, from California in 1920. It was particu-
larly serious on Massachusetts azaleas in 1931 Cercospora
and 1939. Flower buds are dwarfed, turn brown
and dry; scales are silvery gray. Twigs die when Deuteromycetes, Hyphomycetes
lateral leaf buds are infected. Successive crops of
coremia are produced on old dead buds for as Conidia hyaline to pale to medium green or
long as three years, the first crop appearing the brown; long, usually with more than three cross
spring after summer infection. The coremia heads walls; straight or curved, with the base obconate
are dark, and the buds look as if stuck with tiny, or truncate, tip acute to obtuse; thin-walled; not
round-headed pins. Prune out and burn infected formed in a fruiting body but successively on
buds and twigs in late autumn and early spring. slender conidiophores, which emerge in fascicles
Spraying with bordeaux mixture before or groups from stomata and usually show joints or
blossoming and at monthly intervals after bloom scars where conidia have fallen off successively.
may be wise in severe cases. The conidiophores are always colored, oliva-
ceous to brown, pale to very dark (Fig. 1).
This is the largest group of the Dematiaceae,
Calonectria with about 400 species, all parasitic, causing leaf
spots or blights. The teleomorph state, when
▶Cylindrocladium under Blights. known, is Mycosphaerella.
Calonectria colhounii Blight on Cercospora apii Early Blight of celery, general
Leucospermum. on celery and celeriac, first noted in Missouri in
1884 and since found in varying abundance wher-
ever celery is grown. The disease is most severe
Cenangium from New Jersey southward. The name is some-
what misleading; in Florida early blight rarely
Ascomycetes, Helotiales appears before the Septoria disease known as
late blight. Foliage spots appear when plants are
Apothecia small, brown to black, sessile or about 6 weeks old. Minute yellow areas change to
substipitate on bark; spores hyaline, elliptical, large, irregular, ash gray lesions, covered in moist
one-celled; paraphyses filiform. weather with velvety groups of conidiophores
Cenangium ferruginosum Pine Twig Blight, and spores on both sides of leaves. Sunken, tan,
Pruning Disease; Cenangium Dieback of fir elongated spots appear on stalks just before har-
and pine. The fungus is ordinarily saprophytic vest. The disease spreads rapidly in warm, moist
on native pines but may become parasitic weather, the spores being splashed by rain, car-
when their vigor is reduced by drought. The ried with manure or cultivators, or blown by
disease is considered beneficial to ponderosa wind. The life cycle is completed in 2 weeks.
pine in the Southwest because it prunes off the Control Seed more than 2 years old is probably
lower branches; on exotic pines it can be free from viable spores; other seed should be
damaging. treated with hot water, 30 min at 118 to 120  F.
112 Blights

Bordeaux mixture and other copper sprays be purplish; the fungus fruits on underside of
have been recommended. Spray applications leaves. The blight is seldom important enough
should start soon after plants are set and be to warrant control measures.
repeated weekly, or more often. Emerson Pascal
is blight-resistant.
Cercospora carotae Early Blight of
carrot. Lesions on leaves and stems are
subcircular to elliptic, pale tan to gray or
Cercosporidium punctum Stem and Foliage
brown or almost black; lobes or entire leaflets
Blight of fennel.
are killed. The disease is more severe on young
leaves and builds up as the plant grows. Spores,
produced on both leaf surfaces, are spread by
wind. Choanephora
Control Rotate crops and clean up refuse.
Cercospora microsora Linden Leaf Blight, Zygomycetes, Mucorales
general on American and European linden.
Small circular brown spots with darker borders Mycelium profuse; sporangia and conidia pre-
coalesce to form large, blighted areas, often sent; sporangiola lacking. Sporangium pendent
followed by defoliation; most serious on young on recurved end of an erect, unbranched sporan-
trees. giophore with a columella, containing spores
Cercospora sequoiae Arborvitae Blight, Fire provided at both ends and sometimes at the side
Blight, on oriental arborvitae and Italian cypress with a cluster of fine, radiating appendages.
in the South; destructive in ornamental plants. Conidia formed in heads on a few short branches
First reported from Louisiana in 1943, the fungus or an erect conidiophore enlarged at the
was named as a new species of Cercospora in tip; conidia longitudinally striate, without
1945, but it is nearer Heterosporium in spore appendanges.
character. Affected leaves and branchlets are Choanephora cucurbitarum Blossom Blight,
killed, turn brown, and gradually fall off, leaving Fruit Rot, common on summer squash and pump-
shrubs thin and ragged. The lower two-thirds of kin, occasional on amaranth, cowpea, cucumber,
the bush is affected most severely, with a tuft of okra, and pepper; on sweet potato foliage, on
healthy growth at the top. When close to a house, fading hibiscus, on vinca, and other flowers.
the side away from the wall shows most symp- This blight is often found in home gardens in
toms. Plants crowded in nurseries are killed in seasons of high humidity and rainfall. Flowers
1 to 3 years, but in home gardens they may persist and young fruits are covered with a luxuriant
for years in an unsightly condition. Conidio- fungus growth, first white, then brown to purple
phores in fascicles produce conidia after girdling with a definite metallic luster. The fruiting bodies
cankers have killed the twigs. There is often look like little pins stuck through this growth.
a swelling above the girdle that resembles an Both staminate and pistillate flowers are infected,
insect gall. and from the latter the fungus advances into
Cercospora sordida (Mycosphaerella tecomae) young fruits, producing a soft wet rot at the blos-
(see ▶Pseudocercospora sordida)Trumpetvine som end. In severe cases all flowers are blighted
Leaf Blight from New Jersey to Iowa and or fruits rotted.
southward. Control Grow plants on well-drained land;
Pseudocercospora sordida (formerly rotate crops. Remove infected flowers and fruits
Cercospora sordida) (Mycosphaerella tecomae). as noticed.
Trumpetvine Leaf Blight from New Jersey to Choanephora infundibulifera Blossom
Iowa and southward. Small, angular, sordid Blight on hibiscus and jasmine. Leaf Blight on
brown patches run together; edge of leaflets may soybean.
Coryneum 113

Colletotrichum acutatum Twig Blight and

Ciboria Fruit Spot on dogwood
Colletotrichum gloeosporioides Seedling
Ascomycetes, Helotiales, Blight of papaya.
Sclerotiniaceae Colletotrichum dematium Twig Blight on
Stroma a dark brown to black sclerotium in cat-
kins or seed, simulating in shape the stromatized
organ and not resembling a sclerotium externally. Corticium
Apothecia cupulate to shallow saucer-shaped;
brown. Basidiomycetes, Aphyllophorales
Ciboria acerina Maple Inflorescence Blight on
red and silver maple. Apothecia, developed in Hymenium or fruiting surface of basidia
great numbers from stromatized inflorescences consisting of a single resupinate or horizontal
on ground beneath trees, start discharging spores layer. This genus has contained a rather hetero-
when maple flowers appear overhead. Mycelium geneous collection of species; some of the more
spreads through stamens, calyx, and bud scales important have been transferred to the genus
until flower cluster drops. Pellicularia.
Ciboria carunculoides Popcorn Disease of Corticium koleroga Thread Blight.
mulberry, a southern disease, not very important. ▶Pellicularia koleroga.
Sclerotia are formed in carpels of fruit, which Corticium microsclerotia Web Blight.
swells to resemble popcorn but remains green. ▶Pellicularia filamentosa.
Corticium salmonicolor (see ▶Erythricium
salmonicolor). Limb Blight of fig, pear, apple in
Ciborinia Gulf States.
Corticium stevensii Thread Blight.
Ascomycetes, Helotiales, ▶Pellicularia koleroga.
Sclerotiniaceae Corticium vagum, now Pellicularia
filamentosa, teleomorph state of Rhizoctonia
Stroma a thin, flat, black sclerotium of discoid solani, causing black scurf of potatoes and
type in leaves; one to several stalked apothecia damping-off and root rot of many plants. See
arise from sclerotia; apothecia small, brown, both ▶Pellicularia and ▶Rhizoctonia under Rots.
cupulate to flat when expanded. Erythricium salmonicolor (formerly
Ciborinia erythronii and C. gracilis Leaf Corticium salmonicolor). Limb Blight of fig,
Blight of erythronium. Flat black sclerotia are pear, apple in Gulf States. The spore surface is
prominent in leaves. pinkish.

▶ Blackleg.
Cladosporium cladosporioides Blossom Deuteromycetes, Coelomycetes
Blight on strawberry.
Acervuli subcutaneous or subcortical, black,
cushion-shaped or disc-shaped; conidiophores
Colletotrichum slender, simple; spores dark with several cross
walls, oblong to fusoid; parasitic or saprophytic
▶ Anthracnose. (see Fig. 1, chapter ▶ Cankers and Diebacks).
114 Blights

Coryneum berckmansii (see ▶Seimatosporium formed on foliage, dropping out to leave typical
berckmansii). Coryneum Blight of Oriental arbor- shot holes, followed by considerable defoliation.
vitae, also on Italian cypress, causing serious Apricot buds are blackened and killed during
losses in nurseries and home gardens in the Pacific winter; fruiting wood in peaches is killed before
Northwest. growth starts. In late rains leaves and fruit are
Coryneum microstictum (see ▶Seimatosporium peppered with small, round, dead spots. Fruit
lichenicola). Twig Blight of American bladdernut. lesions are raised, roughened, scabby. The fungus
Coryneum carpophilum (Cladosporium winters in twigs, diseased buds and spurs.
beijerinckii) (see ▶Stigmina carpophila, Control In California, the standard spray for
Anamorph, Cladosporium beijerinckii). Peach peach is bordeaux mixture applied in autumn
Shoot Blight, Coryneum Blight of stone fruits, immediately after leaf fall and before the rainy
Shot Hole, Fruit Spot, Winter Blight, Pustular season. On apricots additional sprays are suggested
Spot, general on peach in the West, also on for late January and at early bloom. On almonds at
almond, apricot, nectarine, and cherry. least two spring sprays are recommended, one at
Seimatosporium berckmansii (formerly the popcorn stage of bloom, the other at petal fall.
Coryneum berckmansii). Coryneum Blight of
Oriental arborvitae, also on Italian cypress, caus-
ing serious losses in nurseries and home gardens Cryptocline
in the Pacific Northwest. Small twigs or branches
are blighted, turn gray-green then reddish brown; Deuteromycetes, Coelomycetes
many small branchlets drop, leaving a tangle of
dead gray stems; larger limbs may be girdled. Cryptocline cinerescens Twig Blight of oaks.
Twigs are dotted with black pustules bearing
five-septate spores. As new growth develops
in blighted areas, the spores spread the disease Cryptospora
to young contiguous foliage. Reinfection
continues until the plant is so devitalized it dies. Scomycetes, Amphisphaeriales
The fungus fruits only on scale leaves or young
stems. Perithecia immersed in a stroma, with long necks
Control Remove and destroy blighted twigs. converging into a disc; ascospores long, filiform,
Apply a copper spray in September to healthy hyaline; conidia on a stroma.
bushes as a preventive spray; apply in September Cryptospora longispora (see ▶Servazziella
and repeat in late October to infected bushes. longispora). Araucaria Branch Blight.
Seimatosporium lichenicola (formerly Servazziella longispora (formerly Cryptospora
Coryneum microstictum). Twig Blight of Ameri- longispora). Araucaria Branch Blight. Lower
can bladdernut. Young twigs are killed; the fun- branches are attacked first, with disease spread-
gus winters in acervuli on this dead tissue, and ing upward; tip ends are bent and then broken off;
spores are disseminated in spring. Prune out and plants several years old may be killed. Prune off
burn diseased twigs during the winter. and burn infected branches.
Stigmina carpophila (formerly Coryneum
carpophilum (Cladosporium beijerinckii)).
Peach Shoot Blight, Coryneum Blight of stone Cryptostictis
fruits, Shot Hole, Fruit Spot, Winter Blight, Pus-
tular Spot, general on peach in the West, also on Deutermycetes, Coelomycetes
almond, apricot, nectarine, and cherry. Twig lesions
are formed on 1 -year shoots, reddish spots devel- Spores dark, with several cross walls, formed in
oping into sunken cankers; fruit buds are invaded, acervuli.
and there is copious gum formation. Small spots are Cryptostictis sp. Twig Blight of dogwood.
Cylindrosporium 115

a week in warm rainy weather, and the

Curvularia fungus can survive in the soil for 3 years.
Many gladiolus varieties are more or less
Deuteromycetes, Hyphomycetes resistant; Picardy and some others are very
Conidiophores brown, simple or sometimes
branched, bearing conidia successively on new
growing tips; conidia dark, three-to five-celled,
with end cells lighter, more or less fusiform, Cylindrocladium
typically bent or curved with central cells
enlarged; parasitic or saprophytic. Deuteromycetes, Hyphomycetes
Curvularia cymbopogonis Blight and Leaf
Spot of itchgrass. Leafspots coalesce after 3 or Conidiophores dichotomously branched; spores
4 days to form larger lesions and final blighting hyaline, two-or several-celled.
symptoms. Cylindrocladium clavatum Blight on bird-of-
Curvularia lunata (C. trifolii f. sp. gladioli). paradise.
Gladiolus Flower Blight and Leaf Spot, Cylindrocladium scoparium Cylindrocladium
Curvularia Disease. Suddenly, in 1947, a blight Blight. Damping-off of seedlings and
showed up in Florida as a serious threat to the cuttings – conifers, azalea, magnolia, hydrangea,
gladiolus cut-flower industry, ruining hundreds holly, pyracantha, bottle brush, and poinsettia –in
of acres there and in Alabama in the next few greenhouses under very moist conditions.
months. The disease is now recorded as far north Infected azalea leaves turn black, with petiole
as New York and Wisconsin and on the Pacific bases softened, and drop in a few days; the bark
Coast. The pathogen is usually identified as turns brown. Leaves and stems are covered with
Curvularia lunata, known as a crop pest for brownish mycelial strands and white powdery
many years, especially in the tropics, but studies masses of conidia. Control by proper humidity
indicate it is a special form of C. trifolii, cause of and aeration.
a leaf spot of clover. Cylindrocladium avesiculatum Blight and
Curvularia spots on leaf or stem are oval, tan Leaf Spot of Leucothoe¨ axillaris.
to dark brown, showing on both sides of the leaf,
bordered with a brown ring, slightly depressed
and with a narrow yellowish region between the
spot and normal green of the leaf. Tan centers of Cylindrosporium
spots are covered with black spores resembling
powder. Premature death comes when stems of Deuteromycetes, Coelomycetes
young plants are girdled; florets fail to open when
petioles are girdled. Acervuli subepidermal, white or pale, discoid or
Under favorable weather conditions tan spots spread out; conidiophores short, simple; conidia
on petals turn into a smudgy flower blight. hyaline, filiform, straight or curved, one-celled or
Brown to black irregular lesions appear on becoming septate; parasitic on leaves. Many spe-
corms of blooming stock and develop further in cies have Higginsia or Coccomyces as
storage; the fungus survives in corms from one a teleomorph state.
season to the next. This is a high temperature Cylindrosporium defoliatum Leaf Blight of
fungus, with optimum for growth 75 to 85  F Hackberry. May cause defoliation but usually
and no infection under 55  F. A 13-h dew period unimportant.
is sufficient moisture. Leaf spots show up in 4 to Cylindrosporium griseum On western
5 days, spots on florets and stems in only 2 to soapberry.
3 days. The complete life cycle is as short as Cylindrosporium juglandis On walnut.
116 Blights

several inches upward and down into roots.

Delphinella Scattered dark pycnidia are present in stems, pet-
ioles, leaf blades, and seed capsules, the latter
Ascomycetes, Dothideales probably spreading the blight. Crowns are some-
times developed in a cottony weft of mycelium;
Delphinella balsameae Tip Blight of fir. perithecia develop on decaying stems. Remove
and destroy diseased plants; use seed from
healthy plants.
Dendrophoma Diaporthe phaseolorum Lima Bean Pod
Blight, Leaf Spot, apparently native in New Jer-
Deuteromycetes, Coelomycetes sey, where it was first noticed in 1891, more
abundant on pole than on bush beans. Leaf spots
Pycnidia dark or light brown, superfcial or sub- are large, irregular, brown, often with discolored
merged and erumpent; globose or elongate, borders and large black pycnidia formed in con-
ostiolate; conidiophores elongated, branched; centric circles in dead tissue. Necrotic portions
conidia hyaline, one-celled, elongate to ellipsoid; may drop out, making leaves ragged.
parasitic or saprophytic. Pod lesions spread; pods turn black and wilted,
Dendrophoma obscurans (see ▶Phomopsis with prominent black pycnidia. Seeds are shriv-
obscurans). Strawberry Leaf Blight, Angular eled or lacking. Spores are produced in great
Leaf Spot. The lesions are large, circular to angu- numbers, are disseminated by wind and pickers,
lar, reddish purple, zonate with age, having a dark and enter through stomata or wounds. The dis-
brown center, a light brown zone, and a purple ease is most severe along the coast; optimum
border. Spots may extend in a V-shaped area temperature is around 80  F. The fungus is seed-
from a large vein to edge of the leaf, with black borne, but most lima bean seed is produced where
fruiting bodies appearing in the central portion. the disease does not occur. Use healthy seed;
Not serious before midsummer, the disease may clean up refuse; rotate crops.
be destructive late in the season. The fungus Diaporthe phaseolorum var. sojae Soybean
winters on old leaves. Pod and Stem Blight, widespread. This disease
Phomopsis obscurans (formerly Dendrophoma was formerly confused with the more acute stem
obscurans). Strawberry Leaf Blight, Angular canker caused by D. phaseolorum var. caulivora
Leaf Spot. (▶ Rots). The pod blight is a slower disease,
killing plants in later stages of development. It
can be identified by the numerous small black
Diaporthe pycnidia scattered over the pods and arranged in
rows on stems. The blight is more serious in wet
Ascomycetes, Diaporthales seasons. The fungus winters on the seed and on
diseased stems in the field. Use clean seed; clean
Perithecia in a hard black strom a made up of host up plant refuse; rotate crops.
and fungal elements, first immersed, then Diaporthe phaseolorum var. caulivora Stem
erumpent; ascospores fusoid or ellipsoid, two- Blight of soybean; also causes pod and seedling
celled, hyaline. Anamorph state a Phomopsis blight, stem canker, and seed decay of soybean.
with two types of spores; alpha conidia, hyaline, Diaporthe vaccinii Blueberry Twig Blight. The
one-celled ovate to fusoid, and beta conidia, same fungus that causes cranberry rot blights new
curved or bent stylospores. shoots of cultivated blueberries, entering at tips,
Diaporthe arctii Diaporthe Blight of Larkspur, progressing toward the base, and ultimately gir-
Stem Canker, on annual larkspur and delphin- dling old branches. Pycnidia develop on leaves
ium. Lower leaves turn brown and dry but remain and dead twigs. The disease is seldom serious
attached; brown lesions at base of stems extend enough for control measures.
Didymella 117

Diaporthe vexans Phomopsis Blight of Egg- Control Spray small trees and nursery stock sev-
plant, Fruit Rot, general in field and market, espe- eral times during summer and fall with bordeaux
cially in the South. Destruction is often complete, mixture.
with every above-ground part affected. Seedlings Didymascella tsugae (see ▶Fobrella tsugae).
rot at ground level. The first leaf spots are near the Hemlock Needle Blight. Needles of Canada hem-
ground, definite, circular, gray to brown areas lock turn brown and drop in late summer. Spores
with light centers and numerous black pycnidia. are matured in apothecia on fallen needles with
The leaves turn yellow and die. Stem cankers are new infection in spring. The damage is not heavy.
constrictions or light gray lesions. Fruit lesions Fobrella tsugae (see ▶Didymascella tsugae).
are pale brown, sunken, marked by many black Hemlock Needle Blight. Needles of Canada hem-
pycnidia arranged more or less concentrically. lock turn brown and drop in late summer.
Eventually the whole fruit is involved in a soft
rot or shriveling. Spores winter on seed and in
contaminated soil. There is no fungicidal control. Didymella
Use resistant varieties Florida Market and Florida
Beauty. Ascomycetes, Sphaeriales,

Dichotomophora Perithecia (or perithecia-like stromata) membra-

nous, not carbonaceous; innate; not beaked;
▶ Cankers and Diebacks. paraphyses present; spores two-celled, hyaline.
Dichotomophora lutea Stem Blight, of com- Didymella applanata Raspberry Spur Blight,
mon parsley. Purple Cane Spot, Gray Bark, general on rasp-
berries, also on dewberry, blackberry. Named
because it partially or completely destroys spurs
Didymascella (Keithia) or laterals on canes. The disease, known in North
America since 1891, may cause losses up to 75 %
Ascomycetes, Phacidiales, of the crop of individual plants of red raspberries.
Stictidiaceae Dark reddish or purple spots on canes at point of
attachment of leaves enlarge to surround leaf and
Apothecia brown, erumpent on leaves of coni- bud and may darken lower portion of cane.
fers; spores dark, two-celled, ovoid; paraphyses Affected areas turn brown, then gray.
filiform; asci two-to four-spored. If buds are not killed outright during the winter,
Didymascella thujina Arborvitae Leaf Blight, they are so weakened that the next season’s spurs
Seedling Blight of arborvitae in eastern states are weak, chlorotic, seldom blossoming. Pycnidia
and of giant arborvitae, sometimes called western of the anamorph Phoma state and perithecia are
red cedar. The fungus is a native of North Amer- numerous on the gray bark; ascospores are
ica and occurs abundantly in the West, damaging discharged during spring and early summer; on
seedlings and saplings, often killing trees up to germination they can penetrate unwounded tissue.
4 years old, if they are in dense stands in humid Control Keep plants well-spaced, allowing plenty
regions. Older trees do not die, but foliage of sunlight for quick drying of foliage and canes.
appears scorched, particularly on lower branches, Remove infected canes and old fruiting canes after
and young leaf twigs may drop. Cushionlike, harvest. A delayed dormant spray of lime sulfur or
olive brown apothecia embedded in leaf tissue, Elgetol may be advisable, followed by two sprays
usually upper, are exposed by rupture of the epi- of ferbam or bordeaux mixture, applied when new
dermis. After summer discharge of spores (round, shoots are 6 to 10 inches high and 2 weeks later.
brown, unequally two-celled) the apothecia drop Didymella bryoniae Gummy stem blight and
out of the needles, leaving deep pits. fruit spot; of watermelon.
118 Blights

Didymosphaeria Dothistroma

Ascomycetes, Dothideales Deuteromycetes, Coelomycetes

Perithecia innate or finally erumpent; not beaked; Stroma dark, elongate, innate, becoming
smooth; paraphyses present; spores dark, two- erumpent and swollen, with a stalk extending
celled. into the substratum, composed internally of
Didymosphaeria populina (Venturia populina, dense, vertical hyphae; locules separate, one to
V. tremulae, V. macularis also cause this disease). several in the upper part of the stroma; conidio-
Shoot Blight of polar, Leaf and Twig Blight. phores simple, slender; conidia several-celled,
Young shoots are blackened and wilted. In hyaline, long-cylindrical to filiform.
moist weather dark olive green masses of spores Dothistroma pini Needle Blight on Austrian
are formed on leaves. pine and red pine.

Cryphonectria (Endothia)
Ascomycetes, Diaporthales
Deuteromycetes, Coelomycetes
Perithecia deeply embedded in a reddish to yel-
Pycnidia innate or finally erumpent; black, single,
low stroma, with long necks opening to the sur-
globose, smooth; ostiole present; conidiophores
face but not beaked; paraphyses lacking; spores
slender, simple; conidia dark, two-celled, ellip-
two-celled, hyaline. Conidia borne in hollow
soid or ovoid. Parasitic or saprophytic.
chambers or pycnidia in a stroma and expelled
Some species cause twig blights which are not
in cirrhi.
too important: Diplodia coluteae on bladder
Cryphonectria parasitica Chestnut Blight,
senna; D. longispora on white oak; Sphaeropsis
Endothia Canker, general on chestnut. To most
sapinea (formerly D. pinea) on pine;
gardeners this disease is of only historical impor-
D. sarmentorum on pyracantha.
tance, for practically all of our native chestnuts
Lasiodiplodia theobromae (formerly Diplodia
are gone. The disease, however, persists in
natalensis) (anamorph state of Physalospora
sprouts starting from old stumps and in the chin-
rhodina) causes blight, stem gumming, or stem-
quapin. One of the most destructive tree diseases
end rot of melons, as well as twig blight of peach
ever known, chestnut blight at least served to
and citrus. See further under ▶ Rots.
awaken people to the importance of plant disease
Diplodia gossypina (see ▶Lasiodiplodia
and to the need for research in this field.
theobromae). Blight of slash pine and loblolly
First noticed in the New York Zoological Park
pine seedlings.
in 1904, the blight rapidly wiped out the chestnut
Lasiodiplodia theobromae (formerly Diplodia
stands in New England and along the Allegheny
gossypina). Blight of slash pine and loblolly pine
and Blue Ridge Mountains, leaving not a single
undamaged tree. In 1925 the disease eliminated
chestnuts in Illinois and by 1929 had reached the
Pacific Northwest.
Discula Conspicuous reddish bark cankers are formed
on trunk and limbs, often swollen and splitting
Deuteromycetes, Coelomycetes longitudinally. As the limbs are girdled, the
foliage blights, so that brown, dried leaves are
Discula quercina, Twig Blight of oaks. seen from a distance. The fungus fruits
Diplocarpon (Fabraea) 119

abundantly in crevices of broken bark, first pro- circular lesion, 1/4 inch or less in diameter, with
ducing conidia extruded in yellow tendrils from the raised black dot of a fruiting body in the
reddish pycnidia and later ascospores from peri- center of each spot. If spots are numerous, there
thecia embedded in orange stromata. Fans of is extensive defoliation. Fruit spots are red at first,
buff-colored mycelium are found under affected then black and slightly sunken; the skin is rough-
bark. ened, sometimes cracked. Quince has similar
Ascospores can be spread many miles by the symptoms.
wind, landing in open wounds, but the Twig lesions appear on the current season’s
sticky conidia are carried by birds and insects. growth about midsummer, indefinite purple or
The fungus can live indefinitely as a saprophyte, black areas coalescing to form a canker. Primary
and new sprouts developing from old spring infection comes more from conidia
stumps may grow for several years before they produced in these twig lesions than from asco-
are killed. spores shot from fallen leaves on the ground.
Control All eradication and protective Most commercial varieties of pear and quince
measures have proved futile. Hope for the are susceptible, although some are moderately
future lies in cross-breeding resistant Asiatic resistant.
species with the American chestnut (and Fabraea maculate, Entomosporium
there has been some success in this line) or in maculatum (▶Diplocarpon mespili,
substituting Chinese and Japanese chestnuts for ▶Entomosporium mespili). Pear Leaf Blight,
our own. Entomosporium Leaf Spot, Fruit Spot, generally
distributed on pear and quince, widespread on
amelanchier, sometimes found on apple, Japa-
Diplocarpon (Fabraea) nese quince, medler, mountain-ash, Siberian
crab, cotoneaster, loquat, photinia.
Ascomycetes, Helotiales, Fabraea thuemenii (Entomosporium
Dermateaceae thuemenii). Hawthorn Leaf Blight, wide-spread
on Crataegus species. Symptoms are similar to
Apothecia develop on fallen leaves; small, those of pear leaf blight and for a long time the
disclike, leathery when dry, gelatinous when pathogen was considered identical. Small dark
wet; asci extend above the surface of the disc; brown or reddish brown spots, with raised black
ascospores two-celled, hyaline. The anamorph dots, are numerous over leaves, which drop pre-
stage an Entomosporium with distinctive cruciate maturely in August. In wet seasons trees may be
four-celled conidia, each cell with an appendage, naked by late August.
formed in acervuli (Fig. 1). Control Because the fungus winters in twig can-
Didymascella tsugae (see ▶Fabrella tsugae). kers as well as in fallen leaves, sanitation has
Hemlock Needle Blight. Needles of Canada hem- little effect. Standard recommendation has been
lock turn brown and drop in late summer. to spray three times with bordeaux mixture,
Diplocarpon mespili (Entomosporium mespili) starting when leaves are half out and repeating
(formerly Fabraea maculate, Entomosporium at 2-week intervals. The copper may be some-
maculatum). Pear Leaf Blight, Entomosporium what phytotoxic, causing small reddish spots sim-
Leaf Spot, Fruit Spot, generally distributed on ilar to those of blight, but it does prevent
pear and quince, widespread on amelanchier, defoliation.
sometimes found on apple, Japanese quince, Fabrella tsugae (formerly Didymascella
medler, mountain-ash, Siberian crab, cotoneas- tsugae). Hemlock Needle Blight.
ter, loquat, photinia. Needles of Canada hemlock turn brown and
Pears may be affected as seedlings in nurseries drop in late summer. Spores are matured in
or in bearing orchards. Very small purple spots apothecia on fallen needles with new infection
appear on leaves, later extending to a brownish in spring. The damage is not heavy.
120 Blights

Furcaspora Glomerella

Deuteromycetes, Coelomycetes ▶ Anthracnose.

Glomerella cingulata Cyclamen Leaf and Bud
Starlike botryoblastospores; acervuli become Blight Leaf and Shoot Blight of poplar.
erumpent at maturity and grade into sporodochia
and pycnidia.
Furcaspora pinicola Needle Cast of pine. Gnomonia

▶ Anthracnose.
Gnomonia rubi Cane Blight of blackberry,
Fusarium dewberry, raspberry.

▶ Rots.
Fusarium graminearum Head Blight of wild Hadrotrichum
Fusarium moniliforme var. subgluti- Deuteromycetes, Hyphomycetes
nans Blight of slash pine and loblolly pine
seedlings. Sporodochia cushion-shaped, dark; conidio-
Fusarium solani Stem and Leaf Blight on phores dark, simple, forming a palisade and aris-
Spanish moss. ing from a stroma-like layer; conidia dark, nearly
Fusarium subglutinans Foliar Blight and Collar spherical, one-celled, borne singly; parasitic on
Rot on Chinese evergreen. Seedling Blight on leaves.
pine. Hadrotrichum globiferum Leaf Blight of
Fusarium tabacinum Stem Blight of squash lupine.
and pumpkin.


Gibberella Deuteromycetes, Hyphomycetes

Ascomycetes, Hypocreales, Mycelium light to dark; conidiophores short or

Nectriaceae long; septate, simple or branched, often protrud-
ing from stomata of host; more or less irregular or
Perithecia superficial, blue, violet, or greenish; bent, bearing conidia successively on new grow-
spores hyaline with several cells. Conidial stage ing tips; conidia dark typically with more than
in genus Fusarium with fusoid curved spores, three cells, cylindrical or ellipsoid, sometimes
several-septate. The species causing stalk rots of slightly curved or bent, ends rounded. Parasitic,
corn and producing gibberellic acid are more often causing leaf spots or blights of cereals and
important than those causing blights. grasses.
Gibberella baccata (Fusarium lateritium). Helminthosporium catenarium (Drechslera
Twig Blight of ailanthus, citrus, cotoneaster, fig, catenaria). Leaf Blight or Crown Rot on creeping
hibiscus, hornbeam, peach, and other plants in bentgrass; red leaf lesions and leaf tip dieback;
warm climates, sometimes associated with other eventually entire plant becomes blighted to
diseases. crown.
Hypoderma 121

Helminthosporium gigantea (Drechslera Herpotrichia juniperi Brown Felt Blight of

gigantea). Blight or Zonate Leaf Spot on wild conifers at high elevations; on fir, juniper, incense
rice and grasses. cedar, spruce, pine, yew when under snow. When
Helminthosporium maydis (Cochliobolus the snow melts, lower branches are seen covered
heterostrophus). Southern Corn Leaf Blight, eas- with a dense felty growth of brown to nearly
ily confused with southern corn leaf spot due to black mycelium, which kills foliage by excluding
H. carbonum. The leaf blight occurs throughout light and air as well as by invading hyphae.
the corn areas of the South and north to Illinois, Small, black perithecia are scattered over the
more important on field than on sweet corn. Gray- felt. This pathogen also found on dwarf mistletoe.
ish tan to straw-colored spots with parallel sides
unite to blight most of the leaf tissue. The fields
appear burned by fire. Resistant varieties offer the
only control.
Deuteromycetes, Hyphomycetes
Helminthosporium turcicum (see
Setosphaeria turcica). Northern Corn Leaf Blight
Conidiophores dark, simple, bearing conidia suc-
on field and sweet corn and on grasses; found
cessively on new growing tips; conidia dark, with
from Wisconsin and Minnesota to Florida but
three or more cells, cylindrical, with rough walls
more severe in states with heavy dews, abundant
(echinulate to verrucose); parasitic, causing leaf
rainfall, and warm summers, losses running from
spots, or saprophytic.
a trace to 50 %.
Heterosporium syringae Lilac Leaf Blight.
Setosphaeria turcica (formerly
A velvety, olive green bloom of spores if formed
Helminthosporium turcicum). Northern Corn
in blighted, gray-brown leaf areas, which may
Leaf Blight on field and sweet corn and on grasses;
crack and fall away. Infection is on mature leaves
found from Wisconsin and Minnesota to Florida
and the fungus is often associated with
but more severe in states with heavy dews, abun-
Cladosporium. If necessary, spray after mid-
dant rainfall, and warm summers, losses running
June with bordeaux mixture.
from a trace to 50 %. The disease starts on the
lower leaves and progresses upward. Small, ellip-
tical, dark grayish green, water-soaked spots turn Higginisia
greenish tan and enlarge to spindle-shape, 1/2 to 2
inches wide, 2 to 6 inches long. Spores developing See ▶Coccomyces under Leaf Spots.
on both leaf surfaces after rain or heavy dew give Higginisia hiemalis Cherry Leaf Blight. See
a velvety dark green appearance to the center of ▶ Blumeriella jaapi and ▶Coccomyces
the lesions. Whole leaves may be killed; entire biemalisi, Cherry Leaf Spot.
fields turn dry. The fungus winters in corn residue Higginisia kerriae Kerria Leaf and Twig Blight.
in the field and produces spores the next spring; See ▶ Blumeriella kerriae and ▶Coccomyces
these are spread by wind. kerriae under Leaf Spots.
Control Use a 3-year or longer rotation.

Ascomycetes, Rhytismatales
Ascomycetes, Dothidiales
Ascospores formed in hysterothecia (elongated
Mycelium dark, perithecia superficial; spores perithecia or apothecia) extending along ever-
with several crosswalls, olivaceous when mature. green needles; asci long-stalked; ascospores
122 Blights

one-celled hyaline, fusiform, surrounded by

a gelantinous sheath (see Fig. 1, chapter ▶ Nee- Hyponectria
dle Casts).
Hypoderma lethale (see ▶Ploidoderma Ascomycetes, Hypocreales,
lethale). Gray Leaf Blight of hard pines, from Nectriaceae
New England to the Gulf States.
Ploioderma lethale (formerly Hypoderma Perithecia bright colored, soft; innate or finally
lethale). Gray Leaf Blight of hard pines, erumpent; paraphyses lacking; spores one-celled,
from New England to the Gulf States. light-colored, oblong.
Hysterothecia are short, narrow, black; often Hyponectria buxi Leaf Blight, Leaf Cast of
seen on pitch pine. boxwood.

Deuteromycetes, Hyphomycetes
Ascomycetes, Rhytismatales
Cells reproducing by budding and germinating by
Like Hypoderma but one-celled spores are club- repetition; clamp connections as in Basidiomy-
shaped at upper end, tapering toward base (see cetes and probably anamorph species of
Fig. 1, chapter ▶ Needle Casts). Tremellales. The genus is not well understood.
Hypodermella abietis-concoloris (see ▶Lirula Itersonilia perplexans Petal Blight of chrysan-
abietis-concoloris). Fir Needle Blight on firs and themum and China aster. The fungus was isolated
southern balsam. from greenhouse chrysanthemums in Minnesota in
Hypodermella laricis Larch Needle and Shoot 1951 but apparently has been present, as a parasite
Blight. Yellow spots are formed on needles, or saprophyte, on many other plants. On pompom
which turn reddish brown but stay attached, giv- chrysanthemums the tip half of outer petals turns
ing a scorched appearance to trees. Hysterothecia brown and dries; the diseased tissue is filled with
are very small, oblong to elliptical, dull black, on broad hyphae and clamp connections. Inoculated
upper surface of needles. snapdragons show similar symptoms. Adequate
Lirula abietis-concoloris (formerly greenhouse ventilation seems to prevent trouble.
Hypodermella abietis-concoloris). Fir Needle This fungus has also been reported on dill.
Blight on firs and southern balsam. Itersonilia sp. Leaf Blight, Canker of parsnip,
seasonal in New York and neighboring states.
Plants are defoliated in cool, moist weather.
Spores from leaves produce a chocolate brown
Nectria (Hypomyces) dry rot on shoulder or crown of the root. Good
drainage and long rotation aid in control.
Ascomycetes, Hypocreales

Perithecia bright colored with a subicle (crustlike Kellermannia

mycelial growth underneath); spores two-celled,
light, with a short projection at one end. Deuteromycetes, Coelomycetes
Hypomyces ipomoeae (see ▶Nectria
ipomoeae). Twig Blight of bladdernut. Pycnidia black, globose, separate; immersed in
Nectria ipomoeae (formerly Hypomyces host tissue; ostiolate; conidiophores short, sim-
ipomoeae). Twig Blight of bladdernut. ple; conidia hyaline, mostly two-celled,
Micropeltis 123

cylindrical with an awl-shaped appendage at the Control Sanitation is very important; cut out and
tip; parasitic or saprophytic. burn all diseased canes. A control program for
Kellermania anomala (K. yuccaegena.) Yucca spur blight should suffice for cane blight.
Leaf Blight, general on nonarborescent forms of Leptosphaeria (Melanomma) conithyrium
yucca; in Florida and California on arborescent (Coniothyrium fuckelii) (see ▶Diapleella
forms. coniothyrium) Raspberry Cane Blight, general
Kellermania sisyrinchii (see ▶Scolecosporiella on raspberry, dewberry, blackberry.
sisyrinchii.) Leaf Blight of blue-eyed grass. Leptosphaeria korrae Blight on turfgrass
Scolecosporiella sisyrinchii (formerly (associated with Fusarium blight syndrome); dis-
Kellermannia sisyrinchii.) Leaf Blight of blue- ease is also called spring dead spot.
eyed grass. Leptosphaeria thomasiana Cane Blight of
dewberry, raspberry, in Pacific Northwest.
Leptosphaeria sp. Blight; of Miscanthus.

Deuteromycetes, Coelomycetes
Pycnidia with a radiate shield, rounded; innate or
▶ Leaf Spots.
erumpent; spores hyaline, one-celled.
Linospora tetraspora Leaf Blight of poplar.
Labrella aspidistrae Leaf Blight of aspidistra.

Leptosphaeria Lophodermella

Ascomycetes, Dothideales Ascomycetes, Rhytismatales

Perithecia membranous, not beaked, opening Hymenium on a fleshy-gelatinous stroma under

with an ostiole; innate or finally erumpent; the bark of woody plants; ascospores aseptate.
paraphyses present; spores dark, with several Lophodermella sp. Needle Cast of pine.
cells. Anamorph state a Coniothryium with
black, globose pycnidia and very small, dark,
one-celled conidia, extruded in a black cirrhus. Macrophomina
Diapleella coniothyrium (formerly
Leptosphaeria (Melanomma) conithyrium; ▶ Rots.
(Coniothyrium fuckelii)). Raspberry Cane Blight, Macrophomina phaseolina Ashy Stem Blight,
general on raspberry, dewberry, blackberry. The Charcoal Rot of soybeans, sweet potatoes, many
same fungus causes cankers of apple and rose other plants. ▶ Rots.
(▶ Cankers and Diebacks). On raspberry, brown
dead areas extend into wood; whole canes or
single branches wilt and die; often between Micropeltis
blossoming and fruiting. The fungus enters the
bark at any time during the season, through an Ascomycetes, Dothideales
insect wound or mechanical injury. Smutty
patches on the bark come from small olive A single hymenium, fruiting layer, covered with
conidia of the Coniothyrium stage and larger, an open, reticulate scutellum; paraphyses pre-
dark, four-celled ascospores. Ascospores are sent; spores hyaline, with several cells.
spread by rain; conidia by rain and insects Micropeltis viburni Leaf Blight of viburnum.
124 Blights

cucumber. Gray to brown dead areas in leaves

Monilinia (Sclerotinia) are marked with black pycnidia; leaves may turn
yellow and shrivel. Stem infection starts with
Ascomycetes, Helotiales, a water-soaked oily green area at nodes. The
Sclerotiniaceae stem is girdled, covered with a dark exuded
gum, and the vine wilts back to that point. Fruit
Stroma is a sclerotium formed in fruit by the rot starts gray, darkens to nearly jet black, with
fungus digesting fleshy tissues and replacing gummy exudate.
them with a layer of broad, thick-walled, inter- Control Clean up crop refuse; practice rotation.
woven hyphae forming a hollow sphere enclosing Some varieties are more resistant than others.
core or seed of fruit, which has become a dark, Mycosphaerella fijiensis Black Sigatoka on
wrinkled, hard mummy. Apothecia funnel-form dwarf banana in FL.
or cupulate, rarely flat-expanded, some shade of Mycosphaerella melonis Gummy Stem Blight
brown; asci eight-spored; ascospores one-celled, of cucumbers.
ellipsoidal, often slightly flattened on one side, Mycosphaerella pinodes Pea Blight.
hyaline. Conidia hyaline, one-celled, formed in ▶ Ascochyta pinodes.
chains in grayish masses called sporodochia. Mycosphaerella rabiei (Anamorph, Phoma
Monilinia azaleae Shoot Blight of native or rabiei). Blight of chickpea
pinxter azalea (Rhododendron roseum). Mycosphaerella sequoiae Needle Blight of
Apothecia are formed on overwintered mummied redwood.
fruits (capsules) in leaf mold under shrubs in
moist places. Ascospores infect leaves and suc-
culent shoots when the azalea is in full bloom. Myriogenospora
The conidial stage is common on young develop-
ing fruits in late June and July (New York). Ascomycetes, Hypocreales
Monilinia fructicola Leaf Blight and Shoot
Blight of peach. Ascomata superficial or in a stroma, fleshy,
Monilinia johnsonii Leaf Blight, Fruit Rot of bright-colored; ascus with a thick cap traversed
hawthorn. by a slender pore; ascospores filifrom,
Monilinia laxa Blossom Blight, Brown Rot of multiseptate, often fragmenting.
apricot, almond, cherry, plum, and prune on Myriogenospora atramentosa Blight on turf
Pacific Coast. Blossoms and twigs are blighted grass, centipedegrass.
with a good deal of gum formation. Monilinia
laxa is sometimes coincident with, and confused
with, M. fructicola, which causes a more general Mystrosporium
rot of stone fruits. Both are discussed more fully
under Rots. Deuteromycetes, Hyphomycetes
Monilinia rhododendri (Sclerotinia seaveri).
Twig Blight, Seedling Blight of sweet cherry. Conidia dark, muriform; hyphae long.
Bipolaris iridis (formerly Mystrosporium
adustum). Leaf Blight, Ink Spot of bulbous iris;
Mycosphaerella also on montbretia and lachenalia. Irregular black
patches or blotches appear soon after leaves push
▶ Anthracnose. through the ground; under moist conditions the
Mycosphaerella citrullina (M. melonis) conid- foliage withers and dies prematurely. Inky black
ial stage Didymella bryoniae. Gummy Stem stains appear on husks of bulbs (usually Iris
Blight, Stem End Rot, Leaf Spot of watermelon, reticulata), and yellow dots or elongated sunken
muskmelon, summer squash, pumpkin, and black craters show on fleshy scales. The bulbs
Ovulinia 125

may rot, leaving only the husk and a mass of large, obovoid, one-celled except for basal
black powder. The fungus spreads through the appendage or disjunctor cell; borne singly at
soil, invading adjacent healthy bulbs. tips of short branches of mycelium forming
Control Dig bulbs every year; discard all dis- a mat over surface of petal tissue (see Fig. 1).
eased bulbs and debris; plant in a new location. Ovulinia azaleae Azalea Flower Spot, Petal
Spray with bordeaux mixture. Blight, very destructive to southern azaleas in
Mystrosporium adustum (see ▶ Bipolaris humid coastal regions, occasional on mountain-
iridis). Leaf Blight, Ink Spot of bulbous iris; laurel and rhododendron. Starting as a sudden
also on montbretia and lachenalia. outbreak near Charleston, South Carolina, 1931,
the disease spread rapidly north of Wilmington,
North Carolina, down the coast to Florida, and
Myxosporium around the Gulf. It reached Texas by 1938 and
was in California by 1940; it was reported in
Deuteromycetes, Coelomycetes Maryland in 1945, in Virginia in 1947, and in
Philadelphia in 1959. Petal blight was reported
Conidia hyaline, one-celled, in discoid to from a Long Island, New York, greenhouse in
pulvinate acervuli on branches. 1956, apparently present there since 1952, and in
Myxosporium diedickei Twig Blight of 1959 infected all the azaleas in one New Jersey
mulberry. greenhouse. In both cases the blight started on
Myxosporium everhartii Twig Blight of dog- plants purchased from the South. This is the most
wood. M. nitidum. Twig blight and dieback of spectacular disease that I have ever witnessed,
native dogwood. Prune twigs back to sound with most of the bloom on all the azaleas in
wood; feed and water trees. a town blighting simultaneously and seemingly
overnight under special weather conditions. The
blight does not injure stem or foliage; it is con-
Neopeckia fined to the flowers. The loss is aesthetic and
economic from the standpoint of tourist trade.
Ascomycetes, Dothideales For many years, before a control program was
worked out, the great azalea gardens of the South
Perithecia hairy, not beaked, formed on had to close their gates to visitors far too early in
a mycelial mass; paraphyses present; spores the season.
two-celled, dark. Primary infection comes from very small
Neopeckia coulteri Brown Felt Blight on pines apothecia produced from sclerotia on the ground
only, otherwise similar to brown felt blight under shrubs, usually in January or February,
caused by Herpotrichia, a disease of high alti- occasionally as early as December. Spores shot
tudes on foliage under snow. into the air are carried by wind drift to flowers
near the ground of early varieties, initial spots
being whitish. If you put your finger on such
Ovulinia a spot, the tissue melts away. With continued
high humidity, heavy fog, dew, or rain, conidia
Ascomycetes, Helotiales, are produced over the inner surfaces of petals and
Sclerotiniaceae are widely disseminated to other petals by wind,
insects, and splashed rain. Within a few hours
Stroma a sclerotium, thin, circular to oval, shal- colored petals are peppered with small white
lowly cupulate, formed in petal tissue but falling spots, and white flowers have numerous brown
away; minute globose spermatia; apothecia of spots. By the next day flowers have collapsed into
Sclerotinia type, small; asci eight-spored; a slimy mush, bushes looking as if they had had
paraphyses septate with swollen tips; conidia scalding water poured over them. If the weather
126 Blights

stays wet, small black sclerotia are formed in the

petals in another 2 or 3 days. Infected blooms Pellicularia
seldom drop normally but remain hanging on
the bushes in an unsightly condition for weeks Basidiomycetes, Aphyllophorales
and months, some even to the next season. Many
of the sclerotia, however, drop out and remain in Includes some species formerly assigned to
the litter on the ground ready to send up apothecia Corticium, Hypochnus, and Peniophora. Hyphae
the next winter. stout, very short-celled; mycelium branching at
Both Indian and Kurume varieties are right angles; basidia very stout, formed on
attacked, the peak of infection coming with a resupinate, cottony or membranous layer of
mid-season varieties such as Pride of Mobile or mycelium. Anamorph state a Rhizoctonia, with
Formosa. In some seasons dry weather during sclerotia made up of brown, thin-walled, rather
early spring allows a good showing of azaleas; angular cells, or Sclerotium, with sclerotia having
in other years blight starts early and there is little a definite brown rind and light interior.
color unless azaleas are sprayed. On Belgian aza- Athelia rolfsii (formerly Pellicularia rolfsii
leas in greenhouses blight may start in December. (Anamorph, Sclerotium rolfsii). Southern Blight,
Control Some mulches and soil treatments will Crown Rot. The disease has been known, in its
inhibit apothecial production. Secondary infec- sclerotium stage, for many years on hundreds of
tion is bound to come from some untreated azalea plants. The connection with Pellicularia is recent,
in the neighborhood. Spraying gives very effec- and the name does not have universal agreement.
tive, even spectacular, control if started on time, One strain of the fungus has been called Sclero-
when early varieties are in bloom and midseason tium delphinii in the North, where the disease is
azaleas are showing color. Sprays must be usually designated crown rot. This is, however,
repeated three times a week as long as petal a variable fungus with single spore cultures from
surface is expanding, about 3 or 4 weeks. After the Pellicularia stage producing sclerotia typical
that, weekly spraying is sufficient. Spraying is of Sclerotium delphinii and of S. rolfsii, with inter-
mandatory now for the big azalea gardens, and mediate forms. Sclerotia of the southern blight
the admission fees from the lengthened season strain are very small, round, tan, about the size,
pay for the program many times over. shape, and color of mustard seed, the pathogen
The original successful formula was: Dithane being frequently called the mustard-seed fungus.
D-14 (nabam) 1 1/3 quarts to 100 gallons water, Southern blight affects almost all plants
plus 1 pound 25 % zinc sulfate, 1/2 pound except field crops like wheat, oats, corn, and
hydrated lime, and 1 ounce of spreader Triton sorghum. Fruits and vegetables include Jerusa-
B 1956. Later work showed that the lime could lem artichoke, avocado, bean, beet, carrot, cab-
be omitted, Dithane reduced to 1 quart, and zinc bage, cucumber, eggplant, endive, lettuce, melon,
sulfate to 2/3 pound to prevent injury in periods of okra, onion, garlic and shallot, pea, peanut, pep-
drought. The spray should be a fine mist, applied per, potato, rhubarb, strawberry, sweet potato,
from several directions to get adequate coverage. tomato, turnip, and watermelon. Ornamentals,
Commercial growers should beware of order- too numerous to list in entirety, include ajuga,
ing azaleas from the South unless they are bare- ageratum, amaryllis, azalea, caladium, calendula,
rooted and all flower buds showing color campanula, canna, carnation, cosmos, China
removed. As a matter of fact, any potted or balled aster, chrysanthemum, dahlia, delphinium,
and burlapped plant grown in a nursery near daphne, duranta, gladiolus, hollyhock, hydran-
azaleas could very easily bring along some of gea, iris, jasmine, lemon verbena, lily, lupine,
the tiny sclerotia in the soil, and they might marigold, morning-glory, myrtle, narcissus,
remain viable more than 1 year. All traces of orchids, phlox, pittosporum, rose, rose-mallow,
soil should be washed off roots, and the plants rudbeckia, scabiosa, sedum, sweet pea, star-of-
wrapped in polyethylene for shipping. bethlehem, tulip, violet, and zinnia.
Pellicularia 127

The first sign of blight is the formation of stems, pods, and foliage. Infection starts with
white wefts of mycelium at the base of the stem, small circular spots that appear water-soaked or
spreading up in somewhat fan-shaped fashion scalded. They enlarge to an inch or more, become
and sometimes spreading out over the ground in tan with a darker border, are sometimes zonate.
wet weather. The sclerotia formed in the wefts are The whitish mycelium grows rapidly over the leaf
first white, later reddish tan or light brown. They blade, killing it, and spreads a web from leaf to
may be numerous enough to form a crust over the leaf, over petioles, flowers, and fruit, in wet
soil for several inches around a stem, or they may weather and at temperatures 70 to 90  F; in dry
be somewhat sparse and scattered. weather growth is inconspicuous except on fallen
In the white stage, droplets of liquid often leaves. The fungus is spread by wind, rain, irri-
form on the sclerotia, and the oxalic acid in this gation water, cultivating tools, and bean pickers;
liquid is assumed to kill plant cells in advance of it survives in sclerotial form from season to
the fungus hyphae. This means that the pathogen season.
never has to penetrate living tissue and explains Control Destroy infected plants; clean up refuse.
why so many different kinds of plants succumb so In Florida, do not plant beans between June and
readily to southern blight. Fruits touching the September if web blight has been present. Use
ground, as well as vegetables with fleshy roots, a copper spray or dust.
like carrots and beets, or plants with bulbs or Pellicularia filamentosa f. sp. sasakii See
rhizomes, like onions, narcissus, and iris, seem ▶Thanatephonus cucumeris. Leaf Blight of
particularly subject to this disease. Low orna- grasses, clover, etc.
mentals such as ajuga blight quickly, the whole Pellicularia filamentosa f. sp. timsii See
plant turning black; tall plants like delphinium rot ▶Thanatephonus cucumeris. Leaf Blight of fig.
at the crown and then die back or topple over; Pellicularia rolfsii See ▶ Athelia rolfsii,
bulbs have a cheesy interior, with sclerotia Anamorph, ▶Sclerotium rolfsii. Southern Blight,
forming on or between the scales. Crown Rot. The disease has been known, in its
Control Remove diseased plants as soon as they sclerotium stage, for many years on hundreds of
are noticed. Take out surrounding soil, for 6 plants. The connection with Pellicularia is
inches beyond the diseased area, wrapping it recent, and the name does not have universal
carefully so that none of the sclerotia drop back. agreement. One strain of the fungus has been
Increasing the organic content of the soil reduces called Sclerotium delphinii in the North, where
southern blight, as does the addition of nitroge- the disease is usually designated crown rot. This
nous fertilizers, such as ammonium nitrate. is, however, a variable fungus with single spore
Treating narcissus bulbs in hot water for 3 h, as cultures from the Pellicularia stage producing
for nematodes, kills the fungus in all except the sclerotia typical of Sclerotium delphinii and of
very largest bulbs. S. rolfsii, with intermediate forms. Sclerotia of
Pellicularia filamentosa (see ▶Thanatephorus the southern blight strain are very small, round,
cucumeris), teleomorph state of Rhizoctonia tan, about the size, shape, and color of mustard
solani. This is a variable fungus with some strains seed, the pathogen being frequently called the
or forms causing leaf blights but best known as mustard-seed fungus.
cause of Rhizoctonia rot of potatoes and Southern blight affects almost all plants
damping-off of many plants. ▶ Rots. except field crops like wheat, oats, corn, and
Pellicularia filamentosa f. sp. sorghum. Fruits and vegetables include Jerusa-
microsclerotia (Corticium microsclerotia) see lem artichoke, avocado, bean, beet, carrot, cab-
▶Thanatephorus cucumeris. Web Blight of snap bage, cucumber, eggplant, endive, lettuce, melon,
bean, lima bean, also reported on fig, elder, hibis- okra, onion, garlic and shallot, pea, peanut, pep-
cus, hollyhock, tung oil, and phoenix tree, from per, potato, rhubarb, strawberry, sweet potato,
Florida to Texas. Many small brown sclerotia and tomato, turnip, and watermelon. Ornamentals,
abundant weblike mycelium are found on bean too numerous to list in entirety, include ajuga,
128 Blights

ageratum, amaryllis, azalea, caladium, calendula, roses, and other ornamentals, and some fruits.
campanula, canna, carnation, cosmos, China The disease is recorded on apple, azalea, banana
aster, chrysanthemum, dahlia, delphinium, shrub, blackberry, boxwood, camphor, cherry
daphne, duranta, gladiolus, hollyhock, hydran- laurel, chinaberry, columbine, crabapple, crape
gea, iris, jasmine, lemon verbena, lily, lupine, myrtle, casuarina, currant, dewberry, dogwood,
marigold, morning-glory, myrtle, narcissus, elderberry, elm, erythrina, euonymus, fig,
orchids, phlox, pittosporum, rose, rose-mallow, flowering almond, flowering quince, goldenrod,
rudbeckia, scabiosa, sedum, sweet pea, star-of- gooseberry, guava, honeysuckle, hibiscus, morn-
bethlehem, tulip, violet, and zinnia. ing glory, pear, pecan, pepper vine, persimmon,
The first sign of blight is the formation of pittosporum, plum, pomegranate, quince, rose,
white wefts of mycelium at the base of the stem, satsuma orange, soapberry, silver maple, sweet
spreading up in somewhat fan-shaped fashion potato, tievine (Jacquemontia), tung, Virginia
and sometimes spreading out over the ground in creeper, and viburnum.
wet weather. The sclerotia formed in the wefts are The fungus winters as sclerotia on twigs and
first white, later reddish tan or light brown. They leaf petioles, and in May and June produces
may be numerous enough to form a crust over the threadlike mycelium that grows over lower sur-
soil for several inches around a stem, or they may face of leaves, killing them and causing prema-
be somewhat sparse and scattered. ture defoliation, although often dead leaves hang
In the white stage, droplets of liquid often on the tree in groups, matted together by thread-
form on the sclerotia, and the oxalic acid in this like spider webs. Fruiting patches on leaves are
liquid is assumed to kill plant cells in advance of first white, then buff. The fungus flourishes in
the fungus hyphae. This means that the pathogen moist weather, temperatures 75 to 90  F.
never has to penetrate living tissue and explains Control On figs, one or two applications of tri-
why so many different kinds of plants succumb so basic copper sulfate, or bordeaux mixture, are
readily to southern blight. Fruits touching the satisfactory until the fruit ripens in July. Pruning
ground, as well as vegetables with fleshy roots, out infected branches may be sufficient on tung
like carrots and beets, or plants with bulbs or and pecan, but at least one spray of bordeaux
rhizomes, like onions, narcissus, and iris, seem mixture may be required.
particularly subject to this disease. Low orna- Thanatephonus cucumeris (formerly
mentals such as ajuga blight quickly, the whole Pellicularia filamentosa), teleomorph state of
plant turning black; tall plants like delphinium rot Rhizoctonia solani. This is a variable fungus
at the crown and then die back or topple over; with some strains or forms causing leaf
bulbs have a cheesy interior, with sclerotia blights but best known as cause of Rhizoctonia
forming on or between the scales. rot of potatoes and damping-off of many plants.
Control Remove diseased plants as soon as they ▶ Rots.
are noticed. Take out surrounding soil, for Thanatephonus cucumeris (formerly
6 inches beyond the diseased area, wrapping it Pellicularia filamentosa f. sp. micro-sclerotia
carefully so that none of the sclerotia drop back. (Corticium micro sclerotia)). Web Blight of
Increasing the organic content of the soil reduces snap bean, lima bean, also reported on fig, elder,
southern blight, as does the addition of nitroge- hibiscus, hollyhock, tung oil, and phoenix tree,
nous fertilizers, such as ammonium nitrate. from Florida to Texas. Many small brown scle-
Treating narcissus bulbs in hot water for 3 h, as rotia and abundant weblike mycelium are found
for nematodes, kills the fungus in all except the on bean stems, pods, and foliage. Infection starts
very largest bulbs. with small circular spots that appear water-
Pellicularia koleroga (Corticium stevensii). soaked or scalded. They enlarge to an inch or
Thread Blight, a southern disease, from North more, become tan with a darker border, are some-
Carolina to Texas, important on fig and tung, times zonate. The whitish mycelium grows rap-
sometimes defoliating pittosporum, crape myrtle, idly over the leaf blade, killing it, and spreads
Phacidium 129

a web from leaf to leaf, over petioles, flowers, and temperature than the fungus; shading transplants
fruit, in wet weather and at temperatures 70 to is helpful.
90  F; in dry weather growth is inconspicuous Pestalotia sp. and Penicillium sp. Flower Blight
except on fallen leaves. The fungus is spread by on camellia.
wind, rain, irrigation water, cultivating tools, and Pestalotiopsis funerea (formerly Pestalotia
bean pickers; it survives in sclerotial form from funerea). Tip Blight of conifers, Needle Blight,
season to season. Twig Blight of chamaecyparis, retinospora,
Control Destroy infected plants; clean up refuse. cypress, bald cypress, arborvitae, juniper, yew,
In Florida, do not plant beans between June and and giant sequoia. The fungus is saprophytic on
September if web blight has been present. Use dead and dying tissue and also weakly parasitic,
a copper spray or dust. infecting living tissue through wounds under
Thanatephonus cucumeris (formerly moist conditions. It appears in sooty pustules on
Pellicularia filamentosa f. sp. sasakii). Leaf leaves, bark, and cones.
Blight of grasses, clover, etc.
Thanatephorus cucumeris (formerly
Pellicularia filamentosa f. sp. timsii). Leaf Blight Phacidium
of fig.
Ascomycetes, Helotiales

Apothecia innate, concrete above with the epi-

dermis and slitting with it into lobes; spores one-
celled, hyaline.
▶ Cankers and Diebacks.
Phacidium abietinellum (see ▶Nothophacidium
Penicillium oxalicum Leaf Blight of grass.
abietinellum). Needle Blight of balsam fir.
Phacidium balsameae (see ▶Sarcotrochilia
balsameae). Needle Blight of balsam fir in New
Pestalotia England, of white and alpine fir in the Northwest.
Phacidium infestans Snow Blight of conifer
Deuteromycetes, Coelomycetes seedlings on fir and young pines in the Northeast,
also on arborvitae and spruce; on white and
Acervuli dark, discoid or cushion-shaped, subcu- alpine fir in the Northwest. This native fungus is
taneous; conidiophores short, simple; conidia most damaging in nurseries, attacking foliage
fusiform, several-celled with median cells col- under the snow. The needles turn brown, with
ored, end cells hyaline, a short stalk at the basal a covering of white mycelium, just as the snow
cells and a crest of two or more hyaline append- melts. In late summer and fall brown to nearly
ages, setae, from the apical cell (Fig. 1). Weak black apothecia appear on underside of browned
parasites or saprophytes; some are treated under needles. Ascospores are spread by wind, primary
Leaf Spots. infection being in autumn. Additional infection
Pestalotia funerea (see ▶Pestalotiopsis occurs in late winter, when mycelium grows out
funerea). Tip Blight of conifers, Needle Blight, under the snow from diseased to dormant, healthy
Twig Blight of chamaecyparis, retinospora, needles.
cypress, bald cypress, arborvitae, juniper, yew, Control Spray nursery beds with dormant-
and giant sequoia. strength lime sulfur in late fall; remove infected
Pestalotia hartigii Associated with a basal stem seedlings; dip new stock in lime sulfur before
girdle of young conifers but parasitism not planting.
proven. The stem has a swelling above the gir- Nothophacidium abietinellum (formerly
dling lesions, and the tree gradually turns yellow Phacidium abietinellum). Needle Blight of
and dies. The effect may be more from high balsam fir.
130 Blights

Sarcotrochilia balsameae (formerly Phoma piceina Twig and Needle Blight of Nor-
Phacidium balsameae). Needle Blight of balsam way spruce. May cause defoliation and some-
fir in New England, of white and alpine fir in the times death of forest trees.
Northwest. Phoma sclerotioides Brown Root Rot of alfalfa.
Phoma strobiligena (see ▶Sclerophoma
pythiophila), on cone scales of Norway spruce.
Phaeoacremonium Sclerophoma pythiophila (formerly
Phoma strobiligena), on cone scales of Norway
Phaeoacremonium chlamydosporum Black spruce.
Goo on grape.

Phialophora Phomopsis

▶ Rots. Deuteromycetes, Coelomycetes

Phialophora graminicola Blight on turfgrasses
(associated with Fusarium blight syndrome). Pycnidia dark, ostiolate, immersed, erumpent,
nearly globose; conidiophores simple; conidia
hyaline, one-celled, of two types-ovate or ellip-
Phloeospora soidal and long, filamentous, sickle-shaped or
hooked at upper end (Fig. 1). Anamorph state of
Deuteromycetes, Coelomycetes Diaporthe; parasitic causing spots on various
plant parts.
Pycnidia dark, imperfectly formed, globose, Dendrophoma obscurans (see ▶Phomopsis
innate in tissue, not in distinct spots; conidia obscurans). Strawberry Leaf Blight, Angular
hyaline or subhyaline, several-celled, elongate Leaf Spot.
fusoid to filiform; parasitic or saprophytic. One Phomopsis ambigua (teleomorph, Diaporthe
of the conidial forms linked with Mycosphaerella eres). Twig Blight of pear, widespread.
as a teleomorph state. Phomopsis diospyri Twig Blight of native
Phloeospora adusta Leaf Blight of clematis. persimmon.
Phomopsis japonica Twig Blight of kerria.
Phomopsis juniperovora Nursery Blight, Juni-
Phoma per Blight, Cedar Blight, Canker on red-cedar
and other junipers, cypress, chamaecyparis, Jap-
▶ Blackleg. anese yew (Cephalotaxus), arborvitae, giant
Phoma conidiogena (see ▶Phoma glomerata). sequoia, and redwood. This disease occurs in
Boxwood Tip Blight. Ashy gray necrotic areas at virulent form from New England to Florida and
leaf tips, with pycnidia on both leaf surfaces. through the Middle West; it may also occur on the
Phoma glomerata (formerly Phoma Pacific Coast.
conidiogena). Boxwood Tip Blight. Tips of branches turn brown with progressive
Ashy gray necrotic areas at leaf tips, with dying back until a whole branch or even a young
pycnidia on both leaf surfaces. tree is killed. Trees over 5 years old are
Phoma fumosa Twig Blight, occasional on less seriously injured. Spores produced in quan-
maple. tity in pycnidia on diseased twigs ooze out in little
Phoma macdonaldii Blight, Premature Ripen- tendrils in moist weather, to be spread by
ing of sunflower. splashing water, insects, and workers. Entrance
Phoma mariae Twig Blight on Japanese is through unbroken tissue as well as wounds; the
honeysuckle. stem is killed above and below the point of
Physalospora 131

entrance. Small, sunken lesions give a flattened

appearance to some seedlings. Overhead irriga- Phyllosticta
tion in a nursery is a predisposing factor, and
a large amount of stock can be blighted in Deuteromycetes, Coelomycetes
a very short time. Older trees in home
plantings suffer from twig blight. The fungus Pycnidia dark, with ostiole, in spots in leaves;
winters on infected plant parts and remains viable spores one-celled, hyaline. The characteristics
at least 2 years. are the same as Phoma except that leaves rather
Control Have seedbeds well drained; water by than stems are infected. Other species are listed
ditch irrigation; remove and burn diseased seed- under Leaf Spots.
lings early in the season; keep seedbeds away Phyllosticta batatas Sweet Potato Leaf Blight,
from older cedar trees; do not use cedar branches occasional from New Jersey to Florida, more
or needles for mulching. Spray with fixed prevalent in the South but seldom important
copper or bordeaux mixture plus a wetting enough for control measures. Numerous white
agent, starting when growth begins and repeating spots on leaves are bordered with narrow reddish
to keep new foliage covered. Spiny Greek and zones; pycnidia are numerous; spores are
Hill junipers and Keteller red-cedars are some- extruded in tendrils.
what resistant. Phyllosticta cryptomeriae Needle Blight
Phomopsis kalmiae Mountain-Laurel Leaf found on Cryptomeria.
Blight, Blotch. Circular, brown, often Phyllosticta lagerstroemiae Tip Blight of
zonate areas on leaves, frequently starting near crape-myrtle.
margin or tip, gradually enlarge and coalesce Phyllosticta multicorniculata Needle Blight of
until most of the blade is involved. The fir.
fungus often works down the petiole to Phyllosticta pteridis Tip Blight of fern. Leaves
cause a twig blight. The disease is more lose green color; spots are ash gray with purple
prominent on bushes in the shade or under drip brown margins and numerous black pycnidia in
of trees. Remove blighted leaves or clean up center. A very weak bordeaux mixture has been
fallen leaves. suggested for control; if overhead watering is
Phomopsis longicolla Black Pod Spot and Seed avoided, spraying may not be necessary.
on cowpea.
Phomopsis oblonga Twig Blight on Chinese
elm. Physalospora
Phomopsis obscurans (formerly Dendrophoma
obscurans). Strawberry Leaf Blight, Angular Ascomycetes, Sphaeriales
Leaf Spot. The lesions are large, circular to angu-
lar, reddish purple, zonate with age, having a dark Perithecia with papillate mouths, immerse in sub-
brown center, a light brown zone, and a purple stratum but without well-defined stromata;
border. Spots may extend in a V-shaped area paraphyses present; spores one-celled, hyaline.
from a large vein to edge of the leaf, with black A few species cause blights; many cause rots.
fruiting bodies appearing in the central portion. Botryosphaeria obtusa (formerly
Not serious before midsummer, the disease may Physalospora obtusa). Cane Blight of rose, also
be destructive late in the season. The fungus Black Rot of apple, Canker and Dieback of many
winters on old leaves. plants. ▶ Cankers and Diebacks and also ▶ Rots.
Phomopsis occulta Shoot Blight of Colorado Glomerella cingulata (formerly Physalospora
blue spruce. dracaenae). Dracaena Tip Blight, Leaf Spot. Dis-
Phomopsis vexans Phomopsis Blight of egg- ease starts at the tips of lower leaves and spreads
plant. ▶Diaporthe vexans. down toward the base. Infected areas are sunken
Phomopsis vaccinii Twig Blight of blueberry. and straw-colored, dotted with black specks of
132 Blights

pycnidia. All leaves on the plant may die except first, later large bleached or scalded areas. Dark,
a few at the top. Remove infected leaves as soon watersoaked patches on fruits are covered with
as noticed. Spray with a copper fungicide. white mycelium. The fruit withers but remains
Physalospora dracaenae (▶Glomerella attached; 60 % of green fruit may be infected in
cingulata). Dracaena Tip Blight, Leaf Spot. southwestern commercial plantings. Seed are
Physalospora gregaria Twig Blight of yew. infected from the fruit. Symptoms on squash are
Physalospora obtusa (see ▶ Botryosphaeria somewhat similar; green leaf lesions spreading
obtusa). Cane Blight of rose, also Black Rot of over the blade, a basal stem rot, and wilting. Wet
apple, Canker and Dieback of many plants. soil and high temperatures encourage blight.
Control Place seedbeds on land that has not pre-
viously grown peppers; rotate crops. Avoid over
Phytophthora irrigation.
Phytophthora citrophthora (also P. citricola
Oomycetes, Peronosporales and P. nicotianae var. parasitica). Shoot and
Stem Blight on azalea. Needle Blight and Branch
This most important genus contains many species Dieback of sequoia.
causing destructive blights, cankers, and rots. Phytophthora erythroseptica Leaf Blight of
The name, which means “plant destroyer,” was pink and golden calla. Leaves are wilted and
given in 1876 for the potato blight fungus. distorted; petioles are black and soft.
Sporangia, formed successively on sporangio- Phytophthora ilicis Holly Blight,
phores, slender, sparsely branched hyphae Phytophthora Leaf and Twig Blight, the most
emerging from stomata, germinate either by serious disease of English holly, particularly seri-
a germ tube or by zoospores. The sexual spore ous in the Northwest. For many years the trouble
is an oospore. was ascribed to Boydia insculpta and called
Phytophthora cactorum Lilac Shoot Blight. Boydia canker, but this fungus merely invades
Blossoms and succulent growing tips are blighted tissue killed by Phytophthora. Leaf spots are
and turn brown; suckers are killed back 4 or 5 dark, developing on lower leaves in cool rainy
feet. Blight is most severe in wet springs when weather and progressing upward in late fall and
shrubs are crowded, shaded, and improperly winter. Young twigs die back; black stem cankers
pruned. The same fungus causes a canker, foot kill older twigs. Young plants in nurseries are
rot, and dieback of rhododendron and other plants defoliated and sometimes killed.
and is considered again under Cankers. Avoid Control Choose a planting site with moderate air
planting lilacs and rhododendrons close together. movement; space trees well apart. Prune out all
Prune each year for air circulation and to remove cankered and blighted twigs; prune also for air
dead twigs. movement through trees. Spray with tribasic cop-
Phytophthora capsici Phytophthora Blight of per sulfate, starting the middle of October.
Pepper, Leaf and Stem Blight of Squash, Fruit Phytophthora infestans Late Blight of potato
Rot of pepper, eggplant, tomato, cucumber, and and tomato, general on potato in the Northeast, in
melon. The disease was first found in New Mex- Middle Atlantic and North Central states, some-
ico in 1918 injuring chili peppers; it occurs times in Gulf and western states; on tomato in
chiefly in southwestern and Gulf states. In 1953, humid regions and seasons. Here is a pathogen
however, it was reported that for some years it that has not lost its destructive virulence with pas-
had been causing a leaf blight of squash in North sage of time. In 1946, a whole century after potato
Carolina. blight caused the famous Irish famine, tomato
Pepper plants are girdled at the soil line with blight devastated tomatoes along the eastern sea-
a dark green water-soaked band, which dries and board, both in home gardens and canning fields.
turns brown, followed by wilting and death of the The potato went to Europe from South Amer-
entire plant. Leaf spots are dark green and small at ica shortly before 1600, seemingly leaving its
Late Blight of Potato 133

pathogens at home. For 200 years potatoes

thrived in Europe as the main source of carbohy-
drate food, but in August 1845, the Gardener’s
Chronicle reported: “A fatal malady has broken
out amongst the potato crop. On all sides we hear
of destruction. In Belgium the fields are said to
have been completely desolated. There is hardly
a sound sample in Covent Garden Market.”The
editor went on to describe the decay and to say:
“As to cure for this distemper there is none. One
of our correspondents is today angry with us for
not telling the public how to stop it; but he ought
to consider that Man has no power to arrest the Fig. 4 Late Blight on Potato
dispensations of Providence. We are visited by
a great calamity which we must bear.” And in weather, first on lower leaves. As a spot enlarges
1946 American gardeners were again blaming the the center is shriveled, dry, dark brown to black,
editor, for lack of information on tomato blight. and a downy, whitish growth appears on the
In 1845 the weather was continued gloom and underside of leaves. Similar lesions are formed
fog, with below-average temperatures. The Gar- on stems and petioles, and there is a characteristic
dener’s Chronicle editor was sure blight was due strong odor as tops are blighted. On tubers, first
to potatoes being overladen with water. The Rev. symptoms are small brown to purple discolor-
M. J. Berkeley disagreed. He insisted blight was ations of skin on upper side, changing to
due to a fungus, with the weather contributing to depressed pits when tubers are removed from
spread of a moisture-loving parasite. The argu- soil and put in storage (see Fig. 4). On cutting
ment raged, for this was long before Pasteur and through the potato, a reddish brown dry rot is
his germ theory, and the first time anyone seen.
believed a fungus could be the cause and not the Life History The primary cycle starts with
consequence of plant disease. A French scientist, infected tubers, which have harbored mycelium
Montagne, named the fungus Botrytis infestans, in the dry rot patches over winter. If infected seed
but the first really good description of it was pieces are planted, the fungus grows systemically
published by Berkeley, and it remained for the into the shoots and finally fruits by sending spo-
German de Bary, in 1876, actually to prove rangiophores out through the stomata on lower
the pathogenic nature of the fungus and to leaf surfaces (see Fig. 5). These swell at the tips
erect the new genus Phytophthora to include it. into ovoid bodies, sporangia, then branch and
Meanwhile the disease was making history. produce successively more sporangia. The latter
The loss of the potato crop in 1845 and 1846 may function as conidia, putting out a germ tube,
killed off a million people and caused another but more often are differentiated into a number of
million and a half to emigrate; the first Govern- swarmspores (zoospores), which have cilia
ment Relief program was instigated; and the enabling them to swim about after they are
English Corn Laws were repealed with a change splashed by rain to another leaf. Eventually they
to a policy of free trade and unbounded expansion stop swimming and send a germ tube in through
of commerce. the leaf cuticle or enter through a stoma. Initial
infection in the field also comes from conidia
blown over from sprouts produced on infected
Late Blight of Potato tubers in cull piles. Blighting follows rapidly,
with first symptoms 5 days or less from the time
Symptoms After blossoming, large, dark green, of infection and with the fungus fruiting again in
water-soaked spots appear on leaves in wet a whitish layer on the underside of leaves.
134 Blights

Fig. 5 Late Blight of Potatoes. Sporangiophores of Phytophthora infestans emerging from leaf, bearing sporangia,
sometimes called conidia, which germinate by zoospores

Tubers with only a thin covering of soil may be

infected by swarmspores washing down onto Late Blight of Tomato
them from blighted leaves overhead; they are
also infected during digging if it is done in Although there are potato and tomato strains of
moist weather while tops are still green. Phytophthora infestans, each is capable of
Swarmspores remain viable in the soil several infecting the other host. Ordinarily blight starts
weeks while awaiting favorable conditions. with potatoes in midsummer; when the fungus
Oospores, the sexual spores, are apparently not moves over to tomatoes, it has to go through
required in the life cycle for they are not found several cycles to build up a strain virulent enough
with potatoes grown in the field. They have been to produce general blighting, and by that time the
produced in culture. tomato season is nearly over. Now we know that
Weather Relations This is a disease entirely it is possible for the tomato strain to winter in
dependent on weather conditions. Temperature potato tubers and be ready to inflict damage on
and moisture conditions are right for an epiphy- tomatoes with the first crop of zoospores pro-
totic about 2 years out of 5. Zoospores are pro- duced on potato sprouts. Conversely, tomato
duced only in cool weather, 60  F and under, but seedlings brought up from the South and planted
they invade leaves most rapidly at higher temper- near potato fields can start an epiphytotic of late
atures. Because they are swimming spores, rain is blight on potatoes.
required. A cool, wet July is usually followed by The 1946 tomato blight saga – the one that
blight in August and September. awakened eastern gardeners to the fact that plant
Control Some varieties, such as Kennebec, Essex, disease could be as important to home gardeners
Pungo, and Cherokee, are resistant to the common as to farmers – started in Florida late in Novem-
strain of the fungus but not to some of the newer ber 1945. By January the disease was extremely
strains. Treat potato dumps and cull piles with destructive in tomato seedbeds, and it continued
a weed spray to control sprouts. Delay digging crop so intermittently whenever temperatures ranged
until 2 weeks after tops die, or else kill the tops with from 60 to 70  F and relative humidity was
a weed killer to prevent infection at early digging. nearly 100 % for more than 15 h. Evidence
Pyricularia 135

indicated spores could be wind-borne for as far as

30 miles. The wave of late blight went west to
Alabama, taking 75 % of the early crop, and
rolled up the Atlantic Coast, reaching the Caroli-
nas in May and Virginia and Maryland in June,
again taking 75 % of the early crop. It rolled into
Delaware and New Jersey in July, but did not
reach peak epidemic form until after an extended
rainy period in August, and ended in Massachu-
setts in August and September.
In 1947 a blight-forecasting service was
started, based on weekly graphs prepared by plot-
ting daily the cumulative rainfall and mean tem-
peratures and aided by reports from key Fig. 6 Late Blight on Tomato
pathologists in various states. If conditions are
unfavorable for blight, we can save time and
money by eliminating useless spraying. On lilacs large irregular leaf patches have
Symptoms On seedlings small, dark spots on a lighter zone at margin. There may be some
stems or leaves are followed by death within 2 defoliation.
or 3 days. On mature plants blight starts with
dark, water-soaked leaf spots and large, dark
brown spots on fruit, with most of the leaves Plectosporium
soon hanging lifeless and fruit rotting on the
ground (see Fig. 6). Plectosporium abacinum Blight on Hydrilla.
Control Bordeaux mixture applied to young
tomato plants will either prevent fruit setting or
cause stunting. It can be used after blossoming, or Pyrenochaeta
a fixed copper can be substituted.
Phytophthora meadii Blight and Leaf Spots on Deuteromycetes, Sphaeropsidales,
West Indian holly. Sphaerioidaceae
Phytophthora medicaginis Rot Root on
Medicago spp. Pycnidia dark, ostiolate, nearly globose,
Phytophthora nicotianae (formerly erumpent with a few bristles near ostiole; conid-
Phytophthora parasitica var, parasitica. Leaf, iophores simple or branched; conidia small, one-
Stem and Bub Blight on bougainvillea, dog- celled, hyaline, ovate to elongate; parasitic or
wood, hibiscus, artillery plant, and aluminum saprophytic. See also under ▶ Rots.
plant. Leaf blight; on jojoba. Pyrenochaeta phlogis Stem Blight of Phlox.
Phytophthora parasitica var. parasitica (see
▶Phytophthora nicotianae). Leaf, Stem and
Bub Blight on bougainvillea, dogwood, hibiscus, Pyricularia
artillery plant, and aluminum plant. Leaf blight;
on jojoba. Deuteromycetes, Hyphomycetes
Phytophthora syringae Citrus Blight, also on
lilac, but the more common lilac blight is due to Conidiophores long, slender, simple or rarely
P. cactorum. On citrus trees leaves have semi- branched, septate, single or in tufts; conidia pyr-
transparent spots similar to frost damage. Other iform to nearly ellipsoid, borne singly and
Phytophthora species may be present with attached at broader end; spores hyaline, two-to
P. syringae to cause brown rot of fruits. ▶ Rots. three-celled; parasitic, chiefly on grasses.
136 Blights

Pyricularia grisea Leaf Blight on creeping bent sclerotia made up of short, irregular, angular or
grass and buffelgrass. somewhat barrel-shaped cells.
Rhizoctonia ramicola Silky Thread Blight
a southern disease similar to web blight caused
Pythium by Pellicularia koleroga. Perennial ornamental
hosts in Florida include elaeagnus, erythrina,
▶ Rots. crape-myrtle, holly, guava, pittosporum, pyr-
Pythium myriotylum Blight of tomato. acantha, Carolina jessamine, feijoa, and rhodo-
dendron. Tan spots with purple-brown margins
appear on leaf blades, dead lesions on petioles
and young twigs. When leaves are abscissed, they
Delphinella (Rehmiellopsis) are often held dangling and matted together by
brown fungus threads. Infection recurs annually
Ascomycetes, Dothideales in moist weather with high daytime temperatures.
The fungus winters as mycelium in leaf lesions
Perithecia single, globose, rupturing irregularly; and diseased twigs. Sclerotia are apparently
asci in fascicles, no paraphyses; spores hyaline, lacking in this species.
two-celled. Rhizoctonia sp. (teleomorph, Aquathana-
Delphinella balsameae (formerly tephorus pendulus). Blight on water hyacinth.
Rehmiellopsis balsameae). Tip Blight, Needle Rhizoctonia sp. (teleomorph, Thanatephorus
Blight of balsam fir, on native balsam fir in north- cucumeris). Blight on beet. Needle blight on
ern New England and on ornamental firs in south- pine.
ern New England and New York. Infection is in Rhizoctonia solani Blight of pistachio and
spring with needles of current season shriveled, Cynodon spp., and Foliar Blight of soybean.
curled, and killed, often with a dieback of termi-
nal or lateral shoots and sometimes cankers at
base of infected needles. Satisfactory control on Rhizopus
ornamental firs has been obtained by three sprays,
at 10-day intervals, ofbordeaux mixture, the first ▶ Rots.
application made as new growth starts. Rhizopus stolonifer Seedling Blight on lupine;
Rehmiellopsis balsameae (see ▶Delphinella also caused by Pleiochaeta setosa, Alternaria sp.,
balsameae). Tip Blight, Needle Blight of balsam Aspergillus flavus, Aspergillus niger, and
fir, on native balsam fir in northern New England Curvularia sp.
and on ornamental firs in southern New England
and New York.

Rhizoctonia Ascomycetes, Xylariales

Deuteromycetes, Mycelia Sterilia Perithecia separate, superficial from the first, car-
(Fungi Imperfecti) bonaceous, not beaked, ostioles papillate; spores
dark, one-celled with a small groove.
Sclerotial form of some species of Pellicularia, Rosellinia herpotrichioides Hemlock Needle
Corticium, Macrophomina, and Helicobasidium. Blight. Needle-bearing portions of twigs become
Young mycelium colorless, with branches covered on underside with a grayish brown myce-
constricted at points of origin from main axis, lial mat; black perithecia are produced in this mat
but soon colored, a weft of brownish yellow to in great abundance. Ovoid, hyaline conidia are
brown strands, organized into dense groups, formed on Botrytis-like conidiophores.
Sclerotinia (Whetzelinia) 137

into flower, the ascospores infecting the perianth

Schirrhia and causing flower spotting. From withered
flowers numerous large conidia, germinating
Ascomycetes, Dothideales, with several germ tubes, infect foliage, on
Dothideaceae which large sclerotia are formed late in the sea-
son. Remove infected parts immediately; spray
Asci usually short, cylindrical, and relatively early in the season.
numerous in spherical, ostiolate locules. Ciberinia camelliae (formerly Sclerotinia
camelliae). Camellia Flower Blight, long known
in Japan, first noted in California in 1938, con-
Scleropycnium firmed in Georgia in 1948, although probably
there several years previously, reported in Ore-
Deuteromycetes, Coelomycetes gon in 1949, Louisiana and North Carolina in
1950, South Carolina in 1954. The blight is now
Pycnidia open out to a deep cupulate or discoid widespread in Virginia, confined to certain
structure, tough, dark or black, subepidermal or counties in other states. It was not officially
subcortical, then erumpent; spores hyaline, one- recorded from Texas until 1957 but must have
celled. Largely saprophytic on twigs, sometimes been there earlier. The 1950 outbreak at Shreve-
parasitic on leaves. port, Louisiana, is said to have started on plants
Scleropycnium aureum Leaf Blight of brought in from Texas that probably originated in
mesquite. California.
Floral parts only are affected, infection taking
place any time after tips of petals are visible in
Sclerotinia (Whetzelinia) opening buds. Few to many brownish specks on
expanding petals enlarge until the whole flower
Ascomycetes, Helotiales, turns brown and drops. In early stages darkened
Sclerotiniaceae veins are prominent diagnostic symptoms. When
the flowers rest on moist earth, spermatia are
Apothecia arising from a tuberoid sclerotium produced on petals in shiny black masses. Hard,
which, though formed free on aerial mycelium, dark brown to black sclerotia formed at the base
is sometimes enclosed in natural cavities of of petals frequently unite into a compound struc-
suscept or host, as in hollow stem of perennials. ture simulating petal arrangement. This com-
Interior (medulla) of sclerotium white, pound sclerotium may be an inch or more in
completely enveloped by a dark rind; gelatinous diameter. Although the petals do not melt when
matrix lacking. Conidia wanting but spermatia touched as do azaleas with petal blight, there is
(very small microconidia) formed on a distinctive moist feeling that helps to differen-
sporodochia borne free or enclosed in cavities. tiate flower blight from frost injury. Rarely,
Apothecia some shade of brown; cupulate to fun- a flower blight of camellias is caused by another
nel-form; usually at maturity saucer-shaped to Sclerotinia (S. sclerotiorum).
flat expanded; ascospores hyaline, one-celled, Sclerotia lie dormant on ground or in
ovoid. Species formerly included in Sclerotinia mulching materials until the next winter when,
but possessing monilioid conidia are now in from January on (possibly earlier), after wet
Monilinia. periods with rising temperature, they produce
Botryotinia polyblastis (formerly Sclerotinia one to several apothecia on long or short stipes
polyblastis). Narcissus Fire. A serious flower with brown, saucerlike discs 1/4 to 3/4 inch
blight in England, known here on the Pacific across, rarely up to 1 inch. Spores, discharged
Coast. In England overwintering sclerotia pro- forcibly, are carried by wind currents to flowers,
duce apothecia when Narcissus tazetta comes thus completing the cycle. Spores may be
138 Blights

wind-borne at least 1/3 mile, but presumably Sclerotinia minor Blight of soybean, peanut,
a large proportion of them land on opening petals and Eclipta.
of the bush overhead. The sclerotia remain viable Sclerotinia (Botryotinia) polyblastis (see
in the soil at least 2 or 3 years, sending up more ▶Botryotinia polyblastis). Narcissus Fire.
apothecia each season. No conidia are known; so A serious flower blight in England, known here
there is no secondary infection from flower to on the Pacific Coast.
flower as with azalea blight. The amount of pri- Sclerotinia rolfsii Southern Blight on St.
mary inoculum is very large, however. One after- Johnswort.
noon in New Orleans I collected nearly 1000 Sclerotinia sclerotiorum Shoot and Twig
sclerotia that were producing apothecia from Blight of lilac, grape, pistachio, soybean, peanut,
under a single camellia. and malaviscus; flower blight of camellia resem-
Control The first line of defense is exclusion. bling that caused by S. camelliae but far less
Most southern states have quarantines against serious. Tuber blight and storage rot; of Trillium.
known infected areas; they require that plants be This ubiquitous fungus more often causes stem
shipped bare-rooted, with all flower buds show- rots on its many different hosts. ▶ Rots.
ing color removed. Northern gardeners ordering
plants for greenhouses should insist on the same
precautions even without specific quarantines. Sclerotium
Practically all outbreaks of camellia flower blight
have been traced to plants shipped in cans, pre- Deuteromycetes, Mycelia Sterilia
sumably carrying sclerotia in the soil. The disease (Fungi Imperfecti)
has also appeared on flowers shipped in by air for
camellia shows. Schedules should state that all Asexual fruit bodies and spores lacking; there is
specimens become the property of the show com- merely a resting body, sclerotium, made up of
mittee, to be destroyed at the end of the show; no a compact, rounded mass of light-colored hyphae
blooms should be taken home for propagation. with a brown to black rind; parasitic, often on
Theoretically, because there is no conidial underground plant parts. Pellicularia has proved
stage to spread the fungus, this should be an to be the teleomorph state for some forms.
easy disease to eradicate, but it has not proved Sclerotium bataticola Ashy Stem Blight. See
so in practice. Camellias have thousands of ▶Macrophomina phaseoli under Rots.
flowers produced over a period of months. Sclerotium hydrophilum Blight of wild rice.
They drop into various ground covers, and it is Sclerotium oryzae Blight of wild rice.
almost impossible to find and destroy all infected Sclerotium rhizodes White Tip Blight of grass.
blooms before rotting tissues release sclerotia ▶ Snowmold.
into the litter. Some cities have quarantined Sclerotium rolfsii Southern Blight.
infected properties and provided a host-free ▶Pellicularia rolfsii.
period of 2 years, during which all flower buds
are removed from all camellias in the area, but
this approach has been only partially successful. Septoria
Various chemicals have been tried as ground
treatment to inhibit formation of apothecia. Deuteromycetes, Coelomycetes
Sclerotinia camelliae (see ▶Ciberinia
camelliae). Camellia Flower Blight, long known Pycnidia dark, separate, globose, ostiolate; pro-
in Japan, first noted in California in 1938, con- duce in spots, erumpent; conidiophores short,
firmed in Georgia in 1948, although probably conidia hyaline, narrowly elongate to filiform,
there several years previously, reported in Ore- several septate; parasitic, typically causing leaf
gon in 1949, Louisiana and North Carolina in spots, but also blights and blotches (see Fig. 1).
1950, South Carolina in 1954. There are about 1000 species.
Sirococcus 139

Septoria apiicola (Syn. Septoria apii and The generally destructive Septoria on chrysan-
S. apii-graveolentis) Celery Late Blight, gen- themum is S. chrysanthemi. ▶ Leaf Spots.
eral on celery, also on celeriac. The two species, Septoria petrosellini Leaf Blight of parsley, sim-
singly or together, produce the disease known as ilar to late blight of celery but confined to parsley.
late blight, first reported in Delaware in 1891 and *Recent study indicates these are one species
since causing much crop destruction, one and that the name should be S. apiicola.
California county reporting half a million dollars
loss from celery blight in 1908 and Michigan
a million in 1915. It was not known until 1932 Septotinia
that two distinct species were involved.
Early symptoms are similar. Large leaf spot, Ascomycetes, Helotiales,
due to S. apii, starts as a light yellow area, which Sclerotiniaceae
soon turns brown and dies. Spots are up to 1/4
inch in diameter, with small black pycnidia. In Stroma a definite, small, thin, elongate to angular
small leaf spot, due to S. apiigraveolentis, the black sclerotium maturing in host tissue after it
more common and destructive pathogen, has fallen to ground. Apothecia shallow cup-
pycnidia appear at the first sign of chlorotic spot- shaped, stipitate; spores hyaline, ovoid, one-
ting and are often outside of the indefinite mar- celled. Conidial stage a Septotis, with hyaline
gins of the spots, which are not over 2 mm. If spores, two or more cells, formed on sporodochia.
infection is severe, the spots fuse, and the leaves Septotinia podophyllina Leaf Blight of may-
turn brownish black and rot. Leaf stalks may also apple, found on leaves and stalks of this plant only.
be infected. Pycnidia winter on seed and in plant
refuse in garden and compost. A single pycnid-
ium of the small-spot fungus has an average of Servazziella
3675 spores, extruded in gelatinous tendrils.
A single leaf spot may average 56 pycnidia, and Ascomycetes, Amphisphaeriales
a single plant may have 2000 spots. Thus there
are enormous amounts of inoculum to be spread Perithecia immersed in a stroma, with long necks
by rain, insects, people, and tools. Some years converging into a disc; ascospores long, filiform,
ago on Long Island, when celery was inter- hyaline; conidia on a stroma.
cropped with spinach, it was found that workers Cryptospora longispora (see ▶Servazziella
spread blight spores on their sleeves as they cut longispora). Araucaria Branch Blight.
the spinach in early morning dew. And there is Servazziella longispora (formerly Cryptospora
a case on record where a man walked through his longispora). Araucaria Branch Blight. Lower
own blighted celery before taking a diagonal path branches are attacked first, with disease spread-
across his neighbor’s healthy field. In a few days ing upward; tip ends are bent and then broken off;
blight showed up all along that diagonal path. plants several years old may be killed. Prune off
Control The fungus usually dies in the seed coat and burn infected branches.
while the seed is still viable. Using celery seed
more than 2 years old obviates the necessity for
treatment. Fresh seed can be soaked in hot water Sirococcus
for 30 min at 118 to 120  F. Use crop rotation; do
not plant near where celery was grown the year Deuteromycetes, Sphaeropsidales,
before. Spray with bordeaux mixture or a fixed Sphaerioidaceae.
copper, starting in the seedbed when plants are
just out of the ground. Small, rounded, black, semi-immersed pycnidia
Septoria leucanthemi* Leaf Blight, Blotch on with wide ostioles; conidia hyaline, fusiform,
chrysanthemum, shasta daisy, and oxeye daisy. slightly constricted, 1-septate.
140 Blights

Sirrococcus elavigignenti-juglandacear- Sporodesmium scorzonerae (see ▶ Alternaria

um Canker of black walnut and butternut. scorzonerae). Salsify Leaf Blight. Leaves have
Sirococcus strobilinus Shoot Blight of Picea, many circular ispots, varying from pin point to
Abies, Pinus, and Tsuga spp. 1/4 inch, brown with red borders.

Sphaeropsis Stemphylium

▶ Cankers and Diebacks. ▶ Leaf Spots.

Sphaeropsis sapinea Shoot Blight of pine. Stemphylium vesicarium Stemphylium Blight
of onions. Lesions are nondelineated, light yel-
low to brown, water-soaked and range in length
Dothiora (Sphaerulina) from one centimeter to the entire leaf.

Ascomycetes, Dothideales
Perithecia innate or finally erumpent, not beaked;
paraphyses and paraphysoids lacking; spores Ascomycetes, Dothideales
hyaline, several-celled.
Dothiora wolfii (formerly Sphaerulina Asci in locules in an elongated stroma, which
polyspora). Twig Blight of sourwood, and is erumpent and superficial at maturity; spores
oxydendron. light brown, two-celled. Conidial state
Dothiora taxicola (formerly Sphaerulina taxi). Lecanosticta with brown conidia, two to four
Needle Blight of yew. cells, formed on a conidial stroma resembling
Sphaerulina polyspora (see ▶Dothiora wolfii). an acervulus.
Twig Blight of sourwood, and oxydendron. Mycosphaerella dearnessii (formerly
Sphaerulina taxi (see ▶Dothiora taxicola). Nee- Systremma acicola). Pine Brown Spot Needle
dle Blight of yew. Blight, on southern pines, most serious on
longleaf. The name and classification of the fun-
gus has been in dispute. The conidial stage,
Sporidesmium known since 1876, was first listed as Septoria,
later placed in Lecanosticta. The teleomorph
Deuteromycetes, Hyphomycetes state was named Scirrhia acicola in 1939 but
later transferred to Systremma because of its col-
Conidiophores clustered, dark, short, simple, ored spores.
each bearing a terminal conidium; conidia dark, Most injurious on seedlings, needle blight may
quite large, muriform with many cells, oblong to also injure large trees. Small, gray-green spots on
ovoid; usually saprophytic, sometimes parasitic. needles turn brown and form a narrow brown
Alternaria scorzonerae (formerly band, the needle tips dying. Three successive
Sporodesmium scorzonerae). Salsify Leaf Blight. seasons of brown spot kill longleaf seedlings.
Leaves have many circular ispots, varying from The fungus is more severe on trees in unburned
pin point to 1/4 inch, brown with red borders. areas because of accumulation of inoculum.
Leaves or whole tops die; roots are small and Spray seedlings in plantations with bordeaux
unsalable. The fungus winters as mycelium and mixture every 2 weeks from May to October or
spores in plant refuse. May be the same as November.
Alternaria tenuis. Systremma acicola (see ▶Mycosphaerella
Sporidesmium maclurae Leaf Blight of osage- dearnessii). Pine Brown Spot Needle Blight, on
orange. southern pines, most serious on longleaf.
Volutella 141

Thelephora Volutella

Basidiomycetes, Aphyllophorales Deuteromycetes, Hyphomycetes

Fruiting body leathery, upright, stalked; pileate or Sporodochia discoid, with marginal dark setae;
fan-shaped or much lobed, or in an overlapping conidiophores usually simple, in a compact pali-
series; hymenium on the underside, smooth or sade; conidia hyaline, one-celled, ovoid to
slightly warty; spores one-celled. oblong; parasitic or saprophytic (see Fig. 1).
Thelephora spiculosa Stem Blight found on Pseudonectria pachysandricola (see
azalea, fern, and other ornamentals in a Maryland ▶Volutella pachysandrae, Telemorph). Pachy-
garden. The fungus formed a dense weft of myce- sandra Leaf and Stem Blight. Large areas of
lium on surface of the soil and on plants. leaves turn brown to black, along with portions
Thelephora terrestris Seedling Blight, Smother. of stems, and in wet weather numerous pinkish
The mycelium ramifies in the soil, and the leathery spore pustules appear along stems.
fruiting body grows up around the stem of a seedling Volutella buxi Boxwood Leaf Blight, Nectria
conifer or deciduous tree, smothering it or strangling Canker. Pinkish spore occur as pustules on leaves
it without being actually parasitic on living tissue. and twigs. Leaves often turn straw-colored. See
The disease occurs most often in crowded stands in further under ▶ Cankers and Diebacks.
nurseries. The damage is seldom important. Volutella pachysandrae (formerly,
Pseudonectria pachysandricola). Pachysandra
Leaf and Stem Blight. Large areas of leaves turn
Tryblidiella brown to black, along with portions of stems, and
in wet weather numerous pinkish spore pustules
Ascomycetes, Patellariales appear along stems. The blight is most serious
when pachysandra has been injured or is too
Apothecia opening by a wide cleft; spores dark, crowded or is kept too moist by tree leaves falling
cylindrical, with several cells. into the bed. Spraying once or twice with bor-
Rhytidhysteron rufulum (formerly deaux mixture gives excellent control if severely
Tryblidiella rufula). Twig Blight on citrus. blighted plants have been removed before treat-
Tryblidiella rufula (see ▶Rhytidhysteron ment. Keep pachysandra thinned and sheared
rufulum). Twig Blight on citrus. back periodically.
Blotch Diseases

Diseases designated as blotch have symptoms Cercospora

that are intermediate between blights, where the
entire leaf or shoot dies, and leaf spots, where the ▶ Blights.
necrotic lesions are definitely delimited. Blotches Cercospora concors (see ▶Myrovellosiella
are irregular or indefinite large or small necrotic concors). Potato Leaf Blotch. An unimportant
areas on leaves or fruit. disease; leaflets turn yellow with small blackened
dead areas or larger, irregular brown areas.
Cercospora purpurea (see ▶Pseudocercospora
purpurea). Avocado Blotch, Cercospora Spot,
Alternaria considered the most important avocado disease
in Florida with no commercial variety entirely
▶ Blights. resistant.
Alternaria porri Purple Blotch of onion, also Myrovellosiella concors (formerly Cercospora
on garlic, and shallot, a problem in southern and concors). Potato Leaf Blotch. An unimportant
irrigated areas. Small, white, circular to irregular disease; leaflets turn yellow with small blackened
spots increase to large purplish blotches, some- dead areas or larger, irregular brown areas.
times surrounded by orange and yellow bands, on Pseudocercospora purpurea (formerly
leaves and flower stalks. Leaves often turn yellow Cercospora purpurea). Avocado Blotch,
and die beyond the spots; girdled stalks die before Cercospora Spot, considered the most important
seeds mature. Brown muriform spores form avocado disease in Florida with no commercial
a dusky layer on the blotches. Varieties with variety entirely resistant. Leaf spots are angular,
a waxy foliage are more resistant than those brown to chocolate brown, scattered and distinct,
with glossy leaves. The fungus winters as myce- less than 1/16 inch or coalescing to larger
lium and spores in crop refuse. Rotation, cleaning patches. With a hand lens, grayish spore groups
up plant debris, and seed treatment are can be seen on both sides of the leaf. Successive
recommended. crops of spores are produced in moist periods
Two other species of Alternaria, A. alternata throughout the year. Fruit spots are 1/4 inch or
and A. tenuissima, may cause purple or brown less in diameter, brown to dark brown, irregular,
blotches on onion, and there are physiological sunken, with cracked surfaces and grayish spore
races as well. tufts. Lesions are confined to the rind so that the

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144 Blotch Diseases

flesh is not affected, but the cracks furnish Gloeodes pomigena Sooty Blotch of Fruit on
entrance to anthracnose and other decay organ- apple, crabapple, blackberry, pear, and citrus, in
isms. The fungus winters in leaves, and appears to eastern and central states down to the Gulf, rare in
be progressively more abundant. the West. Fruit may be infected by heavy spore
dissemination from pycnidia on twigs of various
wild trees, including persimmon, prickly-ash,
Cladosporium white ash, bladdernut, hawthorn, red elm, sassa-
fras, maple, sycamore, and willow. On apples,
Deuteromycetes, Hyphomycetes clusters of short dark hyphae make a superficial
thallus on the cuticle, which appears as a sooty
Conidiophores dark, branched variously near brown or black blotch, 1/4 inch in diameter.
upper or middle portion, clustered or single; Numerous spots may coalesce to cover the
conidia dark, one-or two-celled, variable in size apple, a condition known as cloudy fruit. Because
and shape, ovoid to cylindrical, borne singly or in the lesion is superficial the fruit flesh is little
chains of two or three; parasitic or saprophytic. affected, but the grade and market value are
Cladosporium herbarum Leaf Blotch of reduced. On citrus the fungus does not penetrate
lilac. The fungus is usually secondary, sapro- the rind, and spots can be removed by gentle hand
phytic, following blights. rubbing. The disease develops in cool rainy
Cladosporium paeoniae Peony Leaf Blotch, weather during the summer. To control open up
Red Stem Spot, Measles. Leaf and stem spots the trees in the orchards to facilitate quick drying.
are purplish or brownish red. On stems the spots
are raised, upto 4 mm long; on leaves the lesions
are small specks. Small reddish spots are also Guignardia
present on floral bracts and petals. The disease
is widely distributed in commercial plantings and Ascomycetes, Dothideales
may sometimes destroy the value of flowers for
cutting. Cut down tops in fall as for Botrytis Perithecia immersed in substratum, stroma
blight. Spraying the ground with Elgetol in spring lacking, mouths papillate; spores hyaline
before new growth starts has given good control unequally two-celled, with lower cellcut off just
in some fields. before maturity.
Guignardia aesculi Horse-Chestnut Leaf
Blotch, Buckeye Leaf Blotch, general on horse-
Geastrumia chestnut and Ohio buckeye, sometimes on red
and yellow buckeye. Large, reddish brown
Geastrumia polystigmatis Sooty Blotch of blotches in foliage are, usually, surrounded by
Fruit on apple and blackberry. a yellowish area. Numerous pin-point black
dots, pycnidia, distinguish blotch from scorch
due to drought. Petioles often have reddish oval
Gloeodes spots. In a rainy season there is a good deal of
secondary infection from spores spread by wind
Deuteromycetes, Sphaeropsidales, and rain. Blotches appear on nearly every leaflet
Leptostromataceae with extensive defoliation. Primary infection in
spring comes from ascospores developed in
Pycnidia dimidiate, having a radiate cover over fallen overwinter leaves.
the top half only, on a dark subicle or mycelial Control Rake up and burn leaves in fall. Feed
crust; pseudoparaphyses present; conidia hyaline, trees that have been defoliated for successive
one-celled. years.
Septoria 145

Phyllosticta solitaria Apple Blotch, wide-

Mycosphaerella spread on apple and crabapple in eastern states,
serious in the South and in the Ozark section of
▶ Blights. Missouri, Arkansas, Oklahoma, and Texas. The
Mycosphaerella dendroides (Cercospora disease is also called fruit blotch, dryrot, black
halstedii, Anamorph). Pecan Leaf Blotch, on scab, late scab, cancer, and tar blotch. From Kan-
pecan in the South, on hickory in East and sas eastward it is second in importance to apple
South, a foliage disease of nursery and orchard scab. Leaf spots are very small, round, white,
trees. Olive green velvety tufts of conidio- with a single black pycnidium in the center of
phores and spores appear on undersurface of each. Larger elongate lesions are formed on
mature leaves in June and July (in Florida), veins, midribs, and petioles. Leaves do not turn
and yellow spots appear in corresponding yellow, but they drop prematurely if spots are
areas on upper leaf surfaces. Black pimplelike numerous. Cankers on twigs and branches are
perithecia are produced in the tufts about mid- located at leaf nodes or base of spurs. The first
summer, united in groups to give the leaf season they are small, purple to olive in color; the
a shiny black, blotched appearance after the next season this portion is tan and the new area
spores are washed away. In nursery trees, defo- dark purple, often slightly raised. Pycnidia
liation, starting with basal leaves and formed in twig lesions wash to leaves, fruit, and
progressing upward, may be serious. The dis- new shoots, discharged only after heavy rains and
ease is of little consequence to orchard trees in warm weather. Heavily fertilized trees are
unless they have been weakened by more susceptible.
overcrowding, borer attack, or other cause. Fruit blotches are brown, irregular, feathery at
The fungus winters in fallen leaves. To control the margin, studded with numerous pycnidia.
clean up fallen leaves. They afford entrance to secondary decay organ-
Mycosphaerella diospyri Leaf Blotch of Japa- isms and may develop deep cracks, but the blotch
nese persimmon. fungus itself is superficial. It winters in infected
Mycosphaerella lythracearum (Cercospora twigs and bark cankers.
punicae, Anamorph). Leaf Blotch, Fruit Spot of Control Secure healthy nursery stock. Some
pomegranate. The anamorph state has been varieties, including Grimes Golden, Jonathan,
thought the same as that on crape-myrtle Stayman Winesap, and Winesap, are rather
(Cercospora lythracearum), but is now consid- resistant.
ered distinct. Leaf spots are circular, small, dark
reddish brown to almost black, sometimes gray-
ish brown. Septoria

▶ Blights.
Phoma Septoria agropyrina Brown Leaf Blotch on
wheat grasses.
▶ Blackleg. Septoria elymi Speckled Leaf Blotch on
Phoma arachidicola Web Blotch of peanut. wheat grasses. A salt and pepper effect with
numerous pycnidia in pale gray, tan, or fuscous
Phyllosticta Septoria macropoda Purple Leaf Blotch, gen-
eral on blue grasses. Irregular blotches on blades
▶ Blights. are mottled greenish, then gray, tan or brown,
Phyllosticta congesta Leaf Blotch of garden finally bleached nearly white. Pycnidia are
plum. round, flattened, and light brown.
146 Blotch Diseases

losses in California greenhouses in 1953 and

Zygophiala reported from Pennsylvania in 1957. Small, radi-
ate patterns, resembling spider webs, appear as if
Deuteromycetes, Hyphomycetes dipped in oil. Leaves become brittle, turn yellow,
and die prematurely. The same fungus is present
A genus described from banana leaves in as a flyspeck on apple.
Zygophiala jamaicensis Greasy Blotch of car-
nation. A tropical fungus found causing serious

Broomrapes are parasitic seed plants like dodder long as has been believed, for they can live on some
and mistletoe. They are leafless herbs, of the family weeds between crops.
Orobanchaceae, living on roots of other plants and Orobanche ludoviciana Louisiana Broom-
arising from them in clumps of whitish, yellowish, rape on tomato and other plants, including Span-
brownish, or purplish stems. There are 130 or more ish needle and coldenia, becoming a problem in
species, mostly from North Temperate regions, but California. Tomatoes are stunted and do not pro-
few have any garden importance. The seed germi- duce a full crop of fruit.
nates in soil and produces a filiform plant body that Orobanche ramosa Branched Broomrapeh,
grows into the ground penetrating crown or root of Hemp Broomrape, most serious on hemp but
the host plant and forming a more or less tuberous parasitizing tomatoes, lettuce, tobacco, eggplant,
enlargement, from which the flowering shoots Ganra, Melitlotus, Silene, poppy mallow,
arise. Such shoots may be nearly naked, clothed cranesbil, Chaerophyllum, Verbena, Coreopsis,
only with a few scattered rudimentary leaves, fleabank, engelmann daisy, and other hosts in
or they may be covered with conspicuous, California. In small infections destroy the aerial
overlapping scalelike leaves. The seed may remain stems before they set seed; practice crop rotation.
viable in the soil several years but probably not as Deep plowing gives some control.

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Cankers and Diebacks

A canker is a localized lesion or diseased area light gray sunken areas in bark vary from several
often resulting in an open wound and usually on inches to a foot across. The fungus is confined to
a woody structure. Starting as a definite necrotic dead bark; trees are not injured.
spot, it may girdle cane, stem, or tree trunk, Dendrothele acerina (formerly Aleurodiscus
killing the water-conducting tissues so that the acerina). Bark Patch, wide-spread on maple.
most prominent symptom becomes a dieback.
When twigs and branches die back from the tip,
the condition may be a blight, with the pathogen
directly invading the dying area, or it may be
a secondary effect from a canker some distance
Deuteromycetes, Hyphomycetes
Conidiophores are long, slender, pigmented, and
highly branched; clusters of conidia at apex of
each branch; conidia ovoid, one-celled, hyaline.
Amphobotrys ricini Stem Canker on
Basidiomycetes, Aphyllophorales
texasweed and castorbean. Girdling stem canker;
of prostrate spurge.
Hymenium resupinate, of one layer, with
projecting spinose or short-branching cystidia
(swollen sterile cells); spores hyaline. Facultative
parasite on trees. Apioporthe
Aleurodiscus acerina (see ▶Dendrothele
acerina). Bark Patch, wide-spread on maple. Ascomycetes, Diaporthales
Aleurodiscus amorphus Balsam Fir Canker.
Cankers are formed on main stems of saplings, Perithecia in a black, carbonaceous stroma;
which are sometimes killed, but the fungus is also spores two-celled, hyaline; conidia in cavities in
widespread as a saprophyte on dead bark of firs a stroma.
and other conifers. Cankers center around a dead Anisogramma anomala (formerly Apioporthe
branch, are narrowly elliptical with a raised bor- anomala). Canker, Twig Blight of hazelnut.
der; the dead bark is covered with a light-colored Apioporthe anomala (see ▶Anisogramma
layer of the fungus. anomala). Canker, Twig Blight of hazelnut.
Aleurodiscus oakesii Oak Bark Patch, Smooth Apioporthe apiospora Twig Canker, Dieback
Patch of white oak. Irregularly circular, smooth, of elm.

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150 Cankers and Diebacks

larger branches and main stems. Cankers persist

Ascospora for many years, but extension stops after about
10 years.
Ascomycetes, Sphaeriales,
Perithecia with a subicle; paraphyses lacking;
spores two-celled, hyaline. ▶ Blights.
Ascospora ruborum (Hendersonia rubi, Botryodiplodia gallae (see ▶Lasiodiplodia
Anamorph). Cane Spot, Dieback of red and theobromae). Canker of oak.
black raspberry, dewberry. Botryodiplodia theobromae Canker of rose,
and citrus.
Lasiodiplodia theobromae (formerly
Atropellis Botryodiplodia gallae). Canker of oak.

Ascomycetes, Helotiales

Apothecia black, sessile or with short stalk; asci Botryosphaeria

clavate, with longer, hairlike paraphyses; spores
needlelike to slightly club-shaped, hyaline, one- ▶ Blights.
celled. Botryosphaeria dothidea Canker, Gummosis,
Atropellis apiculata Twig Canker. On southern and Dieback on peach, Bradford pear, thornless
pines. blackberry, sequoiadendron and sequoia.
Atropellis arizonica Branchand Truck Canker. Botryosphaeria obtusa Canker, on thornless
On western yellow pine. blackberry.
Atropellis pinicola (syn. A. piniphila) Pine Botryosphaeria ribis Saprophytic on dying tis-
Branch and Trunk Canker, on western white, sue, and var. chromogena, parasitic. Canker,
sugar, and lodgepole pines in Pacific Northwest Dieback of at least 50 woody plants, including
and California. Branches are girdled and killed, apple, avocado, eucalyptus, fig, forsythia, hick-
but not the trees. Perennial cankers are smooth, ory, pecan, pyracantha, quince, rhododendron,
elongated, flattened depressions covered with sequoia, sequoiadendron, sweet gum, and wil-
bark, in which appear very small black apothecia, low. See under ▶ Blights for the disease caused
2 to 4 mm in diameter. on currant and rose, under Rots for apple and
Atropellis piniphila (Cenangium piniphilum, avocado diseases.
Anamorph). Branch and Truck Canker on On redbud, sunken oval cankers nearly girdle
lodgepole and ponderosa pines on Pacific Coast, branches, the fungus entering through wounds,
on cultivated pines in the South. Trees 5 to 25 and dead and dying twigs. On rhododendron
years old are damaged by deformation of main there is a leaf spot and dieback similar to that
stem and branches. Infection is at branch whorls. caused by Phytophthora except that the surface is
Cankers are elongated, flattened depressions cov- roughened by protruding fruit bodies. Cankers on
ered with bark and copious resin. Apothecia have twigs, larger branches, and trunks of willow may
short stalks, are black with brownish discs, 2 to kill trees in a few years. Trunk lesions are very
5 mm across. small, 1/4 to 1/2 inch, and numerous or else large,
Atropellis tingens Branch and Truck Canker of from the union of several small cankers, with
native and exotic hard pines from New England fissured bark. Apples have watery blisters on
and Lake states to Gulf states. Slash pine saplings bark and decline in vigor. Forsythia has affected
are most susceptible. Smaller branches are gir- canes girdled and killed with conspicuous brown
dled; perennial target cankers are formed on dead leaves above the canker.
Ceratocystis (Ceratostomella) 151

Control Prune and burn dead twigs and heavily

infected branches; paint wounds with Ceratocystis (Ceratostomella)
a disinfectant followed by tree paint; avoid inju-
ries. Copper sprays may help. Ascomycetes, Micrascales
Botryosphaeria stevensii Canker, on juniper.
Perithecia with very long beaks, carbonaceous or
leathery; ascospores hyaline, one-celled; brown,
ovoid conidia and one-celled rodlike endospores
Botrytis formed inside tubelike conidiophores and
extruded endwise. Some species are important
▶ Blights. tree pathogens; see ▶Oak Wilt and ▶Dutch Elm
Botrytis cinerea Canker of rose. Disease under Wilts.
Ceratocystis fimbriata f. sp. platani (Endo-
conidiophora fimbriata f. sp. platani). Canker
Stain of London Plane, Plane Blight, on London
Caliciopsis plane and also on American plane or sycamore.
This serious disease started as a killing
Ascomycetes, Coryneliales epidemic in the Philadelphia area about 1935,
destroying city shade trees by the thousands
Stroma lobed, each lobe containing a single loc- there and in Baltimore during the next few
ule, which is finally wide open; perithecia years. The disease now extends from New
stalked; asci on long slender stalks; spores dark, Jersey to North Carolina and Mississippi.
one-celled. Trees show sparse foliage, smaller leaves, and
Caliciopsis pinea Pine Canker on eastern white elongated sunken cankers on trunks and larger
pine and other species, also on Douglas fir. Can- branches. Cross sections through cankers reveal
kers are sharply depressed areas in bark, reddish blue black or reddish brown discoloration of
brown and smoother than rest of bark, up to wood, usually in wedge-shaped sectors. First
several inches in diameter. Small, globose, clus- year cankers may not be more than 2 inches
tered black pycnidia, and stalked perithecia wide and a yard or so long, but they widen
looking like slender black bristles, arise from annually, girdling and killing trees in 3 to
stroma in cankered bark. The disease is most 5 years. Several cankers coalescing around the
serious on suppressed saplings. trunk kill more quickly. Once infection starts,
the tree is doomed.
Ascospores and the two types of conidia are
produced in moist spring weather (see Fig. 1).
Encoelia (Cenangium) They may be spread by rain a short distance, but
most dissemination is by man in pruning opera-
▶ Blights. tions, and ordinary tree paint carry viable spores.
Cenangium singulare (see ▶Encoelia Some beetles may be vectors. Infection is solely
pruinosa). Sooty-Bark Canker of aspen, on through wounds.
Populus tremuloides in Rocky Mountain area. Control Do not try to save trees where trunk has
Encoelia pruinosa (formerly Cenangium been invaded; diseased branches may sometimes
singulare). Sooty-Bark Canker of aspen, on be removed, cutting at least 3 feet from infected
Populus tremuloides in Rocky Mountain area. area. Do not prune unless absolutely necessary
Cankers on older trees, at any point on trunk up and then only in winter when trees are less sus-
to 60 to 70 feet may extend 10 to 15 feet before ceptible. Use tree wound dressing fortified with
they girdle the tree. The bark is sooty black with a disinfectant.
a thin white outer layer. Ceratocystis sp. Canker and Dieback on poplar.
152 Cankers and Diebacks

Fig. 1 Spore Formation of

Some Canker Fungi.
Coniothyrium, small dark
spores on short
conidiophores in
pycnidium; Seiridium,
(formerly Coryneum), dark,
septate spores in acervulus;
Cytospora, sausage-shaped
spores in valsoid pycnidia
expelled in cirrhi;
Endoconidiophora, spores
formed on inside of
conidiophores; Nectria,
twocelled bright ascospores
in reddish perithecia
clustered on bark; Phoma,
hyaline spores in pyncnidia
formed in spots on bark

Chondropodium Coniothyrium

Deuteromycetes, Coelomycetes Deuteromycetes, Hyphomycetes

Pycnidia stromatic, stalked, columnar, externally Pycnidia black, globose, separate, erumpent,
black, hard, internally gelatinous; conidiophores ostiolate; conidiophores short, simple; conidia
simple; conidia hyaline, with several cells, cres- small, dark, one-celled, ovoid or ellipsoid; para-
cent-or sickle-shaped; weakly parasitic or sitic or saprophytic (see Fig. 1).
saprophytic. Coniothyrium fuckelii (Anamorph, Diapleella
Chondropodium pseudotsugae Bark Canker of coniothyrium). Rose Commom Canker, Stem
Douglas-fir. This is a superficial canker with Canker, widespread on rose, also causing rasp-
outer layers of bark killed over small, circular to berry cane blight (see ▶Leptosphaeria under
elliptical areas, in which pycnidia project as Blights), sometimes associated with apple rots,
short, blunt, black spines. Trees are not notice- peach cankers, and stem canker of Virginia
ably injured. creeper. Of the three species of Coniothyrium
that cause rose cankers, C. fuckelii is by far the
most common. Any plant part may be affected.
Pycnidia have even been found within blackspot
Colletotrichum lesions on leaves, but this is primarily a cane
disease, starting as a red or yellow spot on bark,
▶ Anthracnose. drying out and turning brown as it increases in
Colletotrichum acutatum Canker and Dieback size, with the epidermis somewhat wrinkled and
on Japanese maple. perhaps rupturing irregularly over sooty masses
Seiridium (Coryneum) 153

of very small, olive brown spores. The stem may purple margin, contrasting sharply with the green
be girdled with dieback to that point. of the cane. The center of the spot turns light
Stem cankers are found around insect punc- brown as the cells die, and little longitudinal
tures, thorn pricks, leaf or thorn scars, or abra- slits appear over the developing pycnidia. Spores
sions caused by tying, but the majority of cankers are olive brown, nearly twice the size of C.
are formed at the cut end of a cane when a stub fuckelii, and released through epidermal slits
has been left in pruning above a leaf axil or bud. instead of being spread in a sooty mass under
Roses cut properly close to a bud seldom develop the epidermis. Cankers formed under the winter
this canker. A rose stub usually dies back to the protection of soil are black when roses are first
first node, and since this fungus is a weak para- uncovered in spring, which explains the name
site, it starts most readily in such dead or dying Brandfleckenkrankheit, meaning fire-spot
tissue. When a cut is made close to the node, it is disease.
quickly callused over, and the callus is a good C. wernsdorffiae is a cold temperature fungus,
defense against wound fungi. infecting rose canes under the winter covering,
Control Prune out cankered and dying stems as entering through insect wounds, thorn scars,
soon as noticed. Make all cuts just above a bud or scratches, and occasionally through dormant
leaf axil, not only at spring pruning but in cutting buds. During a 4-year investigation at Ithaca,
flowers for the house or cutting off dead blooms I found no infection on canes not hilled with
during the season. earth or other moist cover over winter and no
Coniothyrium rosarum Rose Graft Canker. natural infection during the summer.
This is a disease of roses under glass, starting at Control Omit the usual winter protection of soil
the union of stock and scion in the warm moist or other materials that keep canes moist. If brand
propagating frame and continuing in a large canker is a problem, just fasten canes of climbers
amount of dead wood when plants are removed down near the ground, uncovered, and hope for
to the greenhouse bench. Some consider the path- the best. Loss from winter injury will be less than
ogen a form of C. fuckelii. Having measured from the canker. Cut out diseased canes carefully.
spores of the type specimen, in the Kew Herbar-
ium, I think they are distinct species, but that
some cases of graft canker are due to the common Seiridium (Coryneum)
canker fungus.
Coniothyrium wernsdorffiae Rose Brand Can- ▶ Blights.
ker, a rather rare but very serious disease. The Coryneum cardinale (see ▶Seiridium
pathogen was named in Germany in 1905 and cardinale (Leptosphaeria sp., Telemorph)).
was not reported in this country until 1925, Coryneum Canker of Cypress, Bark Canker of
although it was subsequently shown to have cypress, incense ceder, common juniper and ori-
been collected in Canada in 1912 and in Pennsyl- ental arborvitae.
vania and Minnesota in 1914 and 1916. In 1926 Coryneum foliicola Twig Canker, Fruit Rot,
a severe epiphytotic appeared at Ithaca, New widespread on apple, affecting twigs, foliage
York, in the Cornell rose garden, infecting about and fruit.
90 % of the climbers so seriously that the canes Seiridium cardinale (formerly Coryneum
had to be cut to the ground. Since then it has been cardinale (Leptosphaeria sp., Telemorph)).
reported from a few other states, but in several Coryneum Canker of Cypress, Bark Canker of
instances it has been confused with common cypress, incense ceder, common juniper and ori-
canker. ental arborvitae. This disease, since its discovery
Small, dark reddish spots on canes enlarge and in 1927, has been gradually exterminating Mon-
acquire a more or less definite reddish brown or terey Cypress in most parts of California and is
154 Cankers and Diebacks

also serious on Italian cypress. Twigs, branches, Amphiporthe castanae (formerly

and whole trees turn sickly, lose their leaves, and Cryptodiaporthe castanea). Dieback, Canker of
finally die. The fungus attacks living bark and Asiatic Chestnut, widespread, chiefly on seed-
cambium, girdling twig and branch. Cankers lings or on larger trees in poor sites. Canker starts
appear first at base of lateral twigs; they are as a brown discoloration of bark of the trunk,
slightly sunken, dark, resinous, rough, with limb, or twig, often girdling twig and then invad-
black spore pustules. Conidia have dark median ing larger branch. Leaves on girdled branches
cells, five cross-walls (see Fig. 1). They are wilt without yellowing, turn brown, and die.
spread by tools, in nursery stock, by wind and Bark splitting over callus formation at edge of
rain, and perhaps by birds and insects. Infection diseased area forms pronounced canker. Conidia,
appears first in upper parts of trees, usually in two-celled, fusoid, are formed in pustules in bark;
spring during moist weather. Yellowing and beaked perithecia are formed in groups by
browning of foliage together with gummy ooze midsummer.
at the cankers form conspicuous symptoms. Control Maintain vigor; plant on well-drained,
Control Drastic surgery, removing wood well fertile soil. Prune out diseased portions several
below infected parts, and spraying foliage inches below affected area.
heavily with bordeaux mixture help some, but Crytodiaporthe aculeans, Sporocybe rhois,
with heavy infection the price of saving healthy Anamorph (see ▶Amphiporthe aculeans). Die-
trees is the removal and destruction of all dis- back, Canker of sumac.
eased specimens. California citizens, threatened Cryptodiaporthe castanea (see ▶Amphiporthe
with the loss of the famous native stands of Mon- castanae). Dieback, Canker of Asiatic Chestnut,
terey cypress at Point Lobos and Cypress Point, widespread, chiefly on seedlings or on larger
voluntarily destroyed their own plantings by the trees in poor sites.
thousands. Cryptodiaporthe salicella Twig and Branch
Canker of willow.

Ascomycetes, Diaporthales
Ascomycetes, Rhytismatales
Production of perithecial ascocarps produced in
a stroma of fungal and substrate tissues or directly Apothecia effuse, splitting irregularly; paraphy-
from somatic hyphae on the substrate. Asco- ses present; spores hyaline, one-celled.
spores are hyaline to brown and one-to-several- Cryptomyces maximus Blister Canker on com-
septate. mon and purple osier.
Cryphonectria parasitica Canker on oak.


Cryptodiaporthe Ascomycetes, Diaporthales

Ascomycetes, Sphaeriales, Valsaceae Perithecia in a circle in a stroma, with long necks

converging in a common canal; spores one-
Like Diaporthe but without blackened zones in celled, hyaline; conidia borne on surface of
substratum; spores hyaline, two-celled. stroma.
Amphiporthe aculeans (formerly Cryptosporella umbrina Rose Brown Canker,
Cryptodiaporthe aculeans; Sporocybe rhois, a widespread and serious rose disease, first
Anamorph). Dieback, Canker of sumac. reported in Virginia in 1917 but known from
Cryptosporium 155

Control The best time to take care of brown

canker is at spring pruning. Cut out every dis-
eased cane possible. A dormant lime sulfur
spray, immediately after pruning, kills spores
that may have been spread in the process
and may inhibit the fungus in initial lesions.
Copper or sulfur sprays largely prevent
summer infections. Brown canker is more likely
to be serious where roses are overprotected for
winter with salt, hay, leaves, or other material
added to the mound of soil. I have no trouble
with brown canker when roses are left unhilled
over winter.
Cryptosporella viticola Dead-Arm Disease of
grapes, Branch Necrosis, widespread, especially
in the Northeast, serious in Illinois, important in
California. Small, angular spots with yellowish
margins and dark centers are formed on
leaves, stems of flower clusters and canes. The
latter may split to diamond-shaped cankers, and
by the next season the arm is dead or producing
yellowed, dwarfed and crimped foliage. Lesions
on cluster stems advance into fruit late in the
season causing rotting. Pycnidia are developed
Fig. 2 Brown Canker on Rose
on old wood; infection is often through pruning
herbarium specimens to have been present since wounds.
1903. The fungus was first placed in Diaporthe Control Make pruning cuts at least 6 inches
because of occasional two-celled spores. below the lower margin of the infected part.
Symptoms are most noticeable on canes, Spray with bordeaux mixture when spores are
starting with very small purplish spots, the center extruded.
soon turning white with a reddish purple margin
(see Fig. 2). Many small spots may be grouped on
a single cane. During the winter, and especially
on portions of canes covered with earth, cankers Cryptosporium
or girdling lesions are formed, often several
inches long, with tan centers and purplish bor- Deuteromycetes, Coelomycetes
ders. In moist weather the surface of these large
cankers is covered with yellow spore tendrils Acervuli erumpent, becoming cup-shaped or
from pycnidia just under the bark; asci are also disclike; stroma brownish; conidiophores simple
extruded in tendrils from perithecia. or branched; conidia hyaline or subhyaline, one-
Leaf spots are small purplish specks or larger celled filiform.
dead areas, cinnamon buff to white, bordered Cryptosporium minimum Canker on rose, not
with purple and with black pycnidia in the center. common.
Marginal spots are subcircular. Buds are some- Cryptosporium pinicola (see ▶Gelatirosporium
times blighted; exposed petals of flowers have piricola). Canker, Branch Mortality of Abies spp.
cinnamon-buff spots without the purple border. Gelatirosporium piricola (formerly
Infection is through wounds and also uninjured Cryptosporium pinicola). Canker, Branch Mor-
tissue. tality of Abies spp.
156 Cankers and Diebacks

Cytospora chrysosperma (Teleomorph, Valsa

Cylindrocarpon sordida). Cytospora Canker of poplar, aspen,
cottonwood, willow, occasional on mountain-ash,
▶ Rots. maple, cherry, and elder. Cankers form on trunks
Cylindrocarpon didymium Bole Canker on and large branches, most often on trees of low
apple. vigor. Bark is discolored in more or less circular
Cylindrocarpon cylindroides Canker, Branch areas; sapwood is reddish brown. In old cankers
Mortality of Abies spp. exposed wood is surrounded by layers of callus
tissue. In moist weather spring spore tendrils are
extruded from pycnidia in dead bark. Perithecia
Cylindrocladium are found infrequently in aspen, arranged circu-
larly around a grayish disc; they are flask-shaped
▶ Blights. with long necks pushing through the bark. Twigs
Cylindrocladium scoparium Crown Canker of and small branches may die back without
rose. The cane is attacked at or just below the a definite canker. The fungus is often present on
union of stock and scion, the bark darkening into healthy trees, not becoming pathogenic until the
a black, water-soaked punky region. The cankers trees are weakened by neglect, drought, pollarding
girdle but do not kill the canes; there are fewer or other causes. Entrance is through wounds. Lom-
and more inferior blooms. The disease was long bardy and Simon poplars are frequently killed.
thought confined to greenhouse roses but has Control Remove dead and dying branches and
appeared once or twice in outdoors fields. The trees with extensive cankers. Avoid wounds; feed
fungus lives in the soil and enters through wounds and water as necessary. Plant poplars that are less
in the presence of sufficient moisture. Before susceptible than Lombardy. Rio Grande cotton-
planting of fresh stock, greenhouse benches wood is resistant to twig blight.
should be washed with boiling water and soil Cytospora kunzei (Teleomorph, Leucostoma
sterilized or changed. kunzei). Cutospora Canker of spruce. Twig
The same fungus injures seedling conifers in Blight, common and serious New England to the
nursery rows, causing damping-off, root rot, stem Midwest. Cankers start around bases of small
canker and needle blight to white pine and Doug- twigs or on trunks. Browning and death of Colo-
las-fir. See under ▶ Blights for a discussion of the rado blue spruce branches starts near the ground
pathogen on cuttings of azaleas and other and progresses upward, a large flow of resin on
ornamentals. affected limbs. Needles drop immediately or per-
sist for a time. Cankers are formed near resin
spots and yellow tendrils extruded. Spores are
splashed by rain and wind to other branches;
Cytospora infection is mostly through wounds.
Another form of the pathogen, Valsa kunzei
Deuteromycetes, Coelomycetes var. superficialis, occurs on pine and variety
kunzei on balsam fir, Douglas-fir, larch and
Cosmopolitan species, anamorph state of Valsa. hemlock.
Pycnidia in a valsoid stroma with irregular cavi- Control There are no satisfactory control mea-
ties, incompletely separated; conidia hyaline, sures except removal of diseased branches and
one-celled, allantoid, expelled in cirrhi perhaps carefully excising cankered bark.
(see Fig. 1). Spraying with bordeaux mixture has been
Cytospora abietis Canker, Branch Mortality of recommended but is seldom very effective.
Abies spp. Avoid wounding ornamental trees with lawn
Cytospora pruinosa Canker, Dieback of ash, mowers; sterilize pruning tools between cuts;
on twigs and branches. feed to renew vigor.
Dasyscyphus 157

Cytospora leucostoma Canker of black cherry. Dasyscyphus pseudotsugae (see ▶Lachnellula

Cytospora nivea Canker, Dieback of poplar pseudotsuga). Canker on Douglas-fir. Swollen
and willow, similar to that caused by C. open cankers, 2 to 3 inches long, are formed on
chrysosperma; occasional. suppressed saplings.
Cytospora leucosperma Branch Canker of Dasyscyphus resinaria (see ▶Lachnellula
elder. resinaria). Canker on balsam fir. Swollen cankers
Cytospora sp. Canker on alder and pecan. at base of branches; younger stems girdled and
Cytospora spp. Cytospora Canker of Italian killed.
Prunce, causing severe injury to prune and apri- Dasyscyphus willkommii (see ▶Lachnellula
cot in Idaho orchards since 1951, also present on willkommii (Trichoscyphella willkommii syn.
cherries, peach, apple and willows. Lachnellula wilkommii)). European Larch Can-
Some orchards have been lost, others hard hit. ker. Found in Massachusetts in 1927 on nursery
Symptoms are yellow to brown flags of dead stock from Great Britain.
leaves and erumpent, gummy cankers or elon- Lachnellula agassizi (formerly Dasyscyphus
gated necrotic streaks in the bark. All suspicious agassizi). Common on blister-rust lesions of
wood should be cut out, hauled out of the orchard, white pine; saprophytic on dead branches.
and burned. Lachnellula pseudotsuga (formerly
▶Valsa cincta for further discussion of can- Dasyscyphus pseudotsugae). Canker on Doug-
kers on stone fruits. las-fir. Swollen open cankers, 2 to 3 inches long,
are formed on suppressed saplings.
Lachnellula resinaria (formerly Dasyscyphus
resinaria). Canker on balsam fir. Swollen cankers
Dasyscyphus at base of branches; younger stems girdled and
Ascomycetes, Helotiales, Helotiaceae Lachnellula subtilissima (formerly
Dasyscyphus calycina (Trichoscyphella
Apothecia stalked, white and hairy on the outside hahniana)). On larch and fir, ordinarily
with a bright disc; paraphyses filiform; asci a saprophyte but can be a weak parasite; occa-
inoperculate; spores elliptical to fusoid. sional on blister-rust cankers.
Dasyscyphus agassizi (see ▶Lachnellula Lachnellula willkommii (formerly
agassizii). Common on blister-rust lesions of Dasyscyphus willkommii (Trichoscyphella
white pine; saprophytic on dead branches. willkommiisyn. Lachnellula wilkommii)). Euro-
Dasyscyphus calycina (Trichoscyphella pean Larch Canker. Found in Massachusetts in
hahniana) (see ▶Lachnellula subtilissima). On 1927 on nursery stock from Great
larch and fir, ordinarily a saprophyte but can be Britain. Infected trees were removed and the
a weak parasite; occasional on blister-rust fungus not seen again until 1935, near the
cankers. original location. Perennial branch or trunk
Dasyscyphus ellisiana Canker of Douglas-fir cankers are flattened depressions, swollen on
and pine in eastern United States. This is the flanks and on the opposite side of the
a native fungus on twigs and branches of native stem. Neighboring bark is somewhat cracked
and introduced pines and on basal trunk and dark with heavy exudation of resin. Cup-
and branches of Douglas-fir. Bark on trunk shaped apothecia are 3 to 6 mm across with
may be infected for 10 to 15 feet, with white hairs and orange to buff discs, very short
copious resin flow and numerous swellings, but stalks. Young trees may be killed; older trees
trees are not killed. Apothecia are short-stalked, usually survive. Frost wounds are a contributing
covered with white hairs, with an orange to yel- but not an essential factor. Promptly remove all
low disc, 2 to 4 mm across. Remove trees with trees showing cankers; continue periodic
trunk cankers. inspection.
158 Cankers and Diebacks

sesbania, indigo, spiny amaranth, vetch and

Dermea (Dermatea) soybean.
Diaporthe pruni Twig Canker on black cherry;
Ascomycetes, Helotiales D. prunicola on American plum.

Apothecia small, brownish to black with

a circular opening; innate at first, on a stromoid Dichotomophthora
base, rupturing host at maturity; spores one-
celled, hyaline, globose to oblong. Cup fungi Deuteromycetes, Hyphomycetes
(ascocarpcup-shaped); excipulum of subglobose
cells; sclerotia absent. Conidiophores brown, branching dichotomous to
Dermatea acerina (see ▶Dermea acerina). subdichotomous, elongated, terminal branches
Bark Canker of maple, occasional. 4–8 lobed each lobe bearing single conidium;
Dermatea balsamea (see ▶Dermea balsamea). conidia dark, ovoid to elongate–ovoid, 1 to 6
Twig Canker of hemlock. celled.
Dermatea livida (see ▶Pezicula livida). Bark Dichotomophthora portulacae Stem Canker
Canker of redwood. and Root Rot on common purslane.
Dermea acerina (formerly Dermatea acerina).
Bark Canker of maple, occasional.
Dermea balsamea (formerly Dermatea Pseudomassaria (Didymella)
balsamea). Twig Canker of hemlock.
Dermea pseudotsugae Branch Canker on fir. ▶ Blights.
Pezicola livida (formerly Dermatea livida). Didymella sepincoliformis (see
Bark Canker of redwood. ▶Pseudomassaria sepincolaeformis). Dieback
of rose.
Pseudomassaria sepincolaeformis (formerly
Didymella sepincoliformis).
Diaporthe Dieback of rose.

▶ Blights.
Cryphonectria cubensis (formerly Diaporthe Diplodia
cubensis). Canker of Eucalyptus spp.
Diaporthe eres Canker, Dieback of English ▶ Blights.
holly in the Northwest. The fungus name is Diplodia sp. Rose Dieback, sometimes after
a species complex that may include a Diaporthe drought and other contributing factors. In Texas
on rose petals and one causing a peach constric- the disease is most evident in autumn, progressing
tion disease. on roses in storage or overwintering in the ground.
Diaporthe cubensis (see ▶Cryphonectria Canes die from tip downward, often starting in the
cubensis). Canker of Eucalyptus spp. flower stem. Diseased wood turns brown or black,
Diaporthe eres Canker, Dieback of English and is somewhat shriveled. Pycnidia are produced
holly in the Northwest. in dead canes. Improve general rose vigor; use
Diaporthe helianthi Canker of sunflower; also fungicides as for blackspot. May also cause
leaf spot of sunflower. canker of Russian olive.
Diaporthe oncostoma Canker, Dieback of Diplodia camphorae Canker, Dieback of cam-
black locust. phor-tree.
Diaporthe phaseolorum var. caulivora Canker Diplodia infuscans (see ▶Sphaeropsis
of painted spurge, prickly sida, redweed, morn- hyalina). Ash Canker and Dieback, northeastern
ing-glory, black nightshade, jacquemontia, hemp states.
Dothiorella 159

Diplodia juglandis Dieback, widespread on slightly darker, sunken areas, often at base of
branches of walnut. twigs and limbs, and become elongated. The
Diplodia mutila Stem Canker on Laburnum. bark is killed to the cambium; sapwood is
Branch Dieback on juniper. brown. If a stem is completely girdled, it dies;
Diplodia natalensis (see ▶Lasiodiplodia otherwise, callus formation goes on through the
theobromae). Stem Canker of prickly-ash; Die- summer, over the canker. In time diseased bark
back of citrus twigs, also causing citrus stem-end turns brown and cracks. Spores are extruded in
rot. amber tendrils, drying to brown, and are washed
Diplodia quercina Canker and Blight of oaks. to wounds in the wood.
Diplodia sophorae Dieback of pagoda tree. Control Destroy infected stock in nurseries and
Diplodia sycina Canker, Dieback of fig. plantations; do not move stock from a nursery
Lasiodiplodia theobromae (formerly Diplodia where the disease is known. Avoid pruning and
natalensis). Stem Canker of prickly-ash; Die- other wounds so far as possible; sterilize tools
back of citrus twigs, also causing citrus stem- between cuts. Spraying nursery trees with bor-
end rot. deaux mixture in spring may be helpful.
▶ Rots. Dothichiza populea (see ▶Discosporium
Sphaeropsis hyalina (formerly Diplodia populeum). Dothichiza Canker of poplar; Euro-
infuscans). Ash Canker and Dieback, northeast- pean Poplar Canker, widespread but sporadic as
ern states. a branch and trunk canker.

Discella Dothiora

Deuteromycetes, Coelomycetes Ascomycetes, Pseudosphaeriales

Pycnidia cupulate or discoid; spores 2-celled, Ascocarps hairy and phragonosporous or

hyaline. muriform ascospores are colored.
Discella carbonacea (see ▶Discella Dothiora polyspora (see ▶Sydowia
microsperma). Twig Canker of willow. dothideoides). Canker of aspen.
Discella microsperma (formerly Discella Sydowia dothideoides (formerly Dothiora
carbonacea). Twig Canker of willow. polyspora). Canker of aspen.

Dothichiza Dothiorella

Deuteromycetes, Coelomycetes Deuteromycetes, Coelomycetes

Pycnidia innate, finally erumpent; conidiophores Pycnidia dark, globose, grouped in a subcortical
lacking; conidia hyaline, one-celled. stroma; conidiophores simple, short; conidia hya-
Discosporium populeum (formerly Dothichiza line, one-celled, ovoid to ellipsoid; parasitic or
populea). Dothichiza Canker of poplar; Euro- saprophytic on wood.
pean Poplar Canker, widespread but sporadic as Botryodiplodia gallae (formerly Dothiorella
a branch and trunk canker. Lombardy poplars are quercina). Dothiorella Canker of oak, very
most susceptible, but hosts include black and destructive to red and white oaks in Illinois,
eastern cottonwoods, balsam, black and Norway affecting twigs, branches, and occasionally
poplars. Japanese poplars are rather resistant. trunks. Cankers are dark brown, elongated,
Young trees in nurseries are most injured, can- sunken, often with cracks at the margin. Pustules
kers often starting around wounds. They start as of pycnidia develop in bark and erupt through
160 Cankers and Diebacks

cracks, spores oozing on the surface. Sapwood

has dark streaks. Fusarium
Dothiorella fraxinicola Branch Canker of ash.
Dothiorella quercina (see ▶Botryodiplodia ▶ Rots.
gallae). Dothiorella Canker of oak, very destruc- Fusarium moniliforme var. subglutinans (see
tive to red and white oaks in Illinois, affecting ▶Fusarium subglutinans). Pitch (Branch) Can-
twigs, branches, and occasionally trunks. kers and Shoot Dieback on southern pine species,
Dothiorella sp. London Plane Canker, first loblolly and pond pines.
noted in New York City in 1947. Infected trees Fusarium oxysporum Stem Canker on peanut.
have sparse, undersized foliage and narrow, lon- Fusarium solani Stem Canker of sweet potato,
gitudinal cankers on trunk and branches, varying black walnut, oak, and poinsettia.
from 1 to 4 inches wide and often extending from Fusarium subglutinans (formerly Fusarium
ground level to branch top. The bark is rough, moniliforme var. subglutinans). Pitch (Branch)
deeply fissured; inner bark is brown, dry; sap- Cankers and Shoot Dieback on southern pine
wood is only superficially discolored. Branches species, loblolly and pond pines.
wilt and die back.
Dothiorella ulmi Dieback, Wilt of elm. ▶ Wilt

Deuteromycetes, Coelomycetes
Pycnidia one to several in a stroma, spherical or
▶ Blights. flattened, subepidermal, erumpent; opening sep-
Endothia gyrosa Branch Canker on oak. arately or with a common pore; conidiophores
simple, short; conidia hyaline, one-celled, fusoid;
parasitic or saprophytic.
Fusicoccum amygdali Twig Canker of peach,
Epicoccum increasingly important on peaches in North
Atlantic coastal area. Leaf spots are large, irreg-
▶ Leaf Spots. ular or circular, often zonate, brown with
Epicoccum nigrum Canker on thornless scattered pycnidia near center. Cankers at buds
blackberry. and bases of young twigs result in death of the
distal portions; trunks of young trees may be
girdled. Infections occur throughout the season
at bud scales, stipules, fruit and leaf scars. Prune
Eutypa only in winter.
Fusicoccum elaeagni Canker on Russian-olive.
Ascomycetes, Xylariales, Diatrypaceae

Stroma effuse; perithecia with necks at right

angles to surface. Gibberella
Eutypa armeniacae syn. E. lata Cytosporina
Dieback of apricot and of grape; Twig Canker ▶ Blights.
on cherry and chokecherry. Anamorph state Gibberella baccata Twig Canker of acacia,
reported from California in 1962, perithecia in ailanthus, apple, boxwood, mimosa, mulberry,
1965. Bark cankers with gum are formed at prun- and also on other plants where twig blight is the
ing wounds. most important symptom. ▶ Blights.
Griphosphaeria 161

Gloeosporium Griphosphaeria

▶ Anthracnose. Ascomycetes, Amphisphaeriales

Gloeosporium sp. Canker on holly.
Gloeosporium sp. (Gnomonia rubi, Perithecial wall carbonaceous, mouths papillate;
Teleomorph). Canker on thornless blackberry. spores dark, with several cells.
Discostroma corticola (formerly
Griphosphaeria corticola (Anamorph,
Seimatosporium lichenicola). Rose Canker, Die-
Glomerella back. Cankers are formed near base of canes,
often showing dark glistening pustules of conidia.
▶ Anthracnose. Occasionally when the canker has girdled the
Glomerella cingulata Camellia Dieback, Can- cane, a large gall forms above the lesion (see
ker, widespread; sometimes on azalea, black- Fig. 3). It resembles crown gall but is apparently
berry, bittersweet, rose, raspberry, soapberry, due to interference with downward transfer of
mountain-ash, and English ivy; also causing bitter food. Cut out infected canes.
rot of apple (▶ Rots) and anthracnose of various Griphosphaeria corticola (see ▶Discostroma
hosts (▶ Anthracnose). Camellia tips die back; corticola (Anamorph, Seimatosporium
leaves wilt, turn dull green and finally brown. lichenicola)). Rose Canker, Dieback. Cankers
The stem dries out, turns brown, and there is are formed near base of canes, often showing
a girdle of dead bark. Elliptical cankers are pre- dark glistening pustules of conidia.
sent on older wood. Infection is solely through
wounds, principally leaf scars in early spring but
also through bark wounded by cultivating tools or
lawn mowers, frost cracks, or the graft union.
Governor Moulton, Professor Sargent, and
some other varieties are rather resistant; Flora
Plena, Prince Eugene Napoleon, and many others
are highly susceptible. Spraying with bordeaux
mixture to prevent infection through leaf and bud
scars gives fair control.

Cryptosporiopsis (Glutinium)

Deuteromycetes, Coelomycetes

Pycnidia innate, without a stroma; spores borne at

tip and sides of conidiophores, hyaline, one-
Cryptosporiopsis pruinosa (formerly
Glutinium macrosporum). Canker, Fruit Rot of
Glutinium macrosporum (see
▶Cryptosporiopsis pruinosa). Canker, Fruit Rot
of apple. Fig. 3 Discostroma Canker on Rose
162 Cankers and Diebacks

commonly attacked, balsam poplar less fre-

Hendersonula quently. This is usually a forest, rather than
a home garden, disease. Trees less than 30 years
Deuteromycetes, Coelomycetes old, growing on poor sites, are most susceptible.
Trunk cankers start as small, yellow to reddish
Pycnidia black, stromata, one to several per stroma, brown, slightly sunken areas, centering around
locules occurring at different levels in stroma; a wound, there grow together to form a canker
conidophores long, flexuous; conidia often extruded marked off by vertical cracks. The bark is mottled,
in cirrhi; at first one-celled, hyaline to yellowish, gray, with black patches where the blackened cor-
later becoming three-to four-celled and dark. tex is exposed. Conidia appear in blisterlike stro-
Hendersonula toruloidea Canker on Arbutus mata on first-and second-year cankers, whereas
menziesii. perithecia are formed on third-year cankers in
hard, black stromata covered with a white pruinose
coat. Ascospores are ejected in winter. Eliminate
Hymenochaete infected trees when thinning stands.

Basidiomycetes, Aphyllophorales
Pileus, fruiting structure, resupinate, of several
layers, with long, stiff, usually brown setae Deuteromycetes, Coelomycetes
Hymenochaete agglutinans Hymenochaete Kabatina juniperi Blight on eastern red cedar;
Canker on apple, birch, hazelnut, sweetgum, mis- conidia produced in black acervuli on discolored
tletoe, and various young hardwoods. When an foliage.
infected dead stem comes in contact with a live
one, the mycelium forms a thin leathery fruiting Lachnellula
body around the living stem, holding it to the
dead stem. This resupinate structure is deep Ascomycetes, Helotiales
brown in the center, with a yellow margin. The
stem is constricted at the point of encirclement, Apothecia mostly cup-shaped.
and the sapling usually dies in 2 or 3 years. If the Lachnellula willkommii (Syn. Trichoscyphello
dead stem is removed before girdling, a sunken willkommii). Canker of European larch
canker appears on one side, but this may be over- (▶Dasyscypha).
grown with callus and disappear. Do not leave
severed stems in contact with living seedlings or
saplings in nursery stands. Leptosphaeria

▶ Blights.
Hypoxylon Diapleella coniothyrium (formerly
Leptosphaeria coniothyrium). Canker on thorn-
Ascomycetes, Xylariales less blackberry.
Leptosphaeria coniothyrium (see ▶Diapleella
Perithecia in a pulvinate stroma, often confluent coniothyrium). Canker on thornless blackberry.
and crustose; ascospores with one cell, rarely
two, blackish brown; conidia in superficial layer
on surface of young stroma. Leucostoma
Hypoxylon mammatum Hypoxylon Canker of
poplar. Aspen and large-tooth aspen are most Leucostoma cincta Canker on apple.
Monochaetia 163

black, Japanese, and English walnut. The disease

Macrophoma was first described from Connecticut in 1923, but
evidently was responsible for slow dying of but-
Deuteromycetes, Coelomycetes ternuts long before that. If trees have been previ-
ously weakened, the fungus proceeds rapidly;
Like Phoma, with discrete pycnidia arising otherwise there is the slow advance of a weak
innately, but with much larger spores; conidia parasite. Dead limbs are sprinkled with small,
hyaline; one-celled. black acervuli, looking like drops of ink and
Diplodia tumefaciens (formerly Macrophoma occasionally, in wet weather, developing spore
tumefaciens). Branch Gall Canker of poplar. horns of olive gray conidia. In the teleomorph
Nearly spherical round galls, not over 1 1/2 state, which is rare, perithecia are embedded in
inches in diameter, at base of twigs, which usu- the bark singly or in groups. Mycelium invades
ally die; not serious. bark and wood, with a dark discoloration, and
Macrophoma candollei Associated with Die- grows slowly down a branch to the trunk. When
back of boxwood but apparently saprophytic the latter is reached, the tree is doomed. In final
only. The large black pycnidia are, however, stages trees have a stag-headed effect from loss of
quite striking on straw-colored leaves. leaves.
Macrophoma cupressi Dieback of Italian Control Remove diseased branches promptly,
cypress. cutting some distance below infection; remove
Macrophoma phoradendron Defoliates mis- trees developing trunk cankers; keep the rest
tletoe, but it grows back. growing well with food and water.
Macrophoma tumefaciens (see ▶Diplodia
tumefaciens). Branch Gall Canker of poplar.

Massaria Deuteromycetes, Hyphomycetes

Ascomycetes, Pyrenulales Hyaline mycelium, branched; conidiophores sim-

ple, septate; conidia hyaline, one-celled, pro-
Spores dark, with several cells, oblong-fusiform, duced singly or in clusters.
with mucous sheath. Meria laricis, Dieback and Blight, on western
Massaria platani (see ▶Splanchrorema larch seedlings.
platani). Canker, widespread on branches of
American, London, and California plane trees.
Splanchrorema platani (formerly Massaria Monochaetia
platani). Canker, widespread on branches of
American, London, and California plane trees. Deuteromycetes, Coelomycetes

Acervuli dark, discoid or cushion-shaped, subcu-

Melanconis taneous; conidia several-celled, dark median
cells, hyaline end cells, and a single apical
Ascomycetes, Diaporthales appendage; parasitic.
Monochaetia mali (see ▶Seiridium unicorne).
Perithecia in an immersed black stroma; paraph- Canker, Leaf Spot of apple.
yses present; spores two-celled, light; conidia Seiridium unicorne (formerly Monochaetia
superficial on a stroma. mali). Canker, Leaf Spot of apple. Fungus enters
Melanconis juglandis Walnut Canker, Butter- through deep wounds and grows into wood, then
nut Dieback, widespread on butternut, also on attacks resulting wound callus and produces
164 Cankers and Diebacks

numerous fruiting bodies on exposed wood and Nectria coccinea var. faginata Nectria Beech
callus layer. Killing of successive callus layers Bark Canker on beech in the Northeast. The dis-
results in a canker similar to European apple ease occurs solely in connection with the woolly
canker. The disease is not common enough to beech scale insects (Cryptococcus fagi and C.
be serious. fagisuga), but it has caused high mortality in
Canada, killing 50 % of beech stands; it is epi-
demic in Maine on American beech and is now
Nectria present in much of New England and New York.
The scale nymphs, covered with a woolly
Ascomycetes, Hypocreales, white down, cluster thickly around cracks and
Nectriaceae wounds in bark, often making trunk and branches
appear to be coated with snow. The small yellow
Perithecia bright, more or less soft and fleshy, in larvae establish themselves on the bark in
groups, basal portion seated on a stroma; spores autumn, each inserting its sucking organ, stylet,
two-celled, hyaline or subhyaline (see Fig. 1). into the living bark, which shrinks and cracks.
Nectria cinnabarina Dieback, Twig Canker, Nectria enters through these cracks and kills sur-
Coral Spot, cosmopolitan on hardwoods, most rounding tissue in bark and cambium. When the
common on maples but also found on ailanthus, cells are dead, the insects can no longer obtain
amelanchier, apple, crabapple, apricot, ash, food; therefore, they disappear.
blackberry, chokecherry, beech, birch, elm, hick- White pustules of sporodochia are pushed out
ory, horsechestnut, mimosa, linden, paper mul- through dead bark, bearing elongate, three-to
berry, pear, peach, sophora, locust, and honey nine-celled, slightly curved macroconidia. Red
locust. It may also appear in stem cankers on perithecia, slightly lemon-shaped, appear in clus-
vines and shrubs–ampelopsis, barberry, box- ters on the bark, often so abundant that the bark
wood, callicarpa, cotoneaster, currant, goose- appears red. After ascospores are discharged, the
berry, fig, honeysuckle, kerria, California laurel, upper half of the perithecium collapses and sinks
rose, and syringa. The fungus is widespread as into the lower. The eventual canker is a deeply
a saprophyte. On ornamental trees and shrubs it is depressed cavity surrounded by callus. After the
weakly parasitic, producing cankers around cambium dies, the leaves wilt; the twigs,
wounds and at base of dead branches or causing branches, and roots finally die.
a dieback of twigs and branches. Control Ornamental trees can be sprayed or
On maple, the fungus is more pathogenic, scrubbed to kill the insects. A dormant lime sulfur
killing twigs, small branches, young trees, and spray is very effective. Oil sprays will kill the
girdling larger branches. It is more frequent on scale but may injure beech. Late summer
Norway maple and boxelder; it may also invade spraying for crawlers can supplement the dor-
red, sycamore, Japanese, and other maples. First mant spray.
symptoms are small, depressed, dead areas in Nectria desmazierii (Fusarium buxicola,
bark near wounds or branch stubs. Conspicuous Anamorph). Canker and Dieback of boxwood
flesh-colored or coral pink sporodochia, formed (see Fig. 4).
in dead bark, bear conidia. Later the pustules turn Nectria ditissima Sometimes reported but not
chocolate brown and form pockets, in which confirmed in the United States; reports probably
perithecia are produced. The canker is most refer to Nectria galligena.
common in severely wounded or recently Nectria fuckeliana Canker on fir.
pruned trees. Sapwood has a greenish discolor- Nectria galligena (Cylindrosporium mali,
ation. Open cankers are eventually formed with Anamorph). European Nectria Canker, Trunk
successive rolls of callus. Remove diseased wood Canker, widespread on apple, pear, quince,
and bark, cutting beyond the greenish aspen, beech, birch, maple, hickory, Pacific dog-
discoloration. wood, and various other hardwoods. This is one
Nummularia 165

after leaf fall in autumn with bordeaux mixture to

prevent infection through leaf scars.
Nectria magnoliae Nectria Canker, similar to
the preceding but found on magnolia and


▶ Anthracnose.
Neofabraea perennans (Gloeosporium
Fig. 4 Volutella Blight or “Nectria” Canker on Boxwood perennans) (see ▶Pezicula malicorticis). Peren-
nial Canker of apple, also bull’s-eye rot of fruit.
Pezicula malicorticis (formerly Neofabraea
perennans (Gloeosporium perennans)). Peren-
of the more important diseases of apple and pear nial Canker of apple, also bull’s-eye rot of fruit.
in Europe but is less serious in this country. In The disease is much like northwestern anthrac-
eastern United States it is primarily an apple nose. It often follows after winter injury or starts
disease; on the Pacific Coast it is more common at pruning cuts where aphids congregate, or may
on pear. appear after an application of wound dressing.
Young cankers are small, depressed or flat-
tened areas of bark near small wounds or at base
of dead twigs or branches, darker than the rest of Nummularia
the bark and water-soaked. Older cankers are
conspicuous and somewhat like a target, with Ascomycetes, Xylariales
bark sloughed off to expose concentric rings of
callus. Cankers on elm, sugar maple and birch are Stroma superficial, composed entirely of fungus
usually circular; those on oak irregular; on bass- elements, covered with a conidial layer when
wood elongate, pointed at ends. If the canker is young. Perithecia flask-shaped, embedded in
nearly covered with a callus roll, it indicates that stroma; spores one-celled, dark.
the infection is being overcome. Biscogniauxia marginata (formerly
Small red perithecia are formed singly or in Nummularia discreta). Blister Canker of apple,
clusters on bark or on wood at margin of cankers. crabapple, pear, mountain ash; also reported on
Ascospores discharged during moist weather are serviceberry, birch, elm, magnolia, and honey
disseminated by wind and rain. Creamy-white locust. This is a major apple disease east of the
sporodochia protruding through recently killed Rocky Mountains, especially in Upper Missis-
bark of young cankers produce cylindrical sippi and Lower Missouri River valleys, where
macroconidia and ellipsoidal microconidia. Inva- millions of apple trees have been killed. Large
sion is through bark cracks or other wounds in and small limbs are affected. Cankers are dead
living or dying, but not dead, wood. Infection is areas, up to 3 feet long, mottled with living wood
slow, with annual callus formation; only the and dotted with numerous round cushions of stro-
smallest branches are likely to be girdled. Youn- mata, looking like nailheads. Perithecia, with
ger, more vigorous apple trees receiving nitroge- dark ascospores, are buried in the stromata;
nous fertilizer appear to be more susceptible. hyphae bearing small, light-colored conidia
Control Remove and destroy small branches grow over the surface. The fungus enters through
with cankers. Clean out trunk cankers and cut branch stubs, bark injuries, and other wounds.
back to sound bark; treat with bordeaux paste. Control Avoid especially susceptible varieties
On the West Coast spray pome fruits immediately like Ben Davis. Shape trees early to prevent
166 Cankers and Diebacks

large pruning wounds on older trees; the canker

seldom appears on trees less than 10 years old. Pezicula
Shellac pruning cuts immediately; sterilize tools
between cuts. Ascomycetes, Helotiales
Nummularia discreeta (see ▶Biscogniauxia
marginata). Blister Canker of apple, crabapple, Apothecia similar to Dermatea but lighter.
pear, mountain ash; also reported on service- Pezicula carpinea Bark Canker of hornbeam.
berry, birch, elm, magnolia, and honey locust. Pezicula corticola Superficial Bark Canker and
Fruit Rot, rather common on apple and pears.
Hyaline, one-celled conidia of the Myxosporium
Ophionectria (Scoleconectria) stage are formed in acervuli.
Pezicula pruinosa Canker on branches of
Ascomycetes, Hypocreales amelanchier.

Perithecia red to white, globoid, with a round

ostiole, superficial, paraphyses lacking; spores
needle-shaped to filiform, light colored.
Ophionectria balsamea (see ▶Thyronectria
Ascomycetes, Helotiales
balsamea). Bark Canker of balsam fir.
Ophionectria scolecospora (see
Asci borne in hymenial layers, covered with
▶Scoleconectria cucurbitula). Bark Canker of
a membrane until mature, then splitting;
balsam and alpine firs.
apothecia remain embedded in a stroma; paraph-
Scoleconectria cucurbitula (formerly
yses present; asci clavate.
Ophionectria scolecospora). Bark Canker of bal-
Phacidiella coniferarum (Anamorph,
sam and alpine firs.
Phacidium coniferarum). Phomopsis Disease of
Thyronectria balsamea (formerly
conifers. The fungus is usually saprophytic, but it
Ophionectria balsamea). Bark Canker of balsam
is parasitic on Douglas-fir and larch in Europe
and on living pine in Maine.

Deuteromycetes, Hyphomycetes
▶ Blights.
Conidia in heads; conidiophores unequally verti- Phacidiopycnis boycei (formerly Phomopsis
cillate at tip in whorls; globose conidia formed in boycei). Phomopsis Canker of lowland white
chains, one-celled, hyaline or brightly colored in fir. Branches or main stem of saplings may be
mass; parasitic or saprophytic. girdled and killed; there is often swelling at base
Penicillium vermoeseni Penicillium Disease of of canker where dead tissues join living.
Ornamental palms, serious in southern California The reddish brown needles of dead branches
with symptoms varying according to type of palm. are prominent against living foliage.
On queen palm (Arecastrum or Cocos plumosa) Phacidiopycnis piri (Teleomorph,
the disease is a trunk canker, which may remain Potabiamyces pyri, formerly Phomopsis dis-
inconspicuous for several years but leads to weak- color). Pear branch canker and fruit rot.
ening and breaking of trunk. Infected trees should Phomopsis alnea Canker of European black
be removed at an early stage. On Canary date palm alder.
the disease is a leafbase rot, and on Washington Phomopsis amygdali Branch Dieback on
a bud rot. ▶ Rots. almond.
Physalospora 167

Phomopsis boycei (see ▶Phacidiopycnis

boycei). Phomopsis Canker of lowland white fir. Phragmodothella
Phomopsis discolor (see ▶Phacidiopycnis piri,
see ▶Teleomorph, see ▶Potabiamyces pyri). Ascomycetes, Dothideales
Pear branch canker and fruit rot.
Phomopsis elaeagni (Syn. Phomopsis Asci in locules immersed in groups in
arnoldia). Canker of Russian-olive. a cushionlike stroma; spores hyaline, many-
Phomopsis gardeniae (Teleomorph, Diaporthe celled.
gardeniae). Gardenia Canker, Stem Gall, wide- Dothiora ribesia (formerly Phragmodothella).
spread in greenhouses. Although not reported Dieback,Black Pustule on currant, flowering cur-
until about 1933, this seems to be the most com- rant, and gooseberry.
mon gardenia disease. Symptoms start with Phragmodothella ribesia (see ▶Dothiora
brown dead areas on stem, usually near the soil ribesia). Dieback,Black Pustule on currant,
line. The canker is first sunken, then, as the stem flowering currant, and gooseberry.
enlarges, swollen with a rough, cracked outer
cork. The stem is bright yellow for a short dis-
tance above the canker, a contrast to its normal Physalospora
greenish white. When stems are completely gir-
dled, the foliage wilts and dies; the plant may live ▶ Blights.
a few weeks in a stunted condition. Flower buds Botryosphaeria corticis (formerly
fall before opening. When humidity is high, black Physalospora corticis). Blueberry Cane Canker,
pycnidia on cankers exude yellowish spore in Southeast on cultivated blueberries. The fun-
masses. Entrance is through wounds; spores gus enters through unbroken bark, probably
may be spread on propagating knives. Infection through lenticels, with cankers starting as red-
often starts at leaf joints at the base of cuttings dish, broadly conical swellings, enlarging the
after they have been placed in a rooting medium. next year to rough, black, deeply fissured cankers
Because the cankers may be only slightly visible that girdle the shoots. The portions above cankers
on rooted cuttings, the disease may be widely are unfruitful and finally die. Avoid very suscep-
distributed by the sale of such cuttings. tible varieties like Cabot and Pioneer.
Control Use sterilized rooting medium. Use Botryosphaeria obtusa (formerly
steam for a sand and peat mixture. Destroy Physalospora obtusa (Sphaeropsis malorum)).
infected plants; sometimes it is possible to wait Dieback, Canker of hardwoods, New York
until blooms are marketed. Apple-Tree Canker, Black Rot of Apple. The
Phomopsis lirella (Teleomorph, Diaporthe fungus attacks leaves, twigs, and fruits, is more
vincae). Canker, Dieback of vinca, and important east of the Rocky Mountains, and is
periwinkle. found on many plants, including alder,
Phomopsis lokoyae Phomopsis Canker of ampelopsis, birch, bignonia, bittersweet,
Douglas-fir mostly on saplings in poor sites in callicarpa, catalpa, ceanothus, chestnut, currant,
California and Oregon. Long, narrow cankers, cotoneaster, hawthorn, Japanese quince, maple,
somewhat pointed at ends, develop during the peach, pear, and persimmon. On hardwoods the
dormant season after young shoots are infected. canker is similar to that caused by P. glandicola
If the tree is not girdled during the first season, the on oaks. Limbs are girdled with large areas of
canker heals over. rough bark with numerous protruding black
Phomopsis mali Bark Canker of pear, and apple. pycnidia. For the fruit rot phase of this disease
The bark is rough. ▶ Rots.
Phomopsis padina (Telomorph, Diaporthe Botryosphaeria quercuum (formerly
decorticans). Canker, Twig Blight of sour cherry. Physalospora glandicola (Sphaeropsis quercina,
Phomopsis sp. Shoot Dieback on peach. Anamorph)). Sphaeropsis Canker, Dieback of
168 Cankers and Diebacks

red, chestnut, and other oaks. Shade and orna- scab to form the disease complex known as wil-
mental trees of all ages may be killed. Infection low blight in New England and New York.
may start anywhere through wounds but more Physalospora obtusa (Sphaeropsis malorum)
often on small twigs and branches, passing to (see ▶Botryosphaeria obtusa). Dieback, Canker
larger branches and trunk. Twigs and branches of hardwoods, New York Apple-Tree Canker,
die; leaves wither and turn brown; infected bark Black Rot of Apple.
is sunken, and wrinkled, with small black Physalospora rhodina (see ▶Botryosphaeria
pycnidia breaking through. On larger stems the rhodina). Black Rot Canker of tung in Mississippi
bark has a ridge of callus around the canker, the and Louisiana.
sapwood in this area turning dark with black
streaks extending longitudinally for several
inches. Numerous water-sprouts grow from Phytophthora
below the dead crown. The fungus winters on
dead twigs, producing a new crop of conidia in ▶ Blights.
spring, readily infecting most trees weakened by Phytophthora cactorum Bleeding Canker of
unfavorable environmental conditions. maple, beech, birch, elm, horsechestnut, linden,
Control Prune out diseased portions at least 6 oak, sweetgum, and willow; Crown Canker of
inches below cankers. Fertilize and water to dogwood; Dieback of rhododendron; Trunk Can-
improve vigor. Remove seriously diseased trees. ker of apple, almond, apricot, cherry, and peach.
Botryosphaeria rhodina (formerly Bleeding Canker, first noticed in Rhode Island
Physalospora rhodina). Black Rot Canker of on maple about 1939 and found in New Jersey the
tung in Mississippi and Louisiana. Black, sunken next year, is now present on many trees in the
cankers on trunks, limbs, twigs, and shoots, may Northeast. The most characteristic symptom is
girdle and kill trees. Rogue and burn diseased the oozing of a watery light brown or thick red-
specimens. dish brown liquid from fissures in bark at the root
Glomerella cingulata (formerly Physalospora collar and at intervals in trunk and branches.
miyabeana). Willow Black Canker, accompany- When dry, this sap resembles dried blood, hence
ing scab to form the disease complex known as the name, bleeding canker. Sunken, furrowed
willow blight in New England and New York. cankers are more definite on young trees than on
Starting in leaf blades, the fungus proceeds older trees with rough bark. Symptoms are most
through petioles into twigs; it also causes cankers prominent in late spring and early fall, with trees
on larger stems, followed by defoliation. Pinkish in moist situations most often affected. The fun-
spore masses of the anamorph Gloeosporium gus lives in the soil and advances upward from
state are formed on dead twigs and branch can- a primary root infection. Wilting of leaves and
kers and then short-necked perithecia, which blighting of branches is evidently from a toxin.
overwinter. Remove and destroy dead twigs and Mature trees have fewer, smaller, yellow-green
branches during the dormant period. Spray 3 leaves, and there is an acute dieback of branches.
times with bordeaux mixture, starting just after Reddish-brown areas with intense olive-green
leaves emerge in spring. margins are found in wood extending vertically
Physalospora cortices (see ▶Botryosphaeria from roots to dying branches, marked at irregular
corticis). Blueberry Cane Canker, in Southeast intervals with cavities containing the watery
on cultivated blueberries. fluid.
Physalospora glandicola (Sphaeropsis Control Although there is no real “cure,”
quercina, Anamorph) (see ▶Botryosphaeria injecting trees with Carosel, a mixture of helione
quercuum). Sphaeropsis Canker, Dieback of orange dye and malachite green, has inhibited the
red, chestnut, and other oaks. fungus and neutralized the toxin. In some cases
Physalospora miyabeana (see ▶Glomerella trees recover without treatment. Avoid heavy
cingulata). Willow Black Canker, accompanying feeding; this seems to encourage the spread of
Pseudovalsa 169

disease and causes chronic cases to become Phytophthora cinnamomi Basal Canker of
acute. Maple, particularly Norway maple. Trees have
Crown Canker, collar rot, is the most serious a thin crown, fewer and smaller leaves, and die
disease of dogwood reported in New York, New a year or two after cankers are formed at the base
Jersey, and Massachusetts. The first symptom is of the trunk. Sapwood is reddish brown; the roots
a general unhealthy appearance, with leaves decay. Remove diseased trees. Plant new Norway
smaller and lighter green than normal, turning maples in good soil, well drained, rich in organic
prematurely red in late summer. Leaves may matter; treat injuries at base of trunk promptly.
shrivel and curl during dry spells (normal leaves See under ▶ Rots and ▶ Wilt Diseases for other
often do likewise). Twigs and large branches die, manifestations of this pathogen.
frequently on one side of the tree. The canker Phytophthora syringae Pruning Wound Can-
develops slowly on the lower trunk near the soil ker of almond.
level. Inner bark, cambium and sapwood are
discolored; the cankered area is sunken; the
bark dries and falls away, leaving wood exposed. Plenodomus
Trees die when the canker extends completely
around the trunk base or root collar. The fungus Deuteromycetes, Coelomycetes
lives in the soil in partially decayed organic mat-
ter, and spores are washed to nearby trees. Pycnidia dark, immersed, irregular in shape and
Entrance is through wounds. The disease affects opening irregularly; conidia hyaline, one-celled,
transplanted dogwoods, seldom natives growing oblong; parasitic.
in woods. Aposphaeria fuscomaculans (formerly
Control Transplant carefully, avoiding all unnec- Plenodomus fuscomaculans). Canker on apple.
essary wounds; avoid hitting base with lawnmower, Plenodomus fuscomaculans (see ▶Aposphaeria
by using a wire guard around the tree. It is difficult fuscomaculans). Canker on apple.
to save trees already infected, but cutting out small
cankers and painting the wound with bordeaux
paste is worth trying. If trees have died from Pseudonectria
crown canker, do not replant with dogwoods in
the same location for several years. Ascomycetes, Hypocreales
Rhododendron Dieback, is a disease in which
terminal buds and leaves turn brown, roll up, and Perithecia superficial, blight-colored, smooth;
droop as in winter cold. A canker encircles the spores one-celled, hyaline.
twigs, which shrivel with the terminal portion Pseudonectria rouselliana Nectria Canker of
wilting and dying. In shady locations leaves boxwood, Leaf Cast, Twig Blight. The perithecia
have water-soaked areas, changing to brown, are formed on dead leaves, but the fungus is
zonate spots. Do not plant rhododendrons near thought to be the teleomorph state of Volutella
lilacs, for they are blighted by the same fungus. buxi, which see.
Prune diseased tips well below the shriveled part,
and spray after blooming with bordeaux mixture,
two applications 14 days apart. Pseudovalsa
Trunk Canker of Apple, is an irregular canker
often involving the entire trunk and base of scaf- Ascomycetes, Diaporthales
fold branches, the first outward symptom a wet
area on bark. Trees must be 5-years old or older Perithecia in a stroma; spores ark, with several
for infection. Grimes Golden and Tomkins King cells.
are especially susceptible, often being Pseudovalsa longipes Twig Canker on coast
completely girdled. live oak and white oak.
170 Cankers and Diebacks

Sclerotinia (Syn. Whetzelinia)

Rhabdospora sclerotiorum Basal Canker on Euonymus.
Stem Canker and Wilt on sage.
Deuteromycetes, Coelomycetes

Pycnidia separate, not produced in spots, Septobasidium

erumpent, ostiolate; conidiophores short, simple
conidia hyaline, filiform to needle-shaped, with Basidiomycetes, Septobasidiales
several cells; parasitic or saprophytic.
Rhabdospora rubi (see ▶Septocyta ruberum). All species are on living plants in association
Cane Spot, Canker of raspberry. with scale insects; the combination causes dam-
Septocyta ruberum (formerly Rhabdospora age to trees. Fungus body variable, usually resu-
rubi). Cane Spot, Canker of raspberry. pinate, dry, crustaceous or spongy, in most
species composed of subiculum growing over
bark; a middle region of upright slender or thick
Scleroderris pillars of hyphae supports the top layer, in which
hymenium is formed. Basidium transversely sep-
Ascomycetes, Helotiales tate into two, three, or four cells, rarely one-
celled; basidiospores elliptical, colorless, divided
Apothecia black, opening with lobes, crowded into two to many cells soon after formation, bud-
together or with a stroma, short-stalked; spores ding with numerous sporidia if kept moist. Some
hyaline, elongate, with several cells. species with conidia.
Ascocalyx abietina (formerly Scleroderris The fungus lives by parasitizing scales,
lagerbergii ¼ Gremmeniella abietina). Canker obtaining food via haustoria. The insects pierce
on pine. the bark to the cambium, sometimes killing
Grovesiella abieticola (formerly Scleroderris young trees. The fungus kills a few scales but
abieticola). Canker of balsam fir, on Pacific protects many more in its enveloping felty or
Coast. An annual canker, starting in autumn and leathery covering, a symbiotic relationship.
ceasing when cambium is active in spring, is Spores are spread by scale crawlers and by
formed on twigs, branches, and trunks of sap- birds. Most felt fungi are found in the South,
lings. Only twigs and small branches are girdled, abundant on neglected fruit, nut, or ornamental
and if this does not happen before spring, the trees, rare on those well kept.
wound heals over. Small black apothecia with Septobasidium burtii Felt Fungus on southern
short stalks appear on dead bark. Ascospore hackberry, beech, pear, apple, and peach. This is
infection is through uninjured bark or leaf scars. a perennial growth, with a new ring added to the
Scleroderris abieticola (see ▶Grovesiella patch each summer. Probasidia are formed during
abieticola). Canker of balsam fir, on Pacific the winter, and four-celled basidia in spring.
Coast. Septobasidium castaneum Felt Fungus abun-
Scleroderris lagerbergii Syn. Gremmeniella dant on willow and water oaks, and holly; may
abietina (see ▶Ascocalyx abietina). Canker on injure azaleas. The surface is smooth, shiny,
pine. chocolate brown to nearly black.
Scleroderris lateritium Canker on pine. Septobasidium curtisii Felt Fungus, wide-
spread on many trees in the Southeast, commonly
on sour gum (tupelo) and American ash, also on
Sclerotinia hickory, hawthorn, Japanese quince, and others.
The felt, purple-black throughout, is mounded
▶ Blights. over the insects.
Strumella 171

Septobasidium pseudopedicellatum Felt Fun-

gus, on citrus twigs, sometimes on main stem or Stegonsporium
branches of hornbeam. Surface is smooth, buff-
colored over dark brown pillars. Deuteromycetes, Hyphomycetes

Stegonsporium sp. Maple Canker, Dieback.

Solenia (Henningsomyces) Reported from New Jersey. Large branches die
back with conspicuous flagging. Black tarlike
Basidiomycetes, Aphyllophorales fruiting bodies are formed in cankers.

Fruiting layers erect, cylindrical, formed in

groups, membranous.
Cyphellopsis anomala (formerly Solenia Strumella
(Henningsomyces) anomala). Bark Patch, Can-
ker, widespread on alder. Deuteromycetes, Hyphomycetes
Merismodes ochracea (formerly Solenia
ochracea). Bark Patch of birch, hornbeam, hick- Sporodochia wartlike, gray to black, of
ory, and alder. interwoven hyphae; conidiophores dark,
Solenia (Henningsomyces) anomala (see branches; conidia dark, one-celled, ovoid to
▶Cyphellopsis anomala). Bark irregular.
Patch, Canker, widespread on alder. Conoplea globosa (formerly Strumella
Solenia ochracea (see ▶Merismodes coryneoidea). Strumella Canker of oak, espe-
ochracea). Bark Patch of birch, horn beam, hick- cially the red oak group, also on American
ory, and alder. beech and chestnut, occasional on pignut and
hickories, red maple, and tupelo. Primarily
a forest disease, this canker may become impor-
Sphaeropsis tant on red and scarlet ornamental oaks. Starting
as a yellowish discoloration of bark around a dead
Deuteromycetes, Hyphomycetes branch or other point of infection, the canker
develops into a diffuse lesion or into a target
Pycnidia black, separate or grouped, globose, canker with concentric rings of callus. Whitish
erumpent, ostiolate; conidiophores short; conidia mycelium is present near outer corky bark, and
large, dark, one-celled, ovate to elongate, on fili- the infected portion of the trunk may be flattened
form conidiophores. Some species have or distorted. Target cankers may be up to 2 feet
Physalospora as the teleomorph state. wide and 5 feet long. The small black nodules
Sphaeropsis sapinea Bleeding Canker on pine. bear no spores while trees are living, but after
Sphaeropsis tumefaciens Canker and Gall on death dark brown spore pustules are formed,
Carissa. which blacken with age. New pustules are formed
Sphaeropsis ulmicola Sphaeropsis Canker of yearly. Canker eradication has been unsuccessful
American elm. The disease spreads downward in forest stands. The diseased trees should be
from small twigs to larger branches with removed and utilized before spores can spread
a brown discoloration of wood just under the infection.
bark. Secondary shoots sometimes develop Strumella coryneoidea (see ▶Conoplea
below the cankers. Trees weakened by drought globosa). Strumella Canker of oak, especially
or poor growing conditions are particularly sus- the red oak group, also on American beech and
ceptible. Prune out infected wood, cutting well chestnut, occasional on pignut and hickories, red
below cankers. maple, and tupelo.
172 Cankers and Diebacks

Sydowia Tympanis

Ascomycetes, Dothidiales Ascomycetes, Helotiales, Helotiaceae

Asci usually short, cylindrical, and relatively Ascocarp cup-shaped; sclerotia absent;
numerous, in spherical, ostiolate locules. expiculum usually, if parallel hyphae.
Sydowia polyspora Twig Dieback on fir. Tympanis confusa Canker on pine.

Thyronectria Valsa

Ascomycetes, Hypocreales Ascomycetes, Diaporthales

Stroma valsoid with several perithecia, bright- Many perithecia in a circle in a stroma in
colored; spores muriform, hyaline to subhyaline. bark; flask-shaped with long necks opening to
Thyronectria austro-americana Canker, Wilt the surface; spores hyaline, one-celled, curved,
of honeylocust. Slightly depressed cankers rang- slender.
ing from pinhead size to 1/2 inch grow together Leucostoma cincta (formerly Valsa cincta).
and enlarge to girdle a branch. Underlying wood Perennial Canker of peach, Dieback, also on
is streaked reddish brown for several inches from nectarine. The fungus is apparently infective dur-
the canker, and there is often a gummy exudate. ing the dormant season, entering through
Some trees die, but many survive. wounds, dead buds, leaf scars, and fruit spurs. It
Thyronectria balsamea Canker on fir. forms a canker complex with V. leucostoma and
Thyronectria berolinensis Cane Knot Canker sometimes the brown-rot fungus. It is more com-
of fruiting and flowering currants. mon in northern latitudes than in southern, but is
not important in well-cared-for orchards.
Leucostoma kunzei (formerly Valsa kunzei).
▶Cytospora kunzei.
Trichothecium Leucostoma persoonii (formerly Valsa
leucostoma). Apple Canker, Dieback, Twig
▶ Rots. Blight on apple, apricot, peach, pear, quince,
Trichothecium roseum Canker of rose. plum, cherry, willow, and mountain-ash. The
fungus is a weak parasite entering through
wounds or twigs killed by frost.
Valsa cincta (see ▶Leucostoma cincta). Peren-
Tubercularia nial Canker of peach, Dieback, also on nectarine.
Valsa kunzei (see ▶Leucostoma kunzei).
Deuteromycetes, Hyphomycetes ▶Cytosporakunzei.
Valsa cincta (see ▶Leucostoma cincta). Peren-
Forms bright colored cushions, mostly on wood nial Canker of peach, Die back, also on nectar-
or bark; fine branching conidiophores bearing ine. Valsa kunzei (see ▶Leucostoma kunzei).
small, elipsoidal hyaline conidia. ▶Cytospora kunzei.
Tubercularia ulmea Canker on Russian olive Valsa leucostoma (see ▶Leucostoma
and honeylocust. persoonii). Apple Canker, Dieback, Twig Blight
Volutella 173

on apple, apricot, peach, pear, quince, plum, Volutella buxi Boxwood “Nectria” Canker,
cherry, willow, and mountain-ash. Valsa salicina Volutella Blight. The teleomorph state of the fun-
(Cytospora salicis). Twig and Branch Canker of gus is supposed to be Pseudonectria rouselliana,
willow. which see. As a canker the disease often follows
Valsa sordida ▶Cytospora chrysosperma. after winter injury, with salmon-pink spore pus-
tules on dying twigs, branches, and main stems.
As a blight, the fungus spreads rapidly in moist
Vermicularia weather in summer, attacking healthy twigs when
humidity is high and often discernible at
Deuteromycetes, Coelomycetes a distance by a straw yellow “flag.” On such
yellowing branches the backs of leaves and the
Like Colletotrichum but setae are scattered bark of twigs are both covered with the pinkish
throughout the acervuli, not just marginal; spores spore pustules.
hyaline, globose to fusoid. Control Cut out branches where the bark has
Vermicularia ipomoearum Stem Canker of been loosened by winter ice and snow. Have
morning glory. a yearly “housecleaning,” brushing out accumu-
lated leaves and other debris from interior of
bushes and cutting out all twigs with pink pus-
Volutella tules. If there are signs of disease, follow cleaning
with thorough spraying, from ground up through
▶ Blights. interior of bushes, with lime sulfur.
Club Root

Plasmodiophora and drop. The root system becomes a distorted

mass of large and small swellings, sometimes
Plasmodiophoromycetes, several roots swollen like sweet potatoes, and
Plasmodiophorales sometimes joined in one massive gall. Lateral
and tap roots are scabby and fissured, with rot
This genus, founded on the club root organism, starting from secondary fungi.
has a somewhat doubtful taxonomic position. When diseased roots decompose, small spher-
Formerly considered a slime-mold, one of the ical spores are liberated in the soil; they are capa-
Myxomycetes, then placed in the Chytridiales, ble of surviving there many years between crops.
lowest order of true fungi, it is now placed in In spring, with suitable temperature and moisture,
a separate order, Plasmodiophorales. the resting spores germinate, each becoming
Thallus amoeboid, multinucleate in host cell; a motile swarm spore with a flagellum. This
spores lying free in host cell at maturity; fre- whiplike appendage is soon lost, and the organ-
quently causing hypertrophy; parasitic on vascu- ism becomes amoebalike, moving by protoplas-
lar plants. mic streaming until it reaches a root hair or other
Plasmodiophora brassicae Club Root of cab- root tissue. The plasmodium continues to grow
bage and other crucifers; finger-and-toe-disease, and divide until it reaches the cambial cells, in
on alyssum, brussels sprouts, cabbage, Chinese which it develops up and down the root. The
cabbage, candytuft, cauliflower, hesperis, hon- swelling is produced by division of plasmodia
esty, peppergrass, garden cress, mustard, radish, and of the infected cells. Eventually the
rutabaga, stock, turnip, and western wallflower. multinucleate plasmodium breaks up into many
Club root was present in western Europe as small resting spores, each rounded up around
early as the thirteenth century, but the true cause a single nucleus. They are set free by the millions
was not known until the classic paper of the when the root rots, and are spread in soil clinging
Russian Woronin in 1878. The disease was to shoes or tools and in drainage water, manure,
important in the United States by the middle of and plant refuse. Spores are not seed-borne.
the nineteenth century, and is now present in at Long-distance spread is probably by infected
least 37 states. Losses come from death of the seedlings. Infection takes place chiefly in
plants and also from soil infestation, for suscep- a neutral to acid soil, pH 5.0 to 7.0, at tempera-
tible crucifers cannot be grown again on the same tures below 80  F, and when moisture of soil is
land for several years, unless it is treated. The first above 50 % of its water-holding capacity.
symptom is wilting of tops on hot days, followed Control Inspect seedlings carefully before plant-
by partial recovery at night; affected plants may ing. Dispose of infested crops with caution; rest-
be stunted and not dead; outer leaves turn yellow ing spores passed through animals are still viable.

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_24, 175

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176 Club Root

A long rotation of crops has been recommended, high to use potatoes as a following crop. Most
combined with adding lime to soil, which must be turnip and rutabaga varieties are relatively resis-
applied in large amounts, about 6 weeks before tant to strains of the club root organism present in
the cabbage crop is set. This brings the pH too the United States.

Damping-off is the destruction of young seed- rolfsii, Fusarium equiseti, and Phytophthora
lings by soil organisms. There are two types. may be important on occasion. A synergistic
Pre-emergence damping-off rots the sprouting interaction of Pythium myriotylum, Fusarium
seed before it breaks through the soil; it is recog- solani, and Meloidogyne arenaria causes
nized by bare spaces in what should be uniform damping-off of peanut which has been reported
rows. Such a poor stand may be due to poor in Florida. See under Rots for details.
viability of seed, but more often it is due to soil Also, Caloscypha fulgens (anamorph state,
fungi functioning in cold, wet soils when germi- Geniculodendron pyriforme) causes damping-off
nation is slow. Post-emergence damping-off is of spruce seed, Colletotrichum gloeosporioides of
the rotting or wilting of seedlings soon after papaya, Colletotrichum acutatum of flowering dog-
they emerge from the soil. Succulent stems have wood and Fusarium moniliforme var. intermedia
a water-soaked, then necrotic and sunken, zone at of pine.
ground level; the little herbaceous plants fall over Damping-off is prevented by starting seed in
on the ground or, in woody seedlings, wilt and a sterile medium, such as vermiculite, perlite, or
remain upright. Root decay follows. This type of sphagnum moss, or by treating the soil or the seed
damping-off is most common in greenhouses or before planting. Commercial operators treat soil
outdoors in warm humid weather and where seed- with steam or electricity.
lings are too crowded. Tree seedlings in nursery Seed treatment, the coating of seed with
rows are subject to this type of damping-off, and a protectant dust, is crop insurance. In some sea-
so are perennial flowers started in late summer for sons, good stands can be obtained without it, but
the next year. it scarcely pays to take a chance. Seed disinfec-
Many fungi living saprophytically in the upper tion is used to kill organisms of anthracnose
layers of soil can cause damping-off. Pythium and other specific diseases carried on seed. The
debaryanum, P. mastophorum and Rhizoctonia damping-off organisms are in the soil, not on the
solani are probably most common, but other seed, and coating the seed with a chemical is
species of these two genera and Aphanomyces, intended to kill or inhibit fungi in the soil imme-
Botrytis, Cylindrocladium, Diplodia, Fusarium, diately surrounding the seed and so provide
Macrophomina, Helminthosporium, Sclerotium temporary protection during germination.

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_25, 177

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Dodders are seed plants parasitic on stems and have died. Such tangles are conspicuous in
other parts of cultivated or wild plants. They are weeds along roadsides.
leafless, orange to yellow twining vines, without In ornamental plantings host plants are not
chlorophyll and hence incapable of manufactur- often killed but exhibit stunting and pallor, symp-
ing their own food. They are called love vine, toms of starvation. Minute scales or rudimentary
strangle weed, gold thread, hairweed, devil’s leaves form on the dodder tendrils followed by
hair, devil’s ringlet, pull down, clover silk, and dense clusters of beautiful white blossoms (some-
hell-bind, the last being most appropriate. There times pale pink or yellow), which ripen seed in
are about 40 species in the United States, causing late summer, with as many as 3000 seed being
serious agricultural losses in clovers, alfalfa, and produced on a single plant.
flax, and becoming more and more important in Cuscuta spp. Much of the dodder infesting
gardens on ornamentals and sometimes vegeta- ornamentals is not readily identified as to spe-
bles. Dodders belong to the single genus Cuscuta, cies, but it is widespread on a great many shrubs,
family Cuscutaceae, close to the morning-glory perennials and annuals. It is found very com-
family. monly on chrysanthemum, also strangling any
Dodder seed is grayish to reddish brown, other plant in the vicinity. Many hours may be
resembling small legume seed but roughened spent cleaning up ivy and trumpet-vine, petunias
with three flattened sides. It germinates as ordi- and asters. Dodder is reported on camellias in
nary seed but is synchronized to start a little later the South. It is even a pest of house plants, if
than its host seedlings. The parasite is a slender, field soil has been used for the potting mixture.
yellowish, unbranched thread with the growing Dodder has, however, one virtue for plant
tip circling around in search of support. When it pathologists. It is used as a bridge between
touches the host it twines like a morning-glory plants to carry viruses and MLOs in testing
and puts out little suckers, haustoria, into the their host range.
stem of the victim, after which its original Cuscuta americana on citrus C. californica
connection with the soil dries up (see Fig. 1). on beet.
Although seedlings can live for a few weeks Cuscuta coryli Hazel Dodder. C.
without a susceptible host, they finally die if epithymum. Clover Dodder on legumes.
a connection is not established. Successful para- Cuscuta exaltata on redbud, ilex, and sumac.
sites continue to twine and to spread orange ten- Cuscuta gronovii Common Dodder on but-
drils from one plant to the next, often making tonbush, cucumber, raspberry, members of the
a tangle of matted orange hairs many feet across, potato family, and many garden ornamentals,
with a black region in the center where plants including hedge plants.

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_26, 179

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180 Dodder

Cuscuta pentagona (C. arvensis) Field Dod-

der, widely distributed, most common and seri-
ous east of Mississippi on many cultivated and
wild herbaceous plants.
Cuscuta planifera Littleseed Alfalfa Dodder,
on some legumes in the West.
Control Avoid dodder-infested seed. Commer-
cial seed containing one or more dodder seed per
5-g sample is prohibited entry into the United
States. Many states have laws regulating sale of
infested seed, but it may still be included inad-
vertently in a seed packet. If any contamination
with rough, flat-sided seed is found, do not use
any of the lot. Commercial dealers sometimes
clean infested seed by screening or treating with
an iron powder, which sticks to the rough dodder
seed so it can be drawn out by magnets. Before
breaking new ground for a garden on native sod,
examine it carefully. If dodder is found, burn over
the area, then hoe lightly but repeatedly for sev-
eral weeks to allow buried seed to germinate and
die. When dodder is present on cultivated plants,
Fig. 1 Dodder on Oleander the only thing to do is to remove and burn infested
parts before seed is formed. Pulling off the orange
tendrils is not sufficient. All parts of the plant
attacked must be cut off and burned, for even
Cuscuta indecora Bigseed Alfalfa Dodder on a small fraction of a tendril left twined around
alfalfa from Colorado westward, also on sweet a stem will start growing again.
pea and tomato. A fungus, Colletotrichum destructivum, has
Cuscuta paradoxa on rose, Texas and been found to parasitize dodder and offers
Florida. a slight possibility of biological control.
Downy Mildews

Downy mildews, sometimes called false mil- Basidiophora entospora Downy Mildew of
dews, are Oomycetes, in the order Peronosporales aster, China aster, goldenrod, and erigeron.
and all in the family Peronosporaceae except Aster losses are reported by commercial growers
Phytophthora in the Pythiaceae. They form in the South, but apparently this is not an impor-
mycelium in higher plants and produce sporan- tant garden problem.
giophores that protrude through stomata in great
numbers, their sporangia making white, gray, or
violet patches on the leaves. The downy effect Bremia
distinguishes these mildews from the true or pow-
dery mildews that form white felty or powdery Oomycetes, Peronosporales
The sporangiophores are often branched; they Dichotomous branching of sporangiophores; tips
bear a single sporangium at the tip of each branch enlarged into discs bordered with sterigmata
simultaneously, or successively in Phytophthora. bearing sporangia; swarm spores rare; germina-
Sporangia germinate by swarm spores or with tion usually by a germ tube protruded through an
a germ tube as a conidium. An oospore, resting apical papilla (Fig. 1).
spores with external ridges or knobs, is formed in Bremia lactucae Downy Mildew of lettuce and
an oogonium, large globular multinucleate other composites, endive, cornflower, centaurea,
female cell, after it is fertilized by the antherid- celtuce, escarole, romaine, and various weeds.
ium, a smaller male cell. The oospores are set free First noticed around Boston in 1875, the disease
by weathering and decay of host parts. is serious in greenhouses and in states where
outdoor winter crops are grown. Light green or
yellowish areas on upper surface of leaves are
Basidiophora matched by downy patches on the under surface.
Affected portions turn brown, and leaves die, the
Oomycetes, Peronosporales older ones first. Entrance is through stomata. The
disease is worse in damp, foggy, cool weather
Sporangiophore a single trunk with a swollen (43 to 53  F).
apex from which short branches grow out, each Control The pathogen has numerous physiolog-
bearing a nearly globose sporangium; germina- ical races so that lettuce varieties like Imperial 44
tion by swarm spores; oospore wall not confluent and Great Lakes that are resistant in some local-
with that of oogonium. Mycelium is intercellular, ities may not be so in others. Avoid excessive
haustoria small, knoblike (Fig. 1). irrigation; eliminate crop residue and weeds.

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182 Downy Mildews

Fig. 1 Downy Mildews Fruiting from Stomata on Under- dichotomous branching; Peronospora, sporangia on
side of Leaves. Basidiophora, sporangiophoore with swol- sharply pointed terminal branches; Plasmopara, on obtuse
len apex; Bremia, sporangiophore tip enlarged to a disc, tips; a and a1, sporangium; b1, zoospore

Peronospora arthuri Downy Mildew of gode-

Peronospora tia, clarkia, gaura, and evening primrose.
Peronospora destructor Onion Downy Mil-
Oomycetes, Peronosporales dew, Blight, general on onion, shallot, Vidalia
sweet onion, and garlic. One of the more serious
Mycelium intercellular; haustoria in a few spe- diseases of onion, reported in the United States in
cies short and knoblike, but in most filamentous 1884. All varieties are susceptible, but red onions
and more or less branched. Sporangiophore with have some resistance. Reduction in yield may be
erect trunk two to ten times dichotomously as high as 75 %. The first sign of onion mildew is
branched, with branches somewhat reflexed and the production of conidiophores with a purplish
terminal branches sharp-pointed; sporangia col- tinge a short distance back from tips of older
ored, lacking an apical papilla, germinating from leaves. Leaves turn yellow, wither, and break
an indeterminate point on the side. Oospores over; seedstalks may be infected. Onion mildew
smooth or variously marked, germinating by is sporadic, abundant in years of heavy rainfall.
germ tubes (see Fig. 1). Spores, produced in great numbers in rain or
Peronosclerospora sorghi (formerly when plants are wet with dew, lose vitality
Peronospora sorghi). Downy mildew, on sweet quickly when exposed to sun. Low temperature,
corn. optimum 50  F, favors infection. The fungus
Peronospora antirrhini Snapdragon Downy winters as mycelium in bulbs, in overwintering
Mildew, reported from California, Oregon, Okla- plants in mild areas, or as oospores in soil. Peren-
homa, Pennsylvania, and Maryland. nial onions in home gardens are considered an
Peronospora arborescens Downy Mildew of important source of primary inoculum, but
prickly-poppy on leaves, buds, and capsules. Yel- oospores have been known to survive 25 years
low or light brown blotches on upper leaf surface in soil.
turn dark, with light gray mold on the underside. Control Calred is a resistant variety adapted to
The fungus winters in old plant debris in soil. California. More onion seed is being produced in
Remove and burn infected plants. Use clean seed. Idaho, where dry summers preclude mildew.
Peronospora 183

Peronospora dianthicola Carnation Downy mycelium in seed and oospores in soil. There are at
Mildew, common in California on seedlings. least three races.
Leaves turn pale, curl downward; terminal Peronospora myosotidis Forget-Me-Not
growth is checked, and plants may die. There is Downy Mildew, also on lappula. Pale spots on
a white growth on lower leaf surfaces. upper surface of leaves, with downy growth
Peronospora effusa Spinach Downy Mildew, underneath.
Chard Blue Mold, found wherever spinach and Peronospora oxybaphi, on sand verbena and
swiss chard are grown, absent some seasons, four-o’clock.
nearly destroying the crop in others. Large pale Peronospora parasitica,on garden cress;
yellow spots grow together to cover all or part of P. leptosperma, on artemisia; P. linariae, on
the leaf; lower leaves are infected first, and then linaria; P. lophanthi, on agastache.
the blight is scattered through the plant. Gray to Peronospora parasitica Downy Mildew of cru-
violet mold forms on underside of leaves; some- cifers, general on cabbage, Chinese cabbage,
times the whole plant decays and dries. Initial broccoli, cauliflower, horseradish, radish, turnip,
infection comes from oospores in the soil; it cress, peppergrass, also on sweet alyssum, arabis,
requires humidity above 85 % and a mean tem- arugula, stock, and hesperis. Chief damage is to
perature between 45 and 65  F for a week. Sec- cabbage seedlings or plants grown for seed. Leaf
ondary infection is from conidia. The fungus is an lesions are light green, then yellow, with downy
obligate parasite and does not live over on hosts mold on both sides of the leaf in the widening
other than spinach. yellow zone but not in the dead, shrunken, gray or
Control Plant on well-drained, fertile ground; tan central portion. Secondary fungi often cover
do not crowd; if overhead irrigation is used, dead parts with a black sooty mold. Fleshy roots
water early on sunny days; practice a 2-to of turnips and radishes may be discolored inter-
3-year crop rotation. Resistant varieties such as nally. Warm days and cool nights favor the dis-
Califlay and Texas Early Hybrid 7 are being ease. The pathogen lives between crops in
introduced. perennial plants or winter annuals. There are sev-
Peronospora farinosa (formerly Peronospora eral strains of P. parasitica; one, often reported as
schactii). Beet Downy Mildew, on beet, sugar P. matthiolae, blights stock in greenhouse and
beet, and swiss chard. Inner leaves and seedstalks nursery. Leaves wilt; tender stems and flower
are stunted and killed, covered with violet down. parts are stunted and dwarfed.
The disease appears on the Pacific Coast during Control Avoid crowding plants; keep foliage
the fall rainy season. Oospores can survive in the dry. Spray cabbage seedlings; repeat two or
soil several years. three times a week until plants are set in field.
Peronospora fragariae Strawberry Downy Treat heading cabbage every 6 or 7 days begin-
Mildew.Peronospora grisea, on veronica, a ning 1 to 3 weeks before harvest.
grayish mildew on underside of leaves. Peronospora pisi Pea Downy Mildew. Water-
Peronospora manshurica. Soybean Downy soaked tissue and white growth appear on any
Mildew, general. Yellow-green foliage spots aerial plant part. The mycelium winters in vetch
turn brown, with a grayish mold underneath; stems, fruiting there in spring, and spores are
there may be premature defoliation. The patho- disseminated back to peas. The disease is not
gen winters as mycelium in seed and oospores in important enough for control measures.
soil. There are at least three races. Peronospora potentillae Downy Mildew of
Peronospora grisea on veronica, a grayish mil- agrimony and mock strawberry.
dew on underside of leaves. Peronospora radii Downy Mildew of Margue-
Peronospora manshurica Soybean Downy rite daisy, Argyranthemum frutescens (formerly
Mildew, general. Yellow-green foliage spots turn Chrysanthemum frutescens).
brown, with a grayish mold underneath; there may Peronospora rubi Downy Mildew of black-
be premature defoliation. The pathogen winters as berry, dewberry, and black raspberry.
184 Downy Mildews

Peronospora rumicis Rhubard Downy Mil- Control Use seed grown in the West where mil-
dew. A European disease reported from Califor- dew is not present; plan a 2 to 3-year rotation.
nia on garden rhubarb. Fungus winters in Copper dusts are satisfactory.
rootstalks and grows up into new leaves.
Peronospora schactii (see ▶Peronospora
farinosa). Beet Downy Mildew, on beet, sugar Plasmopara
beet, and swiss chard.
Peronospora sorghi (see ▶Peronosclerospora Oomycetes, Peronosporales
sorghi). Downy mildew, on sweet corn.
Peronospora sparsa Rose Downy Mildew, Sporangiophores with monopodial branches, with
chiefly on roses under glass, rarely outdoors. obtuse tips, arising more or less at right angles;
Young foliage is spotted, leaves drop; flowers haustoria unbranched and knoblike; sporangia
are delayed or unmarketable. Abundant spores (conidia) small, hyaline, papillate, germinating
are produced on undersurface of leaves. To con- sometimes by germ tubes but usually by swarm
trol, keep humidity below 85 % and daytime spores; oospores yellowish brown, outer wall
temperature relatively high. wrinkled, sometimes reticulate, oogonial wall per-
Peronospora statices Downy Mildew on sistent but not fused with oospore wall (see Fig. 1).
statice. Plasmopara acalyphae Acalypha Downy
Peronospora tabacina Blue Mold of tobacco, Mildew.
Downy Mildew; also on eggplant, pepper, and Plasmopara geranii on geranium P. gonolobi
tomato. This is a seedling disease that can be on gonolobus.
controlled by sprays on eggplant and pepper; it Plasmopara halstedii Downy Mildew of bur-
is unimportant on tomato. marigold, centaurea, erigeron, eupatorium,
Peronospora trifoliorum Downy Mildew of gnaphalium, goldenrod, hymenopappus, Jerusa-
lupine, and alfalfa. lem artichoke, ratibida, rudbeckia, senecio,
silphium, verbesina, and vernonia. Zoospores
germinate in soil moisture and invade seedlings
Phytophthora via root hairs; mycelium moving up into stem and
leaves causes early wilting and death. Older
▶ Blights. plants may not die but exhibit a light yellow
Phytophthora phaseoli Downy Mildew of mottling. Sporangiophores project through sto-
lima bean, most important in Middle and North mata on underside of leaves. The fungus winters
Atlantic states, in periods of cool nights, heavy in seed and as oospores in soil.
dews, and fairly warm days. Some seasons it Plasmopara crustosa (formerly Plasmopara
takes 50 to 90 % of the crop; in other years it nivea). Downy Mildew of carrot, parsley, pars-
is of little consequence. The white downy mold nip, and chervil. Yellow spots on upper surface of
is conspicuous on the pod, either in patches or foliage and white mycelial wefts on under surface
covering it completely. The fungus grows turn dark brown with age. The disease is rela-
through the pod wall into the bean, then the pod tively infrequent, important when plants are so
dries, turns black. On leaves the white mycelial crowded they cannot dry off quickly after rain or
weft appears sparingly, but veins are often heavy dew. Control by spacing rows properly.
twisted, purplish, or otherwise distorted. Young Plasmopara nivea (see ▶Plasmopara crustosa).
shoots and flowers are also attacked, bees Downy Mildew of carrot, parsley, parsnip, and
and other insects carrying spores from diseased chervil.
to healthy blossoms. The fungus fruits abun- Plasmopara pygmaea, on anemone, and
dantly on pods, stems, and leaves; spores are hepatica. Fine white mildew covers underside of
splashed by rain. leaves; plants are distorted, stems aborted.
Pseudoperonospora 185

Plasmopara viburni Viburnum Downy Mildew. Toward the end of the growing season thick-
Plasmopara viticola Grape Downy Mildew, walled resting spores, oospores, are produced in
general on grape, also on Virginia Creeper and intercellular spaces of the infected leaves. These
Boston ivy. This is a native disease, endemic in are set free in spring by disintegration of host tissue,
eastern United States, first observed in 1834 on are rain-splashed to other vines, and germinate by
wild grapes. It appeared in France after 1870, production of a short, unbranched hypha bearing
imported with American stock resistant to the a single large sporangium, to start the cycle anew.
Phylloxera aphid, and in a few years had become Control Copper sprays are effective. Apply bor-
as ruinous to the wine industry of Europe as the deaux mixture immediately before and just after
potato blight had been to Ireland. The efficacy of blooming; repeat 7 to 10 days later and possibly
bordeaux was first discovered in connection with when fruit is half grown. Destroy fallen leaves by
this mildew. burning.
In this country downy mildew is most destruc-
tive on European varieties of grape. Pale yellow
spots, varying in form but often nearly circular Pseudoperonospora
and somewhat transparent, appear on upper leaf
surfaces, and a conspicuous white coating Oomycetes, Peronosporales
appears on lower surfaces. The spots turn brown
with age; in dry weather the downy growth is Like Plasmopara but with branches of sporan-
scanty. Young canes, leafstocks, and tendrils giophores forming more or less acute angles; tips
may be infected; flowers may blight or rot; more acute.
young fruits stop growing, turn dark, and dry Pseudoperonospora celtidis Downy Mildew of
with a copious grayish growth. Older fruits have hackberry.
a brown rot but lack the mildew effect. Fruits Pseudoperonospora cubensis Downy Mildew
from diseased vines have less juice; bunches are of cucurbits, destructive to cucumber, musk-
very poorly filled. melon, and watermelon, particularly along the
Initial infection comes from a swarm spore Atlantic seaboard and the Gulf Coast, occasional
stopping on the lower side of a leaf, putting out on gourd, pumpkin, and squash. The disease was
a germ tube and entering through a stoma. In 5 to first noted in 1889 in New Jersey, and in 1896
20 days the mycelium has spread through the leaf destroyed most of the cucumbers on Long Island.
between cells, obtaining food through thin- Irregular yellow spots appear on upper leaf sur-
walled, globular haustoria. The hyphae mass in faces, often on leaves nearest the center of the
compact cushions just beneath the stomata; under hill. The lesion is brown on the opposite side,
humid conditions a few grow out through the covered with a purple growth in rain or dew.
openings and develop into branched conidio- The whole leaf may wither and die, with the
phores (sporangiophores). Each has three to six fruit dwarfed to nubbins and of poor flavor. The
main branches, and they branch again. The ter- fungus does not live in the soil and is not preva-
minal branches end in two to four short, slender lent in the North until July or August. It winters in
sterigmata, each of which produces a single greenhouses or comes up from the South by
multinucleate spore. With moisture, each nucleus degrees. Sporangia are spread by wind and
with adjacent protoplasm is organized into cucumber beetles. The disease is favored by
a swarm spore, motile with two cilia. They high humidity, but temperatures need not be as
swim around for a while, then settle down, absorb cool as for other downy mildews.
their cilia, and put out a germ tube. If they happen Control Resistant cucumbers are of rather poor
to be on the upper side of a leaf, nothing happens; quality. Cantaloupe varieties Texas Resistant
if on the lower surface, the germ tube may reach No. 1 and Georgia 47 combine resistance to
a stoma and start an infection. aphids with resistance to downy mildew.
186 Downy Mildews

Sclerospora farlowii Downy Mildew of Ber-

Sclerospora muda grass, in the Southwest. Short, black, dead
areas prune off tips of leaves without serious
Oomycetes, Peronosporales damage to grass. Tissues are filled with thick-
walled, hard oospores.
Oospore wall confluent with that of oogonium; Sclerospora graminicola on cereals.
sporangiophore typically stout with heavy Sclerophthora macrospora (formerly
branches clustered at apex; mycelium Sclerospora macrospora). Downy Mildew of
intercellular, with small, knoblike, unbranched oats, crazy top of corn, wheat, barley, St.
haustoria; germination by germ tube or Augustinegsrass, Kentucky bluegrass and wild
swarmspores. Common in moist tropic regions grasses. Plants bunch owing to shortening of
on corn, millet, sorghum, and sugar cane. internodes.
Fairy Rings

Several species of mushrooms growing in circles 2 to 12 inches across, white with scattered brown
in lawns and golf greens cause a condition known scales; flesh white; gills green when mature,
as fairy ring, rather common when the soil is quite spores green turning yellow, stem bulbous at
moist and contains a superabundance of organic base with a large ring (annulus). Poisonous,
matter. Less commonly, some of these mush- though other members of this genus, also causing
rooms are responsible for a poor condition of fairy rings, are edible.
other herbaceous plants and of roses. The chief Marasmius oreades Cap 2 inches or less,
symptom in turf is the appearance of continuous convex to plane, thin, tough, withering but not
or interrupted bands of darker green, due to the decaying; gills free from stem; spores white.
fungus mycelium breaking down organic matter Edible.
into products easily assimilated by grass roots. Psalliota (Agaricus) campestris Cap 1 1/2 to
Following the zone of stimulated growth there 3 inches; white, silky, nearly flat; flesh white to
may be a zone of dying grass due to temporary pinkish; gills pink, then brown; spores brownish
exhaustion of nutrients, or to toxic substances purple; stem white, with a ring when young.
from the mushroom mycelium, or because Edible.
a layer has developed that is rather impervious Other Basidiomycetes found on lawns in moist
to water. The green rings are more conspicuous weather include puffballs, which are very good
on underfertilized lawns, and their presence can eating when white and firm inside, and bird’s nest
sometimes be masked by adequate fertilization. fungi, which are tiny cups filled with “seed,”
Breaking off the mushrooms, possibly spiking the resembling a nest of eggs.
sod, is all the control ordinarily recommended.
The following species are merely representative
of the Basidiomycetes found in fairy rings. They are Trechispora
in the order Agaricales, family Agaricaceae.
Cyathus stercoreus (Bird’s Nest Fungus) Basidiomycetes, Aphyllophorales
Fairy Ring on turf.
Lepiota morgani On turf and also in rose Trechispora alnicola Blight, Fairy Ring of
greenhouses, causing poor growth. The caps are Kentucky bluegrass.

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Fruit Spots

Many fruit blemishes are symptoms of rot dis- Helminthosporium

eases and are treated under Rots; others are due to
physiological disturbances; a few others, limited ▶ Blights.
to fruits and known primarily as fruit spots or Helminthosporium papulosum Black Pox on
specks, are included here. apples and pears in eastern states. Fruit spots are
small, sunken, dark, scattered in profusion over
the surface. Blackish papules on bark are
Aureobasidium followed by a pitted or scaly condition. Spray
with sulfur (except at high temperatures).

Yeast-like growth characteristics. Microthyriella

Aureobasidium pullulans Fruit Russet on
apple. Ascomycetes, Hemisphaeriales

Vegetative mycelium lacking; stromata with

Cribropeltis radial structure appearing as black superficial
dots on leaves or stems.
Deuteromycetes, Coelomycetes Microthyriella rubi (see ▶Schizothyrium
pevexiguum). Fly Speck of pome fruits, general
Brown mycellium, branches profusely; black, on apple, also on pear, quince, citrus fruits,
irregularly circular pycnidia; simple, hyaline, banana, Japanese persimmon, plum, blackberry,
clavate conidiophores; pale, oblong, straight or raspberry, and grape.
slightly curved conidia. Schizothyrium pevexiguum (formerly
Cribropeltis citrullina Fly Speck of water- Microthyriella rubi). Fly Speck of pome fruits,
melon fruits. general on apple, also on pear, quince, citrus
fruits, banana, Japanese persimmon, plum, black-
berry, raspberry, and grape. The pathogen has
Zygophiala long been recorded as Leptothyrium pomi, but
this is apparently a misconception. The anamorph
▶ Blotch Diseases. state is Zygophiala jamaicensis, originally iso-
Zygophiala jamaicensis (Schizothyrium pomi). lated from banana and recently reported as caus-
Fly Speck on apple. ing a greasy blotch of carnations. Flyspeck is

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190 Fruit Spots

often associated with sooty blotch on apples, but quince, most prevalent in northeastern states.
the two diseases are distinct. Flyspeck looks like Spots appear on fruits in July or early August,
its name, groups of 6 to 50 very small, slightly deeper red on the colored face of apples, darker
elevated, superficial black dots connected with green on the lighter surface. They are irregular,
very fine threads. Spots may extend entirely slightly sunken, more abundant near the calyx
around blackberry canes and shoots. end of the fruit, usually with centers flecked
with black. The symptoms on quince are more
of a blotch than a definite spot.

▶ Anthracnose. Rhodotorula
Mycosphaerella pomi Brooks Fruit Spot,
Phoma Fruit Spot. Quince Blotch, of apple and Rodotorula glutinis Fruit Russet on apple.

Galls are local swellings, hyperplastic enlarge- exposing the spore-bearing layer. There is sel-
ments of plant tissue due to stimulation from dom more than one diseased shoot on a stem,
insects, bacteria, fungi, viruses, and occasionally and not many on the whole bush; so the disease
physiological factors. Crown gall, a common and does not cause serious damage.
serious problem, is discussed under Bacterial Control Handpicking of affected parts,
Diseases. Cedar galls are treated under Rusts. searching carefully for diseased leaves at base
See ▶ Black Knot for hypertrophy of plum of new growth, removing them before spores
branches. are formed, keeps sporadic infection at
a minimum. Spraying with a low-lime bordeaux
may be effective but is seldom necessary.
Exobasidium Exobasidium oxycocci Cranberry Rose Bloom,
Shoot Hypertrophy on cranberry, and manza-
Basidiomycetes, Exobasidiales nita. The disease appears in cranberry bogs soon
after water is removed in spring. Bud infection
Mycelium intercellular with branched haustoria results in abnormal lateral shoots with enlarged,
entering host cells; basidia extend above the layer swollen, pink or light rose distorted leaves that
of epidermal cells much like the layer of asci in somewhat resemble flowers. Excessive water
Taphrina; each basidium bears two to eight supply promotes the disease. Remove water
basidiospores. Species cause marked hypertro- early in spring. If necessary, spray with bordeaux
phy in the Ericaceae. mixture.
Exobasidium vaccinii Leaf Gall, widespread on Exobasidium rhododendri Rhododendron
flame azalea. Leaf Gall. Large vesicular galls, especially on
Exobasidium burtii Leaf Gall, Yellow Leaf Rhododendron catawbiense and R. maximum.
Spot on azalea and rhododendron. Exobasidium symploci Bud Gall on sweetleaf.
Exobasidium camelliae Camellia Leaf Gall on Exobasidium uvae-ursi Shoot Hypertrophy of
camellia in the Southeast, more common on bearberry.
sasanqua than on japonica. Symptoms are Exobasidium vaccinii Azalea Leaf Gall, Red
a striking enlargement and thickening of leaves Leaf Spot, Shoot Hypertrophy of and romeda,
and a thickening of stems of new shoots. Dis- arbutus (A. menziesii), bearberry, blueberry (fruit
eased leaves are four or more times as wide and green spot), box sandmyrtle, chamaedaphne,
long as normal leaves, very thick and succulent. cranberry, farkleberry, huckleberry, ledum,
Color of the upper surface is nearly normal, but leucothoë, manzanita, and rhododendron. On
the underside is white with a thin membrane that azaleas and other ornamentals the galls are blad-
cracks and peels back in strips or patches der-shaped enlargements of all or part of a leaf,

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192 Galls


▶ Rots.
Fusarium decemcellulare (Teleomorph, Nectria
rigidiuscula). Gall on midge.


Deuteromycetes, Hyphomycetes

Sporodochia erumpent, pale olive-green, cush-

ion-shaped; similar to Kutilakesopsis but differs
by having larger two-celled conidia; teleomorph
state is Nectriella.
Kutilakesa pironii Stem and Leaf Gall, Can-
kers on croton, zebra plant, and Clorodendron.

Fig. 1 Azalea Leaf Gall Nocardia

sometimes a flower bud (see Fig. 1). They are
white or pink, soft and succulent when young, Related to bacteria with mycelial filaments
brown and hard with age. This is seldom a serious breaking up into rod forms.
disease but in wet seasons, particularly in the Nocardia vaccinii Blueberry Bud-Proliferating
South, and in shaded gardens, the number of Gall, first observed in Maryland in 1944,
galls may become rather alarming. On cran- described as a new species in 1952. Galls, similar
berries and blueberries the gall is a small, round, to crown gall, are formed at the soil line. Abnor-
red blister in the leaf, with spores packed in mal buds abort at an early stage or grow into weak
a dense layer on the underside. The fungus is shoots, 1 to 6 inches high, forming a witches’
systemic in blueberries, fruiting on the leaves in broom effect.
June and July.
Control Handpick and destroy galls as they
appear. Spraying is seldom required for cran- Phoma, Phomopsis
berries and other fruits.
Exobasidium vaccinii-uliginosi Shoot and ▶ Blights.
Leaf Gall, Witches’ Broom of rhododendron, Phoma sp. or Phomopsis sp Stem Gall on win-
manzanita, and mountain heath. An excessive ter jasmine, privet, forsythia, and rose, at
number of twigs is formed on infected branches. scattered locations. Both pathogens have been
Leaves are yellowish white covered with a dense reported causing roundish, rather rough stem
mealy fungus growth. The mycelium penetrates enlargements on ornamentals. It has not been
the whole plant so that it is wiser to remove the determined whether more than one fungus is
shrub than to attempt remedial measures. involved.
Synchytrium 193

delphinium, geum, golden-glow, marsh-mari-

Plasmopara gold, and viola. Pick off and burn affected parts.
Synchytrium endobioticum Potato Wart,
▶ Downy Mildews. Black Wart of potatoes, a warty hypertrophy of
Plasmopara halstedii Basal Gall on sunflower. tubers. A European disease wart was found in
1918 in backyard gardens in mining towns of
Pennsylvania, Maryland, and West Virginia. Dis-
Protomyces eased tubers had apparently been brought in by
immigrants. A strict quarantine was placed on
Archiascomycetes, Taphrinales infested districts, and there has been no spread
to commercial potato fields. The disease shows as
Protomyces gravidus Stem gall on ragweed. prominent outgrowths or warts originating in the
Protomyces macrosporus Leaf gall on hedge eyes, varying from the size of a pea to that of the
parsley (Torilis sp.). tuber itself. Numerous yellow sporangia are
released into the soil by decay of the malformed
tissue. The disease, which may affect other spe-
Sphaeropsis cies of Solanum, is spread by contaminated soil or
infected tubers. Buds and adventitious shoots of
▶ Cankers and Diebacks. tomato are infected below the soil line.
Sphaeropsis tumefaciens Canker and Gall on Control By 1953 potato wart had been eradi-
Carissa. cated from more than half of the 1,112 infested
gardens in Pennsylvania. The plan called for
applying copper sulfate the first year, keeping
Synchytrium the land clean and cultivated, applying lime the
next year, growing vegetables the third year, and
Chytridiomycetes, Chytridiales going back to potatoes the fourth year to test
Mycelium lacking; thallus converted into a soros Synchytrium vaccinii Red Leaf Gall on cran-
with a membrane, at maturity functioning in entirety berry, azalea, chamaedaphne, gaultheria, and
as a resting sporangium or divided to form many ledum, from New Jersey northward. On cran-
sporangia in a common membrane; zoospores with berry the disease appears just before blossoms
one cillum at posterior end. Various species cause open. Buds, flowers and young leaves are covered
excrescences on leaves and fruit; potato wart. with small, red, somewhat globular galls about
Synchytrium anemones Leaf Gall, Flower Spot the size of birdshot; affected shoots bear no fruit.
of anemone and thalictrum. Flowers are spotted, The disease is erratic in appearance but is most
distorted, dwarfed, and may fall. Red spots are frequent in bogs that have excessive or uneven
formed on leaves and stems. water supply.
Synchytrium aureum Red Leaf Gall, False Rust Synchytrium sp Stem Gall on castor bean, in
on many plants, 130 species in widely separated Texas. Small red galls on stems, petioles, and
genera, including calypha, artemisia, clintonia, leaves of seedlings.
Leaf Blister and Leaf Curl Diseases

A single genus, Taphrina, is responsible for most undersurface, convex and yellow on the upper
of the hyperplastic (overgrowth) deformities surface. Individual blisters are 1/4 to 1/2 inch
known as leaf blister, leaf curl, or, occasionally, across but often become confluent, causing the
as pockets. leaf to curl. Ascospores are borne on the surface
of the blistered area. The disease is most serious
in a cool wet spring.
Taphrina Control A single dormant eradicant spray,
before the buds swell, controls the disease; later
Archiascomycetes, Taphrinales sprays are ineffective.
Taphrina carnea Birch Red Leaf Blister.
Parasitic on vascular plants, causing hypertrophy. Taphrina castanopsidis California Chinquapin
Asci in a single palisade layer, not formed in Leaf Blister.
a fruiting body; hyphal cells become thin-walled Taphrina communis Plum Pockets, common
chlamydospores; on germination the inner spore on American plums; T. pruni, on European spe-
protrudes from the host and is cut off by a septum cies, not in United States; T. prunisubcordata,
to form an eight-spored ascus, which may in western United States. Leaves, shoots and
become many-spored by budding or the fruits become puffy and enlarged into reddish or
ascospores. white swollen bladders. Fruits are sometimes ten
Taphrina spp Maple Leaf Blister. Leaves after times the size of normal plums. Most garden
expanding in spring show dark spots, shrivel, and plums are of foreign origin and not susceptible
fall. The disease may be locally epidemic; it is to the American species of Taphrina. Bordeaux
more common in shaded locations. mixture applied in spring before flower buds open
Taphrina aceris Western Maple Leaf Blister. gives satisfactory control.
Taphrina aesculi Leaf Blister of California Taphrina coryli Hazelnut Leaf Blister.
buckeye; yellow turning to dull red; witches’ Taphrina deformans Peach Leaf Curl, general
broom formed. on peach, also on nectarine and almond but not on
Taphrina australis American Hornbean Leaf apricot. This is an old disease, known in the
Curl. United States for well over a century but not
Taphrina caerulescens Oak Leaf Blister on var- quite so important since 1900, when a control
ious oak species, with red oak particularly sus- was worked out. Young leaves are arched and
ceptible but often defoliating and sometimes reddened, or paler than normal as they emerge
killing water, willow, laurel, and live oaks in the from the bud, then much curled, puckered, and
South. Blisters start on young partially grown distorted, greatly increased in thickness (Fig. 1).
leaves as gray depressed areas on the Any portion or the entire leaf may be curled,

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196 Leaf Blister and Leaf Curl Diseases

Fig. 1 Peach Leaf Curl;

deformed leaf; palisade
layer of asci formed on
curled portion; germinating

and one or all leaves from a bud. The leaves often Taphrina japonica (T. macrophylla). Leaf Curl
look as if a gathering string had been run along on red alder. Young leaves are enlarged to several
the midvein and pulled tight. Leaves may drop, times normal size and curled. They dry up after
lowering vitality of tree, with partial or total ascospore discharge, and a new crop of healthy
failure to set fruit, and increasing chances of leaves is formed.
winter injury. Young fruits may be distorted or Taphrina populina Leaf Blister, Yellow on
cracked. Defoliation for several seasons kills tree poplar.
outright. Taphrina populina Poplar Yellow Leaf Blister.
The fungus has no summer stage, and the asci Conspicuous blisters, small to large, an inch or
are formed not in a fruiting body but in a layer more in diameter, are brilliant yellow on the
over infected surfaces, giving them a silvery concave side when the asci are fully developed;
sheen. Before leaves fall, ascospores are later the color changes to brown.
discharged from this layer, and land on bark or Taphrina robinsoniana, T. occidentalis,
twigs and bud scales, there to germinate by bud- T. alni Catkin Hypertrophy of alder. Scales of
ding into yeastlike spores, which remain viable catkins enlarge and project as reddish curled
over winter, sometimes for 2 years. In spring they tongues covered with a white glistening layer.
are washed by rain to opening leaf buds. Infection can be reduced with a lime sulfur spray.
Control One spray during the dormant season Taphrina sacchari Maple Brown Leaf Blister.
gives effective control. This is best applied just Taphrina ulmi Elm Leaf Blister. Very small
before the buds swell, but can be done any time blisters on elm leaves. Dusting nursery trees
after leaf fall in autumn when the temperature is with sulfur has helped.
above freezing. Applications after the buds swell Taphrina weisneri Cherry Witches’ Broom,
have little effect. Leaf Curl on wild and cultivated cherries, T.
Taphrina faulliana Leaf Blister of Christmas flavorubra, on sand cherry; T. flectans, on west-
fern; T. filicina, on sensitive fern; T. ern wild cherry; T. farlowii, leaf curl and fruit
struthiopteridis, on ostrich fern. pockets on eastern wild cherry; T. confusa, on
Taphrina flava Yellow Leaf Blister of gray and chokecherry; T. thomasii, witches’ broom of
paper birches in northeastern states. cherry-laurel in California.
Leaf Scorch

According to the dictionary scorching means to Curvularia

heat so as to change color and texture without
consuming. Sometimes leaves are literally ▶ Blights.
scorched in summer heat, and sometimes symp- Curvularia sp Leaf Scorch on pecan.
toms caused by fungi resemble those of a heat
scorch. This section includes some of the latter.

Ceratocystis ▶ Blackspot.
Diplocarpon earlianum Strawberry Leaf
▶ Cankers and Diebacks. Scorch, general where strawberries are grown
Ceratocystis paradoxa Black Scorch, Bud but more prevalent in the South. Dark purplish
Scorch, Heart Rot of coconut, Canary, Washing- spots about 1/4 inch in diameter are scattered
ton, and Guadaloupe palms, also causing profusely over upper surface of leaves in all
a pineapple disease in the tropics. The most strik- stages of development. Later the spots enlarge
ing symptom is a black, irregular, necrotic con- to scorch wide areas of the leaf, and black
dition of the leaf stalk. The tissues look as if they fruiting bodies give a “tar spot” appearance.
had been burned, whence the name black scorch. Scorch spots always lack the white centers so
Furled pinnae of leaf fronds show pale yellow characteristic of Mycosphaerella leaf spot on
spots with broad margins that later converge and strawberry. Lesions are found on petioles, sto-
turn black; infection spreads rapidly, and in lons, and fruit stalks as well as leaves. If the fruit
severe cases the heart leaves dry up. The heart stems are girdled, flowers or young fruits die.
rot discolors trunk tissues and rots the pithy mate- Rarely the disease appears on green berries
rial between cells. Infection is through wounds as a superficial red or brown discoloration
during periods of relatively high humidity, or and flecking. Spores, produced in quantity in
through roots, or sometimes through uninjured acervuli on lesions, are distributed by birds,
fruit strands, petioles, or pinnae. Palms with vital- insects, and pickers on tools and clothing. The
ity lowered, as when the normal crown of leaves fungus winters in old leaves. Teleomorph and
has been reduced but the water supply to the anamorph states are both produced in spring,
leaves is not reduced, are most susceptible. and repeated infections occur throughout the
Control Destruction of infected parts seems to summer in moist weather.
be the chief control measure. It is easier to bury Control Remove all old leaves when setting
than to burn palm trunks. plants in spring. Spray with bordeaux mixture at

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198 Leaf Scorch

10-day intervals, starting in January in Louisiana,

late February in North Carolina. Fairly resistant Septoria
varieties include Catskill, Midland, Fairfax,
Howard 17, Blakemore, Southland. ▶ Blights.
Septoria azaleae Azalea Leaf Scorch, Leaf
Spot. Small, yellowish, round spots enlarge irreg-
Epicoccum ularly, turn reddish brown, with dark brown
centers. Leaves fall prematurely; black fruiting
▶ Leaf Spots. bodies are produced in fallen leaves. The disease
Epicoccum sp Leaf Scorch on pecan. is most severe in greenhouses in fall and winter
and under high humidity.

▶ Rots.
Fusarium sp Leaf Scorch on pecan. Deuteromycetes, Coelomycetes

Pycnidia dark, separate, superficial, or erumpent,

Hendersonia globose, ostiolate; conidiophores short; conidia
hyaline, typically with three or more cells, cylin-
Deuteromycetes; Coelomycetes drical to elliptical; parasitic or saprophytic.
Stagonospora curtisii Narcissus Leaf Scorch,
Pycnidia dark, separate, globose, ostiolate, Red Blotch of Amaryllis, Red Leaf Spot,
immersed then usually erumpent; conidia dark, Red Fire Disease, also on crinum, eucharis,
several-celled, elongate to fusoid; saprophytic or hymenocallis, leucojum, nerine, sternbergia,
parasitic. vallota, and zephyranthes.
Hendersonia opuntiae Scorch, Sunscald, com- Leaf tips of narcissus are blighted for 2 or 3
mon and serious on prickly pear cactus inches as in frost injury and separated off from
(Opuntia). Segments turn reddish brown and healthy basal portions of leaves by a definite margin
die; centers are grayish brown and cracked. or yellow area. Spores formed in pycnidia in the
dead area furnish inoculum for secondary infection,
which consists of lesions in lower portions of leaves,
Pestalotia minute water-soaked or yellowish spots becoming
raised, scabby, and reddish brown. Flower stalks
▶ Blights. may be spotted; brown spots appear on petals.
Pestalotia sp Leaf Scorch on pecan. Bulbs suffer loss in weight due to killing of foliage
a month or two before normal dying down. All types
may be infected but the most susceptible varieties
Pseudopezicula are in the Leedsii and Polyanthus groups. The fun-
gus was described on narcissus in 1878 but was not
Ascomycetes, Helotiales considered a threat to it, nor was it known to be
connected with amaryllis red blotch before 1929.
Hyaline, gelatinous apothecia containing paraph- On amaryllis or hippeastrum red spots are
yses and 20–80 asci; asci contain four reniform, formed on leaves, flower stems, and petals. On
binucleate ascospores; five-spored asci rarely foliage the spots are bright red to purplish, small
observed. at first but often increasing to 2 inches. Leaf or
Pseudopezicula tetraspora Leaf Scorch of flower stalks are bent or deformed at the point of
grapevines. attack. This disease should not be confused with
Stagonospora 199

“red disease” caused by mites. The spores are var- Control Treat suspected narcissus bulbs before
iable in size and number of cells, one to six. They planting. Control secondary infection in the field
are embedded in a gelatinous matrix and are dis- with bordeaux mixture. Discard seriously dis-
seminated in rain. The fungus apparently winters in eased amaryllis bulbs; remove infected leaves
or on bulbs, infecting new leaves as they grow out and bulb scales; avoid syringing and heavy
in spring. watering.
Leaf Spots

Leaf spots are the most prevalent of plant dis- expensive proposition. If the budget is limited,
eases, so common we seldom notice them, and it is more important to have an elm sprayed for
rightly so, for if we should attempt to control all elm leaf beetles, which cause defoliation every
the miscellaneous leaf spots that appear in a small season, than for elm black spot, which may be
suburban garden in a single season, we would serious in only one year out of three or four.
quickly go mad. A typical leaf spot is a rather When it comes to rose blackspot (no relation to
definitely delimited necrotic lesion, often with elm black spot), weekly protection with
a brown, sometimes white, center and a darker a fungicide is necessary, but to save labor it can
margin. When the spots are so numerous they be combined with insecticides.
grow together to form large dead areas, the dis-
ease becomes a blight, or perhaps a blotch, or
scorch. Certain types of lesions are called Actinothyrium
anthracnose, spot anthracnose, blackspot. All of
these have been segregated out in their different Deuteromycetes, Coelomycetes
sections. What is left is a very large collection of
names. Pycnidia superficial, globose, with a more or less
The genus Septoria, for instance, has about fimbriate shield; spores filiform, hyaline.
1000 species, Mycosphaerella 500, Cercospora Actinopelte dryina (see ▶Tubakia dryina).
400, chiefly identified by the hosts on which they On oak.
appear. Cercospora beticola is so named because Actinothyrium gloeosporioides (see
it causes a leaf spot of beet, C. apii for its celery ▶Tubakia dryina). On oak. Leaf Spot on
host. Species recorded in this country as causing sassafras.
a definite disease are listed under their respective Tubakia dryina (formerly Actinopelte dryina).
hosts. They are not repeated here unless the leaf On oak. Very small dark spots between veins.
spot is of some importance or there is some useful Conspicuous in midsummer but not serious.
information that can be added to the name. Tubakia dryina (formerly Actinothyrium
Most leaf spot diseases flourish in wet seasons. gloeosporioides). Leaf Spot on sassafras.
A comparative few may be important enough to
call for control measures other than general san-
itation. Adequate protection usually means sev- Alternaria
eral applications of fungicides, and the cost of
spraying trees and shrubs must be balanced ▶ Blights.
against the expected damage. Calling in a tree Alternaria alternata Leaf Spot of
expert with high-pressure apparatus is often an Calathea spp.

R.K. Horst, Westcott’s Plant Disease Handbook, DOI 10.1007/978-94-007-2141-8_33, 201

# Springer Science+Business Media Dordrecht 2013
202 Leaf Spots

Alternaria alternata (formerly Alternaria holes; there is more or less defoliation. The fun-
fasciculata). Leaf Spot on rose-acacia and gus is sometimes secondary following bacterial
asclepiodora. infection or midge infestation. Rake up and burn
Alternaria alternata (formerly Alternaria fallen leaves.
tenuis). Leaf Spot of magnolia, hibiscus, clarkia, Alternaria chrysanthemi (see ▶Alternaria
and many ornamental and other hosts. The fungus leucanthemi). Leaf Spot on shasta daisy, and
is a general saprophyte and an occasional weak Canada thistle.
parasite. It discolors beet, chard, and spinach Alternaria citri Cherry Leaf Spot, occasional,
seed. more often a rot of citrus fruits. ▶ Rots.
Alternaria angustiovoidea Leaf Spot and Alternaria fasciculata (see ▶Alternaria
Blight of leafy spurge. alternata). Leaf Spot on rose-acacia and
Alternaria brassicae (with large spores) and asclepiodora.
A. brassicicola (with small spores). Black Leaf Alternaria leucanthemi (formerly Alternaria
Spot of crucifers, cabbage, Chinese cabbage, chrysanthemi). Leaf Spot on shasta daisy, and
collards, turnip, garden cress, mustard greens, Canada thistle.
radish, and horseradish; Head Browning leaf Alternaria longipes Brown Spot of tobacco,
and pod spot of cauliflower; Damping-off, Wire- including ornamental flowering tobacco. Small
stem of seedlings. spots on lower leaves rapidly enlarge and turn
Seedlings are subject to pre-or post-emergence brown. The fungus winters on old stalks, which
damping-off, with dark brown to black sunken should be removed and burned.
spots on cotyledons, narrow dark spots on stems, Alternaria oleracea (see ▶Alternaria
followed by wire-stem, a blackening toward the brassicicola). Cabbage Leaf Spot, occasional
base. Leaf spots are small, circular, yellowish, on crucifers.
enlarging in concentric circles with a sooty black Alternaria panax Leaf Spot of schefflera,
color from the spores. In storage the spots unite to Dizygotheca, and Tupidanthurs.
form a moldy growth over the entire leaf. On seed Alternaria passiflorae Brown Spot of passion
pods, spots are purplish at first, later brown; in flower. Minute brown leaf spots, enlarging to an
moist weather entire pods may be infected. inch across are concentrically zoned with various
Cauliflower infection is a browning of the head, shades of brown. Dark green water-soaked spots
starting at the margin of an individual flower or on fruit turn brown; the fruit shrivels, but the
cluster. Spores are blown, splashed by tools, spots stay firm.
spread on feet of men and animals. Seed bears Alternaria polypodii Fern Leaf Spot. Brown,
spores externally, mycelium internally. Wounds circular to ovate, concentrically zonate spots
are not necessary for infection. are formed along margins of fronds. Chains of
Control Hot water treatment of seed, 122  F for spores are spread by syringing or air currents.
30 min, is fairly effective. Use long rotation for Keep foliage dry; remove and burn diseased
cauliflower, avoiding all other crucifers in inter- leaves.
mediate years. Alternaria raphani Radish Leaf Spot. Yellow
Alternaria brassicicola Leaf Spot on Thlaspi. spots with black sporulation, often with centers
Alternaria brassicicola (formerly Alternaria dropping out. Also occurs on turnip.
oleracea). Cabbage Leaf Spot, occasional Alternaria sonchi Leaf Spot of lettuce, esca-
on crucifers. Has been confused with role, endive, and chicory.
A. brassicicola. Alternaria tagetica Leaf Spot of marigold.
Alternaria catalpae Catalpa Leaf Spot, wide- Alternaria tenuis (see ▶Alternaria alternata).
spread in rainy seasons. Small, water-soaked Leaf Spot of magnolia, hibiscus, clarkia, and
spots, up to 1/4 inch, appear over the leaf; they many ornamental and other hosts.
turn brown and sometimes drop out leaving shot Alternaria tenuissima Leaf Spot on blueberry.
Ascochyta 203

Alternaria tenuissima (formerly Alternaria

tomato). Nailhead Spot of tomato, a leaf, stem, Aristastoma
and fruit spot. On leaves and stems the disease is
much like early blight (see ▶A. solani under Deuteromycetes, Coelomycetes
Blights) with small dark brown spots with
yellow margins. But on fruit the disease is quite Pycnidia brown, globose, erumpent, separate,
different. Very small tan spots, 1/16 to 1/8 inch in with dark brown setae near ostiole;
diameter, become slightly sunken, with grayish conodiophores short, simple; conida hyaline,
brown centers and darker margins. Spores pro- several-celled.
duced abundantly on fruit and foliage are spread Aristastoma oeconomicum Zonate Leaf Spot
by winds and splashing rain. Treat seed and spray of cowpea, kidney bean.
as for early blight. Varieties Marglobe, Pritchard, Aristastoma sp. Leaf Spot on desert-rose.
Glovel, and Break O’Day are quite resistant to
nailhead spot. The same fungus causes ghost spot
of apple. Ascochyta
Alternaria tomato (see ▶Alternaria
tenuissima). Nailhead Spot of tomato, a leaf, ▶ Blights.
stem, and fruit spot. Ascochyta abelmoschi (possibly identical with
Alternaria sp. Leaf Spot of schefflera, and A. phaseolorum). Leaf Spot, Pod Spot, Stem
umbrella tree. Spot of okra. Dark, small, water-soaked spots
slowly enlarge, turn brown, with many large
black pycnidia in concentric rings in dead tissue.
Amerosporium Young okra pods are severely infected, and the
mycelium grows into the seed.
Deuteromycetes, Coelomycetes Ascochyta althaeina (see ▶Phoma exigna).
Leaf Spot of hollyhock, rose-mallow.
Pycnidia superficial, discoid to cupulate, hairy; Ascochyta armoraciae Leaf Spot of horse-
spores one-celled, hyaline. radish.
Amerosporium trichellum (see Ascochyta aspidistrae Aspidistra Leaf Spot.
▶Colletotrichum trichellum). Leaf Spot and Large, irregular pale spots on leaves.
Stem Spot on English ivy. Ascochyta asteris (see ▶Phoma exigna). Leaf
Colletotrichum trichellum (formerly Spot of China aster.
Amerosporium trichellum). Leaf Spot and Stem Ascochyta boltshauseri (see ▶Stagonos-
Spot on English ivy. In some cases stems are poropsis hortensis). Leaf Spot, Pod Spot of
girdled, causing collapse and death. beans, on snap, kidney, lima, and scarlet runner
beans, reported in Oregon.
Aschochyta Leaf Spot on big bluestem, little
Annellophora bluestem (both species of And ropogon and on
Deuteromycetes, Hyphomycetes Ascochyta cheiranthi Leaf and Stem Spot of
wallflower. Grayish spots up to 1/2 inch
Conidiophores brown, simple, slender, elongat- long, may girdle stems. Leaf spots are circular
ing by successive proliferations through conidial to elongate, brown with darker brown
scars; conidia brown, multiseptate, obclavate to margins. Dark pycnidia contain hyaline, two-
fusoid. celled spores. Leaves wilt and fall; potted plants
Annellophora phoenicis Leaf Spot of date may be infected. Keep greenhouse on the
palm. dry side.
204 Leaf Spots

Ascochyta clematidina Clematis Leaf and formed on leaves and stems, sometimes cankers
Stem Spot, widespread. On out-door plants at base of young stems. Black pustules in center
stems are infected near the ground and are often of spots discharge spore tendrils in wet weather.
girdled, upper portions dying back. Spores for The fungus winters in old plant refuse, is a weak
initial infection probably come from pycnidia parasite, and is ordinarily too unimportant for
on stumps of old stems. Leaf spots are more control measures.
common in greenhouses, small, water-soaked, Stagonosporopsis hortensis (formerly
then buff with reddish margins. Remove and Ascochyta boltschauseri). Leaf Spot, Pod Spot
destroy infected leaves and stems. of beans, on snap, kidney, lima, and scarlet runner
Ascochyta compositarum Leaf Spot on aster, beans, reported in Oregon. Spots on leaves and
eupatorium, silphium, and sunflower. pods are dark to drab, zonate; light to dark brown
Ascochyta cornicola Dogwood Leaf Spot. pycnidia are numerous.
Ascochyta cypripedii Cypripedium Leaf Spot,
reported on orchid from Wisconsin. Leaf lesions
are narrow, brownish, with a dark brown border. Asteroma
Ascochyta juglandis Walnut Ring Spot. Very
small, round, brown leaf spots between veins, Deuteromycetes, Coelomycetes
ringed with targetlike ridges. The disease is
unimportant in trees sprayed for walnut blight. Pycnidia globose with a radiate subicle, a com-
Ascochyta lycopersici (Didymella pact, crustlike growth of mycelium underneath;
lycopersici) (see ▶Phoma lycopersici) Leaf without an ostiole; spores hyaline, one-celled.
Spot, Ascochyta Blight of tomato, eggplant, and Asteroma garretianum Black Spot on
potato. primrose.
Ascochyta phaseolorum (see ▶Phoma exigua). Asteroma solidaginis Black Spot, Black Scurf
Leaf Spot of snap beans. on goldenrod.
Ascochyta pisi Leaf Spot, Pod Spot of pea. Asteroma tenerrimum Black Spot on
General, but rare in the Northwest. One of three erythronium.
species causing the disease complex known as
Ascochyta blight (also see ▶ Blights). Foliage
spots are circular to irregular, pinhead size to Asteromella (Stictochlorella)
1/2 inch. Stem lesions, at nodes or base, are
brown to purplish black. Brown pycnidia exude Deuteromycetes, Coelomycetes
spore tendrils in wet weather.
Phoma exigua (formerly Ascochyta althaeina). Pycnidia smooth, with ostiole, densely gregari-
Leaf Spot of hollyhock, rose-mallow. ous in asteroma-like spots; spores hyaline, one-
Phoma exigua (formerly Ascochyta asteris). celled.
Leaf Spot of China aster. Spray foliage with Asteromella lupini Leaf Spot on lupine.
bordeaux mixture.
Phoma exigua (formerly Ascochyta
phaseolorum). Leaf Spot of snap beans. Recent Botrytis
isolation and inoculation studies indicate that
the Ascochyta leaf blights of hollyhock, okra, ▶ Blights.
pepper, eggplant, and tomato are all caused by Botrytis fabae Chocolate Leaf Spot on vetch.
strains of the bean pathogen.
Phoma lycopersici (formerly Ascochyta
lycopersici (Didymella lycopersici)). Leaf Spot, Calonectria
Ascochyta Blight of tomato, eggplant, and
potato. Brown spots with concentric rings are See ▶Cylindrocladium under Blights.
Cercospora 205

Calonectria colhounii Leaf Spot on sentry palm.

Calonectria crotalariae Leaf Spot on sentry Cephalosporium
Calonectria theae Leaf Spot on sentry palm. Deuteromycetes, Hyphomycetes

Conidiophores slender or swollen, simple;

Cephaleuros conidia hyaline, one-celled, produced succes-
sively at the tip and collecting in a slime drop,
One of the green algae, possessing chlorophyll produced endogenously in some species; sapro-
but not differentiated into root, stem, and leaves; phytic or parasitic, some species causing vascular
forming motile spores in sporangia. wilts of trees.
Cephaleuros virescens Algal Spot, Red Leaf Cephalosporium apii Celery Brown Spot,
Spot, Green Scurf in the far South or in green- a new disease first reported from Colorado in
houses on acacia, albizzia, ardisia, avocado, bixa, 1943, later from New York and Ohio. Irregular
bischofia, camellia, camphor-tree, cinnamon- light tan or reddish brown shallow lesions are
tree, citrus, grevillea, guava, jasmine, jujube, formed on celery leaf stalks, petioles, and leaflets.
loquat, magnolia, mango, pecan, Japanese per- They may unite to make a scurfy brown streak up
simmon, privet, rhododendron, viburnum. the inside of the stalk and may develop transverse
On some hosts this is a disease of twigs and cracks. Utah and Pascal varieties are most
branches, which may be girdled and stunted, cov- susceptible.
ered with reddish brown hairlike fruiting bodies. Cephalosporium cinnamomeum Leaf Spot of
On magnolia leaves velvety, reddish brown to nephthytis and syngonium. Small circular to
orange, cushiony patches are formed, but in the irregular spots, reddish brown with pale yellow
absence of sporangia (tiny globular heads on fine, borders enlarge, with centers becoming gray and
dense reddish hairs) the leaf spots remain green- papery. In severe cases leaves turn yellow and
ish brown. Occasionally citrus fruits as well as die. Pick off infected leaves. Maintain low tem-
leaves are attacked. perature and humidity.
The sporangia formed on the fine hairs germi- Cephalosporium dieffenbachiae Dief-
nate in moist weather, producing zoospores that fenbachia Leaf Spot. Small red lesions with
enter through stomata and form mycelium-like dark borders appear on young leaves. Spots
chains of algal cells in host tissue. On twigs the sometimes run together, and the whole leaf
alga invades outer cortical tissue, which may turns yellow and dies. Infection is often through
swell abnormally, crack, and afford entrance to mealybug wounds. Avoid promiscuous syring-
injurious fungi. Weakened trees are most suscep- ing; keep temperature and humidity low; control
tible, and disease spread is most rapid in periods mealybugs, and ants that transport them.
of frequent and abundant rains. Twigs may die,
and there may be reduced yield of citrus fruit.
Control Improve draining and other growing Cercospora
conditions; citrus trees sprayed regularly with
copper seldom have algal trouble. If it gets ▶ Blights.
started, follow cleanup pruning with a bordeaux Cercospora abeliae Abelia Leaf Spot, reported
mixture spray in December or January. Repeat from Louisiana. Irregular purple to brown spots;
with bordeaux at start of rainy season or when red defoliation.
stage of the alga is first seen, and spray again Cercospora abelmoschi (see
1 month later. A neutral copper may substitute ▶Pseudocercospora abelmoschi). Leaf Spot on
for bordeaux for the last two applications. The okra, hibiscus.
copper kills beneficial insects parasitic on scales, Cercospora albo-maculans (Syn. Cercosporella
but the oil controls the scale insects. brassicae) (see ▶Pseudocercospora capsellae).
206 Leaf Spots

White Spot of turnip, Chinese cabbage, mustard, Control Crop rotation is highly important. In
and other crucifers, common in the Southeast. a small garden pick off the first spotted leaves.
Cercospora althaeina Leaf Spot of hollyhock Cercospora bougainvilleae (see
and abutilon. Spots circular, angular or irregular, ▶Cercosporidium bougainvilleae). Leaf Spot
1.5 mm, olivaceous to grayish brown, with the first seen in Florida in 1962 and now the most
dead tissue falling out. The fungus winters in old important pathogen of this host.
plant parts. Cercospora brunkii Geranium Leaf Spot,
Cercospora angulata Leaf Spot on philadel- mostly in the South. Spots are circular, light red-
phus, currant, flowering currant, and gooseberry. dish brown with dark brown borders, sometimes
Circular to angular spots, dingy gray centers, dark coalescing to kill entire leaf.
purple to nearly black margins. Cercospora calendulae Calendula Leaf Spot.
Cercospora aquilegiae Columbine Leaf Spot, Spots run together to blight and kill leaves; plants
reported from Kansas, Wisconsin, Oregon. Spots may be destroyed early in the season. Spores
circular to elliptical, reddish brown to nearly enter through stomata of plants more than
black; fruiting is on both sides of the leaf. a month old.
Cercospora arachidicola (Mycosphaerella Cercospora cannabina (see
arachidicola, Teleomorph). Peanut Early Leaf ▶Pseudocercospora cannabina). Leaf Curl and
Spot. Spots light tan aging to reddish or dark Wilt on hemp.
brown with a yellow halo, often confluent. Conid- Cercospora cannabis Leaf Spot on hemp.
iophores on both sides of the leaf, emerging from Cercospora capsici Pepper Leaf Spot,
stomata or breaking through epidermal cells. Stem-end Rot, common in the Southeast, serious
Conidia colorless to pale yellow or olive, with in rainy seasons. Spots 1/7 to 1 inch in diameter are
5 to 12 cells. Control with sulfur-copper dust. first water-soaked then white with dark brown
Cercospora armoraciae Horse-Radish Leaf margins. Leaves turn yellow and drop. The fungus
Spot. Tan to dingy gray lesions with yellow- grows through the pedicel into fruit, causing a rot
brown margin; often slightly zonate. of the stem end. Loss of foliage exposes the fruit to
Cercospora beticola Cercospora Leaf Spot of sunscald. Spray or dust with copper.
beet, general on garden and sugar beets, also on Cercospora circumscissa (Mycosphaerella
swiss chard, spinach. Brown flecks with reddish cerasella, Teleomorph). Leaf Spot, Shot Hole
purple borders become conspicuous spots with of apricot, plum, cherry, cherry-laurel, oriental
ash-gray centers and purple margins. The brittle cherry, and chokecherry. Dead spots are some-
central tissue often drops out, leaving ragged what larger than those caused by other shot-hole
holes. The spots usually remain small but are fungi, but the damage is not serious.
often so numerous that foliage is killed. If suc- Cercospora citrullina Leaf Spot of water-
cessive crops of leaves are lost, the crown of the melon, muskmelon, and other cucurbits. Spots
beet root is elongated and roughened. Leaf spot- are small, circular, black with grayish centers,
ting is of little direct importance except in chard, occurring first on leaves in center of watermelon
where foliage is used for greens. The beet root hills. On cucumber, muskmelon, and squash the
yield is reduced. spots are large and ochre-gray. Defoliation of
The grayish color of the spots is due to long, vines causes reduction in fruit size, but the dis-
thin, septate conidia produced on conidiophores ease is not considered important. Clean up dis-
protruded through stomata in fascicles or groups, eased vines; use a 2-or 3-year rotation; spray or
coming from a knotted mass of mycelium resem- dust as for bacterial wilt.
bling a sclerotium. Conidia are spread by rain, Cercospora concors (see ▶Mycovellosiella
wind, tools, and insects. Infection is through sto- concors). Potato Leaf Spot, Leaf Blotch.
mata; disease spread is most rapid under condi- Cercospora cornicola Dogwood Leaf Spot, in
tions of high humidity that keep stomata open. the Gulf states, often with Septoria florida. Spots
Hot weather favors the disease. irregular without definite borders.
Cercospora 207

Cercospora fusca (see ▶Sirosporium tan, or gray. Avoid syringing; keep plants well
diffusum). Pecan Brown Leaf Spot, prevalent spaced; ventilate greenhouse.
throughout the pecan belt but minor, serious Cercospora rosicola (Mycosphaerella rosicola,
only with high rainfall and in neglected orchards Teleomorph). Cercospora Spot of rose, wherever
where trees lack vigor. roses are grown but more important in the South.
Cercospora lathyrina Leaf Spot on pea and Spots are circular, 1 to 4 mm, but coalescing to
sweet pea, in southern states and north to New irregular areas, purplish or reddish brown with
Jersey and Missouri. Angular to elongate spots pale brown, tan, or gray centers. Perithecia are
have dirty gray centers with a black line border. formed in fallen leaves.
Cercospora lythracearum Leaf Spot on crape- Cercospora smilacis Smilax Leaf Spot. Spots
myrtle, in Texas. Spots circular, pale brown to are more or less circular up to 1/4 inch, dark
gray with a greenish fringe or yellow halo. purplish red, centers fading with age but margins
Cercospora magnoliae (see ▶Cercosporidium remaining definite and dark.
magnoliae). (Mycosphaerella milleri, Cercospora sojina Frog-Eye Disease of soy-
Telleomorph). On magnolia in South. bean. Typical frog-eye spots are formed on
Cercospora melongenae Eggplant Leaf Spot, leaves and elongated reddish lesions on stems,
more common in tropical areas. Yellow lesions changing to brown, gray, or nearly black with
change to large brown areas with concentric rings. age. Pods of late varieties may be infected. The
Cercospora nandinae Nandina Leaf Spot, one fungus winters on diseased leaves and stems.
of the few diseases of this usually healthy shrub. Seed treatment is not effective; crop rotation
Red blotches appear on upper leaf surface with is necessary. Early varieties often escape
centers of older spots almost black. There is injury. There is a wide difference in varietal
a scant sooty fruiting layer on the undersurface. susceptibility.
Reported from Alabama and North Carolina. Cercospora symplocarpi Leaf Spot on snow-
Cercospora personata (Mycosphaerella berry, coralberry, and wolfberry. Very small cir-
berkeleyii, Teleomorph) (see ▶Phaeoisariopsis cular to angular spots, uniformly brown or with
personata). Peanut Leaf Spot, general on peanut. tan centers and brown margins.
Cercospora piaropi Leaf Spot on water- Cercospora sp. Leaf Spot on kalanchoë.
hyacinths. Cercospora zebrina Leaf and Stem Spot on
Cercospora pittospori Pittosporum Leaf Spot, bean, cowpea, groundnut, peanut, birdsfoot tre-
reported from Mississippi, Florida, Louisiana, foil and Lespedeza. Leaf Spot on clovers
and Texas. Spots small, angular, yellow to dull Cercosporidium bougainvilleae (formerly
brown, fruiting in fawn-colored effuse patches on Cercospora bougainvilleae). Leaf Spot first seen
lower surface. in Florida in 1962 and now the most important
Cercospora puderi (see ▶Pseudocercospora pathogen of this host. Lesions are 1 to 5 mm,
puderi). Leaf Spot on rose, reported from Georgia circular, depressed, with brown or tan centers,
and Texas. reddish brown margins and a diffuse chlorotic
Cercospora resedae Leaf Spot, Blight of area.
mignonette, a rapid disease killing much of the Cercosporidium magnoliae (formerly
foliage. Numerous small circular spots, pale Cercospora magnoliae). (Mycosphaerella
yellow with reddish brown borders, run together, milleri, Telleomorph). On magnolia in South.
discoloring the entire leaf. Spores are spread by Leaf spots are small, angular, dark, with narrow
wind and rain; lower leaves are most affected. yellow halo.
Cercospora rhododendri (see Mycovellosiella concors (formerly Cercospora
▶Pseudocercospora handelii). Rhododendron concors). Potato Leaf Spot, Leaf Blotch. Spots
Leaf Spot. none to large irregular brown areas. Fruiting on
Cercospora richardiaecola Leaf Spot on calla undersurface; conidiophores very pale; conidia
lily, sometimes injurious. Spots circular, brown, almost hyaline.
208 Leaf Spots

Phaeoisariopsis personata (formerly

Cercospora personata (Mycosphaerella Cercosporella
berkeleyii, Teleomorph)). Peanut Leaf Spot,
general on peanut. Spots are circular, 1 to 7 mm, Deuteromycetes, Coelomycetes
but may coalesce; dark brown to black, often with
a yellow halo. Conidiophores on both sides of the Conidiophores hyaline, bearing conidia apically
leaf, more numerous on the lower, are arranged or on short branches; conidia hyaline, cylindrical
concentrically in tufts; the epidermis is ruptured. to filiform with several cells (see Figs. 1 and 2);
Spores are pale brown to olivaceous, one-to like Cercospora except for light conidiophores;
eight-septate. In wet seasons vines may be nearly parasitic.
defoliated. Primary infections come from asco- Cercosporella brassicae (see ▶Pseudocer-
spores on overwintered peanut leaves. Sulfur dust cosporella capsellae). Leaf Spot of cabbage, tur-
with 3.5 % copper is recommended; apply every nip, mustard, on West Coast.
10 to 14 days. Pseudocercosporella capsellae (formerly
Pseudocercospora abelmoschi (formerly Cercosporella brassicae). Leaf Spot of cabbage,
Cercospora abelmoschi). Leaf Spot on okra, turnip, mustard, on West Coast. Lesions on
hibiscus. Spots indistinct, but a sooty fruiting of cabbage are black, those on turnip and mustard
spores on under leaf surface. gray with tan margins.
Pseudocercospora cannabina (formerly
Cercospora cannabina). Leaf Curl and Wilt on
Pseudocercospora capsellae (formerly Cercosporidum
Cercospora albo-maculans (Syn. Cercosporella
brassicae)). White Spot of turnip, Chinese Cercosporidium personata (see
cabbage, mustard, and other crucifers, common ▶Phaeoisariopsis personata). Leaf Spot on
in the Southeast. Small, pale, circular slightly peanut.
sunken spots; may coalesce. Phaeoisariopsis personata (formerly
Pseudocercospora puderi (formerly Cercosporidium personata). Leaf Spot on
Cercospora puderi). Leaf Spot on rose, peanut.
reported from Georgia and Texas. Spots are
circular, to 5 mm, with dingy gray centers,
brown or reddish brown margins. Fruiting is
chiefly on the upper surface in dense fascicles of Ciborinia
short conidia.
Sirosporium diffusum (formerly Cercospora ▶ Blights.
fusca). Pecan Brown Leaf Spot, prevalent Ciborinia whetzelii (Syn. Sclerotinia
throughout the pecan belt but minor, serious whetzelii). Black Leaf Spot of poplar, Ink Spot,
only with high rainfall and in neglected orchards from New England States to the Rocky Moun-
where trees lack vigor. Spots are circular to irreg- tains on aspen, black poplar, and other species.
ular, reddish brown, often with grayish concen- Saucerlike, thin black sclerotia are formed in
tric zones. The fungus winters in old spots on leaves, fall to the ground, and produce apothecia
leaves. In Florida the disease appears first in in spring. There is often considerable defoliation,
June or July on mature leaves and may cause and small trees may be killed.
premature defoliation in October. Stuart variety Ciborinia seaveri (Sclerotinia bifrons). Ink
is particularly susceptible; others are more resis- Spot, in western states, producing apothecia on
tant. Control with one application of bordeaux ground under cottonwoods and poplars but path-
mixture between May 15 and June 15. ogenic state confused.
Blumeriella (Coccomyces) 209

Fig. 1 Shot-Hole on
Prunus sp

Fig. 2 Some Leaf-Spot Fungi. Ascochyta, hyaline, two- a perithecium; Phyllosticta, hyaline, one-celled conidia in
celled conidia in pycnidium; Cercosporella, hyaline, sep- pycnidia formed in spots on leaves; Ramularia, hyaline
tate spores on condiophores emerging from a stoma; spores, becoming septate, formed successively on conid-
Cladosporium (formerly Heterosporium), spiny, dark, iophores; Stemphylium, colored muriform spores borne
septate spores; Helmonthosporium, smooth, dark, septate free on mycelium
spores; Mycosphaerella, two-celled hyaline ascospores in

Cladosporium Blumeriella (Coccomyces)

▶ Blotch Diseases. Ascomycetes, Discomycetes

Cladosporium colocasiae Leaf Spot on ele-
phants ear. Blumeriella jaapii (formerly Coccomyces
Cladosporium echinulatum Leaf Spot on hiemalis and Higginisia hiemalis). Cherry Leaf
carnation. Spot, Blight, Shot Hole, general on sweet and
Cladosporium epiphyllum Leaf Spot on locust. sour cherries, the most common and destructive
Cladosporium oxysporum Leaf Spot on tomato. leaf disease of cherries. Leaf spots are circular,
210 Leaf Spots

first purplish, then brown, falling out to give the spots with darker borders. When spots are numer-
shot-hole effect (see Fig. 2). If lesions are numer- ous, leaves turn yellow and die. Similar lesions on
ous, the leaves turn yellow and fall by midsum- young stems may run together into extended
mer, this premature defoliation reducing next cankers, the bark splitting to show black
season’s harvest. The fungus winters in fallen pycnidia, from which ooze out masses of long,
leaves, producing disc-shaped apothecia for pri- white, curved spores. The fungus winters in old
mary infection. Secondary infection comes from dead leaves. Spraying with bordeaux mixture
conidia, formed in whitish masses on the spots in may help.
moist weather, more numerous on the undersur- Coccomyces hiemalis and Higginisia hiemalis
face. New infection continues through the sum- (see ▶Blumeriella jaapii). Cherry Leaf Spot,
mer after harvest. Defoliation prior to ripening Blight, Shot Hole, general on sweet and sour
reduces size and quality of fruit and exposes it to cherries, the most common and destructive leaf
sunscald. Some seasons shoots, spurs, and disease of cherries.
branches are killed, followed by a light crop the Coccomyces kerriae and Higginisia kerriae
next year. Thousands of sour cherry trees have (see ▶Blumeriella kerriae) Kerria Leaf Spot,
been killed. Twig Blight, widespread on kerria from eastern
Control An eradicant spray of a dinitro com- states to Texas.
pound, such as Elgetol, applied to the ground in Coccomyces lutescens (see ▶Blumeriella
early spring, reduces the amount of primary jaapii). Leaf Spot, Shot Hole on cherry-laurel,
inoculum, but summer sprays are also necessary. black cherry, and chokecherry.
On sour cherry this may mean a spray at petal fall, Coccomyces prunophorae (see ▶Blumeriella
another 10 days later, two sprays in June, and jaapii). Leaf Spot, Shot Hole on garden plum
another just after fruit is picked, with more appli- and apricot.
cations, especially on nursery trees, needed in
some seasons. Consult your state experiment
station for suitable materials and schedule for Colletotrichum
your area.
Blumeriella jaapii (formerly Coccomyces ▶ Anthracnose.
lutescens). Leaf Spot, Shot Hole on cherry-laurel, Colletotrichum acutatum Fruit Spot, Crown
black cherry, and chokecherry. Similar to the and Petiole Spot on strawberry.
disease caused by C. hiemalis. Colletotrichum coccodes Leaf Spot and Slight
Blumeriella jaapii (formerly Coccomyces Blight of velvetleaf.
prunophorae). Leaf Spot, Shot Hole on Colletotrichum dematium f. sp. truncata Leaf
garden plum and apricot. Reddish to brown Spot and Stem Canker of Stylosanthes spp.
spots, dark blue initially, produce pinkish spore Colletotrichum elastica (see ▶Colletotrichum
masses on underside of leaves in wet weather. gloeosporioides). Leaf Spot on fig (Fiscus
The shot-hole effect from dropping out of carica). Leaf Spot of basil, flowering dogwood,
dead tissue may be very prominent and accom- cyclamen, jasmine, passion flower, leaf and stem
panied by heavy fruit drop. Spray when shucks spot of calendula and dwarf mistletoe; on many
are off young fruit, 2 or 3 weeks later, and before other hosts as anthracnose.
fruit ripens, with lime sulfur, or with wettable Colletotrichum gloeosporioides (formerly
sulfur. Colletotrichum elastica). Leaf Spot on fig
Blumeriella kerriae (formerly Coccomyces (Fiscus carica). Leaf Spot of basil, flowering
kerriae and Higginisia kerriae). Kerria Leaf dogwood, cycla men, jasmine, passion
Spot, Twig Blight, widespread on kerris from flower, leaf and stem spot of calendula and
eastern states to Texas. Leaves have small, dwarf mistletoe; on many other hosts as
round to angular, light brown or reddish brown anthracnose.
Cryptostictis 211

on lipstick vine, and on weeping fig and leaf spot

Coniothyrium on thyme.

▶ Cankers and Diebacks.

Coniothyrium concentricum (see Cristulariella
▶Microsphaeropsis concentrica). Leaf Spot of
century plant and yucca. Deuteromycetes, Moniliales,
Coniothyrium hellebori Black Spot of Moniliaceae
Christmas rose. Large, irregular, dark brown to
black spots on both sides of leaves, often running Sterile hyphae decumbent; fertile hyphae hya-
together with concentric zonation; many leaves line; ascending in a branched head with conidia
turn yellow prematurely and die; plants are weak- at tips of intermediate branches; spores globose,
ened and fail to mature the normal number of hyaline, one-celled.
leaves. Stems may be cankered, shrivel, and fall Cristulariella depraedans Leaf Spot on sugar
over, with wilting of unopened flower buds. Open and other maples. Spots gray, definite or
petals sometimes have black spots. In wet confluent.
weather in spring and fall the disease can spread Cristulariella moricola Zonate Leaf Spot on
through an entire planting in 2 or 3 days, but Halesia. Leaf Spot on hibiscus and tomato.
continuous moisture is necessary for infection. Cristulariella moricola (Telemorph,
Spray with bordeaux mixture. Grovesinia pyramidalis). Leaf Spot on maple,
Coniothyrium pyrina Leaf Spot, Fruit Spot of tree-of-heaven, apple, bean, blueberry, cherry,
apple, pear. dogwood, hibiscus, sycamore, tung tree, vibur-
Microsphaeropsis concentrica (formerly num, walnut, black walnut, beggar-ticks, trumpet
Coniothyrium concentricum). Leaf Spot of cen- vine, Mexican tea, dayflower, blue waxweed, tick
tury plant and yucca. Spots are zoned, light gray- clover, mistflower, white snakeroot, morning
ish brown, an inch or more in diameter, with glory, Indian tobacco, blue cardinal-flower, beef
concentric rings of tiny black pycnidia. Large steak plant, poke, smart weed, false buckwheat,
portions of leaves may be destroyed. Remove yellow dock, prickly mallow, goldenrod, catbird
and burn diseased leaves. grape, nectarine, grape, maple, serviceberry and
boxelder. Spots yellow-gray with definite

Deuteromycetes, Hyphomycetes Cryptomycina

Hyphae and conidia both dark. Ascomycetes, Rhytismatales

Corynespora cassiicola (Syn.
Helminthosporium vignicola). Soy Bean Target Apothecium splitting irregularly into lobes,
Spot, also on cowpea, tomato, poinsettia, vinca, hyphal layer thin; spores hyaline, one-celled.
and privet; general in South. Circular to irregular, Cryptomycina pteridis Tar Spot of fern,
reddish brown leaf spots, pin point to 1/4 inch, bracken. Spots are usually on lower surface and
often zonate and surrounded by yellow-green between veins; leaves may roll.
halos. Fruit necrotic pitting and freckles are also
found on infected fruit. Dark brown spots on
petioles, pods, and seed. Variety Ogden is mod- Cryptostictis
erately resistant. The same fungus causes reddish
purple spots on azalea, hydrangea and leaf spots ▶ Blights.
212 Leaf Spots

Cryptostictis arbuti (see ▶Seimatosporium Cylindrosporium betulae Brown Leaf Spot of

arbuti). Leaf Spot on Arbutus menziesii, Birch. Sometimes serious enough to defoliate but
Manzanita, ledum. not often present on ornamental trees.
Seimatosporium arbuti (formerly Cryptostictis Cylindrosporium chrysanthemi Chry-
arbuti). Leaf Spot on Arbutus menziesii, santhemum Leaf Spot. Spots are dark brown
Manzanita, ledum. with yellowish margins, increasing to take in the
whole leaf, which hangs down. Similar to more
Cycloconium common Septoria leaf spot.
Cylindrosporium clematidinis Clematis Leaf
Deuteromycetes, Hyphomycetes Spot. Reddish brown spots on lower leaves,
which may drop. Dusting with sulfur has been
Mycelium coiled, spores small, dark, two-celled; suggested.
scarcely different from short hyphae. Cylindrosporium salicinum Willow Leaf Spot.
Cycloconium oleaginum Olive Leaf Spot, Sometimes causing defoliation; can be controlled
Peacock Spot, Ring Spot. Blackish, more or less with bordeaux mixture if necessary.
concentric rings on leaves, especially those weak- Cylindrosporium sp. Leaf Spot on spirea,
ened or old. recorded from a Kansas nursery. Light yellow
lesions turn dark brown, with masses of yellow
Cylindrocladium conidia on underside.

Deuteromycetes, Hyphomycetes
Conidiophores repeatedly dichotomously or tri-
chotomously branched, each terminating in two ▶ Cankers and Diebacks.
or three phialides (cells developing spores); Cytospora sp. Leaf Spot on mulberry.
conidia hyaline, with two or more cells, cylindri-
cal, borne singly; parasitic or saprophytic.
Cylindrocladium avesiculatum Leaf Spot and Dactylaria
Twig Dieback on holly, and Leucothoe¨ sp.
Cylindrocladium colhounii Leaf Spot on bottle- Dactylaria higginsii Leaf Spot on nutsedge.
brush (Callistemon).
Cylindrocladium clavatum Leaf Spot on
bottle-brush (Callistemon). Dichotomophthoropsis
Cylindrocladium pteridis Leaf Spot, Leaf Blight
of Washington palm. Numerous small dark brown Deuteromycetes, Hyphomycetes
spots with light margins are somewhat disfiguring.
Cylindrocladium pteridis Fern Leaf Spot, Leaf Dichotomophthoropsis nymphaearum Leaf
Blotch. Reddish brown lesions run together to Spot on water-lily, and water shield.
cover large areas. Pick off and burn infected fronds.

Cylindrosporium Didymaria

Deuteromycetes, Coelomycetes Deuteromycetes, Hyphomycetes

Acervuli subepidermal, white or pale; conidio- Conidiophores simple, arising from leaf
phores short, simple; conidia hyaline, filiform, surface in loose groups; conidia hyaline, two-
straight or curved, one-celled or septate; parasitic celled, ovate-oblong, borne singly; parasitic on
on leaves. leaves.
Didymosporium 213

Didymaria didyma (see ▶Ramularia didyma).

Leaf Spot on anemone. Angular brown spots.
Ramularia didyma (formerly Didymaria
didyma). Leaf Spot on anemone. Angular brown


Acomycetes, Sphaeriales,

Perithecia separate, innate or finally erumpent,

not beaked; spores two-celled, hyaline.
Didymellina macrospora (Heterosporium
iridis, H. gracilis) (see ▶Mycosphaerella
macrospora, Anamorph). Irish Leaf Spot,
Blotch, Fire on both bulbous and rhizomatous
Fig. 3 Iris Leaf Spot
Didymellina ornithogali (Heterosporium
ornithogali) (see ▶Mycosphaerella ornithogali).
Leaf Spot on star-of-bethlehem.
Didymellina poecilospora A weak parasite Soils deficient in lime apparently favor the dis-
sometimes causing black discoloration of iris ease. Repeated spotting reduces bloom and, after
foliage. a number of years, may kill plants.
Mycosphaerella macrospora (formerly Control It is often sufficient to remove and burn
Didymellina macrospora; Heterosporium iridis, all old leaves at the end of the season; shearing
H. gracilis, Anamorph). Irish Leaf Spot, Blotch, back spotted leaves in midsummer is helpful. If
Fire on both bulbous and rhizomatous iris. The the disease is regularly a problem, spray with
spotting is conspicuous toward the end of the bordeaux mixture, starting when fans are 6 to
season but is not too serious in a normally dry 8 inches high and repeating at 10-to 14-day
season. Usually the spots are confined to the intervals.
upper half of leaves, but if plants are crowded Mycosphaerella ornithogali (formerly
and shaded and the summer is wet, the spotting Didymellina ornithogali; Heterosporium
appears earlier, covers more of the leaf, and is ornithogali, Anamorph). Leaf Spot on star-of-
more damaging. bethlehem. Occasional sooty spots on leaves,
Spots are dark brown at first, surrounded by with foliage blackened and killed in severe
a water-soaked and then yellowing region; they infections.
enlarge into rather oval lesions, up to 1/2 inch
long, with a red-brown border (Fig. 3). Flower
buds and stems of bulbous iris may be attacked. Didymosporium
Tufts of olive conidia turn the centers grayish, the
spores being produced in abundance and splashed Deuteromycetes, Coelomycetes
by rain to neighboring leaves. Infection is
through stomata or directly through the epider- Conidia are slime-spores in acervuli; dark,
mis. The fungus winters as mycelium in old two-celled.
leaves, and in spring produces a fresh crop of Didymosporium arbuticola Leaf Spot on
conidia or perithecia of the Didymellina stage. Arbutus menziesii.
214 Leaf Spots

Dothichiza caroliniana Leaf Spot, Double Spot

Dilophospora of blueberry, found only on Vaccinium australis in
North Carolina, but there causing extensive defo-
Deuteromycetes, Coelomycetes liation. Leaf spots are small, circular, with brown
centers and a dark brown ring, but in late summer
Pycnidia distinct in a stroma; conidia very long, infection spreads to a secondary necrotic area
filiform, with bristlelike hairs at each end. Usu- around the original spot, giving the common
ally found on cereals and sometimes with the name of double spot. Black pycnidia are formed
wheat nematode, causing a disease called twist. sparsely in the spots. All varieties of high bush
Dilophospora geranii (see ▶Pestalozziella blueberries are somewhat susceptible, but Cabot,
subsessilis). Leaf Spot on native geranium. Dixie, Pioneer, and Rancocas are most damaged.
Pestalozziella subsessilis (formerly
Dilophospora geranii). Leaf Spot on native
geranium. Ectostroma

Deuteromycetes, Hyphomycetes
Black stromata formed in leaves and stems.
Deuteromycetes, Coelomycetes Ectostroma liriodendri Tar Spot, widespread
in tulip-trees but perhaps secondary after insect
Pycnidia black, separate, immersed or erumpent, injury.
globose or flattened, ostiolate; conidiophores sim-
ple, slender; conidia hyaline, two-celled, ovoid or Epicoccum
ellipsoid; parasitic or saprophytic. Similar to
Ascochyta but not produced in spots. Deuteromycetes, Hyphomycetes
Diplodia rhododendri (see ▶Encoeliopsis rho-
dodendron). Leaf Spot on rhododendron. Sporodochia dark, rather cushion-shaped; conid-
Encoeliopsis rhododendron (formerly iophores compact or loose, rather short; conidia
Diplodia rhododendri). Leaf Spot on dark, with one or more cells, globose; mostly
rhododendron. saprophytic.
Epicoccum asterinum (see ▶Epicoccum
nigrum). Leaf Spot of yucca; E. neglectum, on
Diplotheca (Stevensea) royal palm; E. nigrum, on Magnolia grandiflora;
E. purpurascens, on amaryllis.
Ascomycetes, Myriangiales Epicoccum nigrum (formerly Epicoccum
asterinum). Leaf Spot of yucca; E. neglectum,
Asci born singly in locules at various levels in on royal palm; E. nigrum, on Magnolia grandi-
a massive stroma; spores dark, several-celled. flora; E. purpurascens, on amaryllis. All of these
Diplotheca wrightii Black Spot, Charcoal Spot may be secondary infections. E. neglectum and
of Opuntia cacti in Florida and Texas uncommon E. purpurascens are also synonyms of E. nigrum.
in the North. Dark spots, 1/4 inch or more in
diameter, are surrounded by a ring of fruiting Exosporium
Deuteromycetes, Hyphomycetes

Dothichiza Conidia on subglobose to convex sporodochia;

spores dark, with two to several cells, somewhat
▶ Cankers and Diebacks. club-shaped.
Glomerella 215

Discogloeum concentricum (formerly conidia hyaline, elongate to filiform, one-to

Exosporium concentricum). Leaf Spot on euony- many-septate, straight or curved, in a slimy
mus and ligustrum (privet) in the South. matrix.
Exosporium concentricum (see Gloeocercospora inconspicua Leaf Spot of
▶Discogloeum concentricum). Leaf Spot on highbush and rabbit-eye blueberry. Circular to
euonymus and ligustrum (privet) in the South. angular brownish spots on leaves, with
sporodochia more frequent on upper surface.
Fusicladium These are flat discs when dry, glistening globules
when wet, containing curved, septate conidia.
Deuteromycetes, Hyphomycetes Gloeocercospora sorghi Copper Spot of turf.
(▶Ramulispora sorghi).
Mycelium forming a stroma under cuticle of host;
conidiophores dark, short; conidia dark, two-
celled, produced successively as pushed-out Gloeosporium
ends of new growing tips. Parasitic on higher
plants, causing scab as well as leaf spots. ▶ Anthracnose.
Fusicladium pisicola Black Leaf of peas, first Asteroma inconspicuum (formerly
reported in Utah in 1921, causing trouble with Gloeosporium inconspicuum). Elm Leaf Spot,
canning peas. Spots start as small, irregular whit- Twig Blight, Anthracnose on American and
ish areas on undersurface of leaflets and stipules, English elms. Subcircular brown spots with
but they darken to gray or black from the closely darker margins and centers are visible on upper
packed layer of dark conidia. The disease is not and lower leaf surfaces.
very important. Cryptocline betularum (formerly
Fusicladium robiniae (see ▶Phaeoisariopsis Gloeosporium betularum). Leaf Spot, Anthrac-
robiniae). Leaf Spot, Seedling Leaf Blight of nose of river birch. Spots are more or less circu-
black locust. lar, 1/8 inch across, brownish with pale centers
Phaeoisariopsis robiniae (formerly and yellow margins.
Fusicladium robiniae). Leaf Spot, Seedling Leaf Gloeosporium betularum (see ▶Cryptocline
Blight of black locust. Spots are small, with light betularum). Leaf Spot, Anthracnose of river
centers and dark margins. There is frequently birch.
defoliation of seedlings, sometimes stunting and Gloeosporium inconspicuum (see ▶Asteroma
death. inconspicuum). Elm Leaf Spot, Twig Blight,
Anthracnose on American and English elms.
Gibbago Gloeosporium mezerei (see ▶Marssonina
daphnes). Leaf Spot on daphne.
Deuteromycetes, Hyphomycetes Gloeosporium rhododendri Leaf Spot on rho-
dodendron, tulip-tree.
Gibbago trianthemae Leaf Spot of horse purs- Gloeosporium ulmicola Elm Leaf Spot. Elon-
lane; a new genus and species, recently described gated spots on midribs, veins, and margins,
(1986), with potential for bioherbicide activity. visible on both leaf surfaces.
Marssonina daphnes (formerly Gloeosporium
Gloeocercospora mezerei). Leaf Spot on daphne. Small brown
spots on both sides of leaves.
Deuteromycetes, Hyphomycetes

Sporodochia formed on surface of host above Glomerella

stomata from hyphae emerging through open-
ings; conidiophores hyaline, simple or branched; ▶ Anthracnose.
216 Leaf Spots

Glomerella cingulata Leaf Spot, widespread on common and less important toward fall. Asco-
queen palm, dracaena, and maranta. Sobralia spores are formed in spring in perithecia on fallen
blight of orchids. Dark discoloration starts at tip dead leaves; conidia are produced as a creamy
of leaves and advances toward base. exudate of spores in summer. The fungus also
Glomerella cingulata Leaf Spot on apple, winters as mycelium in dormant buds.
aucuba, wampi, and croton. See under Control Rake and burn fallen leaves. Chemical
▶ Anthracnose for this fungus on many other control is required only in a wet spring, difficult
hosts. to determine in advance.
Glomerella sp. Black Spot of Vanda orchids.

Ascomycetes, Diaporthales
▶ Anthracnose.
Gnomonia comari Leaf Spot/Blotch and Fruit Perithecia in substratum, beaked, membranous,
Rot of strawberry. separate; spores hyaline, one-celled.
Gnomonia fragariae Leaf Spot, Leaf Blotch of Gnomoniella coryli (see ▶Mamianiella
strawberry. Often associated with Dendrophoma coryli). Leaf Spot on hazel, frequent in northern
causing leaf blight, but not connected. states.
Gnomonia nerviseda (formerly Gnomonia Gnomoniella fimbriata (see ▶Mamianiella
caryae var. pecanae). Pecan Liver Spot. Dark fimbriata). Leaf Spot of hornbeam.
brown circular spots, mostly along midribs on Mamianiella coryli (formerly Gnomoniella
underside of leaves, appear in May and June. In coryli). Leaf Spot on hazel, frequent in northern
autumn the color changes to cinnamon brown, and states. Controlled with bordeaux mixture aided
dark fruiting bodies appear; there may be premature by cleaning up fallen leaves.
defoliation. Spray in May with bordeaux mixture. Mamianiella fimbriata (formerly Gnomoniella
Pecan Vein Spot. Lesions resemble pecan fimbriata). Leaf Spot of hornbeam.
scab on veins or leaf stems; sometimes a narrow
brown lesion extends nearly the length of
a midrib. Defoliation may be moderate or severe. Gonatobotryum
Stuart variety is especially susceptible. Spray
with bordeaux mixture just before and just after Deuteromycetes, Hyphomycetes
pollination; repeat 3 to 4 weeks later.
Gnomonia ulmea, Anamorph, Gloeosporium Conidiophores dark, with spiny inflations at inter-
ulmeum (see ▶Stegophora ulmea). Elm Black vals, around which are borne ovoid, dark, one
Spot, Black Leaf Spot of Elm, general on celled conidia.
American, English, and Chinese elms. Gonatobotryum apiculatum Leaf Spot on
Gnomonia caryae var. pecanae (see witchhazel.
▶Gnomonia nerviseda). Pecan Liver Spot.
Stegophora ulmea (formerly Gnomonia ulmea, Graphium
Anamorph, Gloeosporium ulmeum). Elm Black
Spot, Black Leaf Spot of Elm, general on Deuteromycetes, Hyphomycetes
American, English, and Chinese elms. Spots on
leaves are small but conspicuous, shining coal Synnema or coremium tall, dark, with a rounded
black, and slightly raised. Leaves may turn yellow terminal mass of conidia embedded in mucus;
and drop, with severe defoliation in a wet season, simple, hyaline conidiophores; oblong conidia
especially on Siberian elm. Defoliation in spring reproducing by budding; parasitic.
means death of twigs, but the disease is more Graphium sorbi Leaf Spot of mountain-ash.
Helminthosporium 217

Drechslera erythrospilum (formerly

Guignardia Helminthosporium erythrospilum). Red Leaf
Spot on redtop and bent grasses, widespread in
▶ Blotch Diseases. eastern and midwestern states. Under wet condi-
Guignardia bidwellii f. sp. parthenocissi Leaf tions lesions have small, pale centers with russet
Spot on Boston ivy, pepper-vine, and Virginia borders; in dry weather leaves wither as in
creeper. Spots are numerous, angular, reddish drought but with less evident spotting. Conidia
brown, usually dark brown at margins, with are typically cylindrical, rounded at both ends,
black dots in center, minute pycnidia of the yellowish, and germinate from any or all cells
anamorph Phyllosticta state. Leaves are quite (see Fig. 1).
unsightly and there may be defoliation. Bordeaux Drechslera giganteum (formerly
mixture applied two or three times, starting as Helminthosporium giganteum). Zonate Leaf
leaves are expanding, gives some control, but Spot, Eye Spot on bent grasses, Canada and
the “cure” looks about as bad as the disease. Kentucky bluegrass, and Bermuda grass. The
This fungus is a form of the species causing disease is present in turf and in nursery rows.
black rot of grapes. Spots are small, 1/16 to 1/8 inch, bleached-straw
color in centers. In presence of moisture (dew is
sufficient) the fungus grows periodically into new
Helminthosporium areas, giving the zoned appearance. In continued
wet weather leaves are killed and grass turns
▶ Blights. brown. Metropolitan and velvet bent grasses are
Bipolaris cynodontis (formerly less susceptible. Most injury is in July and
Helminthosporium cynodontis). Bermuda Grass August. The fungus overwinters as dormant
Leaf Blotch, general in South. Olive brown indef- mycelium in old leaves.
inite lesions on dry leaves. Drechslera poae (formerly Helminthosporium
Bipolaris setariae (formerly Helminthosporium vagans). Bluegrass Leaf Spot, Going-Out,
setariae (Drechslera setariae)). Leaf and Petal Melting-out, Foot Rot, general but most injuri-
or Greasy Spot on geranium, areca palm, fish- ous in northeastern states, on bluegrass only.
tail palm, rhapis palm, Calathea spp. Maranta Scattered circular to elongate leaf spots, 0.5 to 3
spp., and Chamaedorea spp. by 1 to 8 mm, have prominent reddish brown to
Bipolaris sorokiniana (formerly black borders; centers are brown changing to
Helminthosporium sativum). Melting-out, preva- straw-colored or white with age. The disease,
lent on bent grass in warm weather. Leaf Spot on favored by cool rainy weather, usually appears
Russian wildrye (Elymus); Spot Blotch on in early spring, sometimes in late fall, and is
switchgrass (Panicum). most severe on close-clipped turf. Grass thins
Bipolaris sorokiniana (formerly out in large areas; roots rot; weeds invade
Helminthosporium sorokiniana). Leaf Spot and exposed soil.
Stem Spot of wild rice. Leaf and Pod Spot on Control Merion bluegrass is quite resistant to
bean. leaf spot and will stand close-clipping. For other
Drechslera catenaria (formerly bluegrasses cut high and fertilize well to help turf
Helminthosporium catenarium). Leaf Spot on withstand the disease.
ribbon-grass. Drechslera siccans (formerly
Drechslera dictyoides (formerly Helminthosporium siccans; Teleomorph,
Helminthosporium dictyoides). Fescue Pyrenophora lolii). Brown Blight on fescue, and
Netblotch, general on fescue. Dark streaks across ryegrass. Leaves die back with numerous dark
green leaves with darker lengthwise streaks form chocolate-brown spots, oval to elongate and
a net pattern. Leaves turn yellow and die back often coalescing. The disease appears in early
from tips. spring in cool, moist weather.
218 Leaf Spots

Drechslera stenacra (formerly Helminthosporium siccans; Teleomorph,

Helminthosporium stenacrum). Leaf Mold on Pyrenophora lolii (see ▶Drechslera siccans).
redtop and bent grasses. Indefinite spots; leaves Brown Blight on fescue, and ryegrass.
dry, withered, in fall. Helminthosporium sorokiniana (see
Drechslera triseptata (formerly ▶Bipolaris sorokiniana). Leaf Spot and Stem
Helminthosporium triseptatum). Leaf Spot, Gray Spot of wild rice. Leaf and Pod Spot on bean.
Leaf Mold on redtop, spike and bentgrasses in Helminthosporium stenacrum (see
Oregon, Washington, and New York. Leaf tips ▶Drechslera stenacra). Leaf Mold on redtop
are killed with vague lesions; gray mold appears and bent grasses.
on dying tissue. Helminthosporium triseptatum (see
Drechslera tritici-repentis (formerly ▶Drechslera triseptata). Leaf Spot, Gray Leaf
Helminthosporium tritici-repentis). Leaf Spot Mold on redtop, spike and bentgrasses in Oregon,
on Russian wildrye (Elymus). Washington, and New York.
Exserohilum rostratum (formerly Helminthosporium tritici-repentis (see
Helminthosporium rostratum). Leaf Spot on ▶Drechslera tritici-repentis). Leaf Spot on
bromelia, areca palm, fishtail palm, rhapis palm, Russian wildrye (Elymus).
sweet sorghum, and Chamaedorea spp. Helminthosporium vagans (see ▶Drechslera
Helminthosporium catenarium (see poae). Bluegrass Leaf Spot, Going-Out, Melting-
▶Drechslera catenaria). Leaf Spot on ribbon- out, Foot Rot, general but most injurious in north
grass. eastern states, on bluegrass only.
Helminthosporium cynodontis (see
▶Bipolaris cynodontis). Bermuda Grass Leaf
Blotch, general in South. Hendersonia
Helminthosporium dictyoides (see
▶Drechslera dictyoides). Fescue Net-blotch, Deuteromycetes, Coelomycetes
general on fescue.
Helminthosporium erythrospilum (see Pycnidia smooth, innate or finally erumpent,
▶Drechslera erythrospilum). Red Leaf Spot on ostiolate; conidia dark, several-celled, elongate
redtop and bent grasses, widespread in eastern to fusoid; saprophytic or parasitic.
and Midwestern states. Hendersonia concentrica Leaf Spot on
Helminthosporium giganteum (see rhododendron.
▶Drechslera giganteum). Zonate Leaf Spot, Hendersonia crataegicola Leaf Spot on haw-
Eye Spot on bent grasses, Canada and Kentucky thorn. Spots irregular, dark brown.
bluegrass, and Bermuda grass.
Helminthosporium rostratum (see
▶Exserohilum rostratum). Leaf Spot on Cladosporium (Heterosporium)
bromelia, areca palm, fishtail palm, rhapis palm,
sweet sorghum, and Chamaedorea spp. Deuteromycetes, Hyphomycetes
Helminthosporium sativum (see ▶Bipolaris
sorokiniana). Melting-out, prevalent on bent grass Conidiophores dark, simple; conidia dark, spiny,
in warm weather. Leaf Spot on Russian wildrye cylindrical, with three or more cells; parasitic,
(Elymus); Spot Blotch on switchgrass (Panicum). causing leaf spots, or saprophytic.
Helminthosporium setariae (Drechslera Acroconidiella escholtziae (formerly
setariae) (see ▶Bipolaris setariae). Leaf and Heterosporium escholtziae). Capsule Spot, Leaf
Petal or Greasy Spot on geranium, areca palm, Spot, Stem Spot of California poppy. Lesions
fishtail palm, rhapis palm, Calathea spp. Maranta faint purplish brown; seed capsules may shrivel.
spp., and Chamaedorea spp. Treat seed with hot water, 125  F, for 30 min.
Kabatia 219

Cladosporium allii (formerly Heterosporium

allii). Leaf Spot on onion, leek, shallot, chive, Illosporium
and garlic; rare in North America. Leaves have
elliptical, depressed, pale brown spots, and Deuteromycetes, Hyphomycetes
yellow and wither from tip downward.
Cladosporium echinulatum (formerly Sporodochia cushionlike, light-colored; conidio-
Heterosporium echinulatum). Fairy Ring Spot, phores hyaline, branched with phialides bearing
Leaf Mold on carnation, occasional in green- conidia apically; spores hyaline, one-celled; par-
houses. Bleached spots on leaves have black asitic or saprophytic, often secondary.
spore groups in ring formation. Syringe as little Illosporium malifoliorum Leaf Spot of apple
as possible and on bright days; control and crabapple.
Cladosporium iridis (formerly Heterosporium
gracile). Leaf Spot on chlorogalum, daylily, Isariopsis
same as H. iridis on iris (conidial state of
Didymellina macrospora). Leaf Spot on iris, Deuteromycetes, Hyphomycetes
blackberry, lily, freezia, and gladiolus.
Cladosporium variabile (formerly Dark, synnemata composed of loose conidio-
Heterosporium variabile). Leaf Spot, pinhead phores with spores at or near tips; conidia dark
“rust” of spinach, cabbage mold, sometimes or pale, with two or more cells, cylindrical to
severe in cold, wet weather. Circular, chlorotic obclavate, often curved; parasitic.
spots with brown or purple margins enlarge and Isariopsis griseola (see ▶Phaeoisariopsis
multiply until they cover most of the leaf, which griseda). Angular Leaf Spot, Pod Spot of
turns yellow, withers, dies. There is a greenish beans, also sweet pea.
black mold on both leaf surfaces, made up of Phaeoisariopsis griseda (formerly Isariopsis
large olive conidia, one-to six-celled, covered griseola). Angular Leaf Spot, Pod Spot of
with warts. Keep plants growing vigorously in beans, also sweet pea. Small, angular brown spots
wall-drained soil. are so numerous they give a checkerboard appear-
Heterosporium allii (see ▶Cladosporium allii). ance to leaves. The fungus forms a gray moldy
Leaf Spot on onion, leek, shallot, chive, and covering over dead areas on underside of leaves.
garlic; rare in North America. Pod spots are conspicuous when present, black
Heterosporium echinulatum (see with red or brown centers, varying from a speck
▶Cladosporium echinulatum). Fairy Ring Spot, to the width of the pod. Small, dark synnemata
Leaf Mold on carnation, occasional in scattered over the surface bear large conidia, with
greenhouses. two to four cells, at top of stalks. They are probably
Heterosporium escholtziae (see wind-disseminated. Control measures are seldom
▶Acroconidiella escholtziae). Capsule Spot, practical. Also, Leaf Spot on kidney bean.
Leaf Spot, Stem Spot of California poppy.
Heterosporium gracile (see ▶Cladosporium
iridis). Leaf Spot on chlorogalum, daylily, Kabatia
same as H. iridis on iris (conidial state of
Didymellina macrospora (Fig. 3)). Leaf Spot on Deuteromycetes, Coelomycetes
iris, blackberry, lily, freezia, and gladiolus.
Heterosporium variabile (see ▶Cladosporium Pycnidia with a radiate shield or scutellum, with
variabile). Leaf Spot, pin-head “rust” of spinach, an ostiole; spores two-celled, hyaline, like a tooth
cabbage mold, sometimes severe in cold, wet at the apex.
weather. Kabatia lonicerae Leaf Spot on honeysuckle.
220 Leaf Spots

Kabatia periclymeni (formerly Leptothyrium

Lasiobotrys periclymeni). Leaf Spot on honeysuckle,
Ascomycetes, Dothideales Leptothyrium californicum Leaf Spot on coast
live oak.
Perithecia in a ring around a sclerotial stroma; Leptothyrium dryinum (see ▶Tubakia
spores dark, two-celled. dryina). Leaf Spot on white oak.
Lasiobotrys lonicerae Leaf Spot on honey- Leptothyrium periclymeni (see ▶Kabatia
suckle. Spot is well-marked with small, dark, periclymeni). Leaf Spot on hon eysuckle,
wartlike stromas. widespread.
Tubakia dryina (formerly Leptothyrium
dryinum). Leaf Spot on white oak.

Deuteromycetes, Coelomycetes
Pycnidia elongate, with a cleft; separate; spores
filiform, with rounded ends, hyaline, continuous Ascomycetes, Diaporthales
to septate on simple conidiophores.
Leptostromella elastica Leaf Spot of rubber- Perithecia innate, beak often lateral, with
plant. The symptoms appear in spots and a shield; paraphyses lacking; spores spindle-
streaks, but infection spreads until the entire shaped to filiform, hyaline.
leaf is involved. Black lines outline spots in Linospora gleditschiae Leaf Spot, Tar Spot on
which small black pycnidia produce long, honey locust in the South. Numerous black
colorless spores. Remove and burn infected fruiting bodies are formed on undersurface of
leaves. leaves.

Deuteromycetes, Coelomycetes
Ascomycetes, Rhytismatales
Pycnidia with a radiate shield, separate; spores
2-celled, hyaline. Fruiting body a hysterothecium, midway between
Leptothyrella liquidambaris (see ▶Tubakia an elongated perithecium and a compressed apo-
dryina). Leaf Spot red on sweetgum. thecium, hard, black, opening with a long narrow
Tubakia dryina (formerly Leptothyrella slit; paraphyses present; hooked at tip; spores
liquidambaris). Leaf Spot red on sweetgum. filiform, septate or continuous. Most species
cause needle casts.
Lophodermium schweinitzii Rhododendron
Leptothyrium Leaf Spot. Large silvery white spots with red,
raised margins have very prominent oval, black
Deuteromycetes, Coelomycetes fruiting bodies on the upper surface. Lower side
of spots is a light chocolate brown. Infected por-
Pycnidium flattened with a more or less radiate tions may fall out, leaving irregular holes. The
shield, opening with a ostiole; spores one-celled, disease is more common on native than on hybrid
hyaline, on simple conidiophores. varieties.
Melasmia 221

There may be premature defoliation and kill-

Macrophoma ing of twigs.
Marssonina rhabdospora (Teleomorph,
▶ Cankers and Diebacks. Pleuroceras populi) (see ▶Cylindrosporium
Macrophoma candollei Leaf Spot of boxwood. populinum). Leaf Spot of poplar.
Conspicuous black pycnidia on dead leaves, usu- Marssonina rosae Anamorph state of the rose
ally straw-colored, sometimes brown or tan. The blackspot fungus, Diplocarpon rosae.
fungus is a weak parasite coming in secondarily Marssonina tremulae Leaf Spot on poplar.
after winter injury or other predisposing factors. Marssonina truncatula (see ▶Didymosporina
aceris). Leaf Spot and Leaf Blight of Norway
Marssonina Piggotia fraxini (formerly Marssonina fraxini).
Ash Leaf Spot, sometimes serious in nursery stock,
▶ Anthracnose. controlled by spraying with bordeaux mixture.
Cylindrosporium populinum (formerly
Marssonina rhabdospora; Teleomorph,
Pleuroceras populi). Leaf Spot of poplar. Mastigosporium
Brown spots on living leaves; beaked pyriform
perithecia formed in fallen leaves over winter. Deuteromycetes, Hyphomycetes
Didymosporina aceris (formerly Marssonina
truncatula). Leaf Spot and Leaf Blight of Norway Conidiophores hyaline, very short, simple;
maple. conidia with four or more cells, with or without
Diplosporonema delastrei (formerly apical appendages: broadly cylindrical with
Marssonina delastrei). Leaf Spot on corncockle rounded or pointed ends; parasitic on grasses.
and campion. Mastigosporium rubricosum Leaf Fleck on
Discella ochroleuca (formerly Marssonina redtop and bent grasses. Spores with rounded
ochroleuca). Leaf Spot on oak, American chest- ends, without appendages.
nut. Spots are circular, yellow to brown with
concentric markings, small on chestnut, up to an
inch on oak. Melanconium
Marssonina daphnes Daphne Leaf Spot.
Small, thick brown spots on both sides of leaf, Deuteromycetes, Coelomycetes
which turns yellow, dies.
Marssonina delastrei (see ▶Diplosporonema Acervuli subcutaneous or subcortical, conic or
delastrei). Leaf Spot on corn-cockle and discoid, black; with setae; conidiophores simple;
campion. conidia dark, one-celled, ovoid to ellipsoid; par-
Marssonina fraxini (see ▶Piggotia fraxini). asitic or saprophytic.
Ash Leaf Spot, sometimes serious in nursery Melanconium pandani Leaf Spot on pandanus.
stock, controlled by spraying with bordeaux
Marssonina juglandis See ▶Gnomonia Melasmia
leptostyla under Anthracnose.
Marssonina ochroleuca (see ▶Discella Deuteromycetes, Coelomycetes
ochroleuca). Leaf Spot on oak, American
chestnut. Pycnidia in a broad, black, flattened stroma that is
Marssonina populi Poplar Leaf Spot. Brown superficial or nearly so, dimidiate; conidiophores
spots with darker margins. simple or branched; spores hyaline or subhyaline,
222 Leaf Spots

one-celled, allantoid or fusoid; parasitic on Monochaetia monochaeta Leaf Spot on chest-

leaves; anamorph states of Rhytisma. nut, white, red, and coast live oaks, winged elm,
Melasmia falcata Tar Spot of persimmon. hickories, especially destructive in the Southeast.
Melasmia menziesiae Leaf Spot tar spot of Spots are large, 1 to 2 inches in diameter, with
azalea. pale green or yellow centers with a red and brown
border or concentric zones of gray, yellow, and
Micropeltis brown.
Symptoms appear most often in late summer
▶ Blights. when loss of green tissue is not so important.
Micropeltis alabamensis Black Leaf Spot on
Lembosina (Morenoella)

Microstroma See ▶Lembosia under Black Mildew.

Lembosina quercina (formerly Morenoella
Basidiomycetes, Exobasidiales quercina). Leaf Spot, Black Mildew of red and
black oaks; twig blight of white oak, common in
Sporodochia small, white, breaking through epi- Southeast. Spots are purplish black, roughly cir-
dermis; conidiophores hyaline, one-celled, some- cular, up to 1/3 inch across, on upper surface and
what clavate, bearing conidia on short sterigmata; irregular brown areas on underside. Mycelium is
spores hyaline, one-celled, small, oblong; superficial in early summer, but by late summer
parasitic. “Conidia” are now known to be there are subcuticular hyphae and a black shield
basidiospores. formed over a flat cushion of fertile cells. Asci are
Microstroma juglandis Leaf Spot, White mature and shield is fissured by spring.
Mold, Downy Spot, Witches’ Broom of pecan, Morenoella quercina (see ▶Lembosina
walnut, and hickory. Yellow blotching of upper quercina). Leaf Spot, Black Mildew of red and
side of leaves and a glistening white coating on black oaks; twig blight of white oak, common in
underside, due to pustules with enormous num- Southeast.
bers of spores, may be accompanied by prema-
ture defoliation. On shagbark hickory the fungus
also invades the stems, causing witches’ broom Mycosphaerella
up to 3 feet across. Leaves formed on them in
spring are yellow-green, with white powder on ▶ Blights.
underside. Leaflets are small, curled, and soon Mycosphaerella angulata Angular Leaf Spot
drop. Prune out witches’ broom; spray with bor- of muscadine grapes. Many small, angular black
deaux mixture. spots, more conspicuous on lower surface of
leaves, which may turn yellow and die.
Mycosphaerella arachidis Peanut Leaf Spot.
Microthyriella ▶Cercospora arachidicola.
Mycosphaerella berkeleyi Peanut Leaf Spot.
▶ Fruit Spots. ▶Cercospora personata.
Microthyriella cuticulosa Black Spot of holly. Mycosphaerella (Anamorph, Pseudcercospora)
Dark spots on leaves of American holly, Georgia. bolleana Leaf Spot of fig, and rubber-tree.
Mycosphaerella (Anamorph, Asteromella
brassicae) brassicicola Ring Spot of crucifers,
Monochaetia chiefly cabbage and cauliflower, sometimes brus-
sels sprouts, broccoli, and turnip. Dead spots in
▶ Cankers and Diebacks. leaves, small to 1/2 inch, are surrounded by
Mycosphaerella 223

a green zone that keeps its color even if the rest of Mycosphaerella colorata Mountain-Laurel
the leaf turns yellow. Small black pycnidia are Leaf Spot. ▶Phyllosticta kalmicola.
deeply embedded in the dead tissue, often in Mycosphaerella (Anamorph, Pseudocercospora
concentric rings. In moist weather conidia ooze cruenta) cruenta Leaf Spot, Leaf Blotch of
from pycnidia in pink tendrils. The fungus win- soybean, and kidney bean. Leaf spots distinct to
ters in old plant refuse, and ascospores are forc- indistinct, circular to irregular, greenish to
ibly ejected from perithecia in spring. The disease yellowish to rusty brown to almost red, some-
is confined to the Pacific Coast and, as black times with gray centers.
blight, is serious on the seed crop in the Puget Mycosphaerella effigurata (Anamorph, Piggotia
Sound area. Sanitary measures and crop rotation fraxini). Ash Leaf Spot, general east of the Plains.
keep it in check. Spots small, purple to brown with yellow borders.
Mycosphaerella caroliniana Leaf Spot, Purple Mycosphaerella fragariae Strawberry Leaf
Blotch on oxydendron (sourwood). Reddish or Spot, Black-Seed Disease, general on straw-
purple spots on foliage in midsummer have dry, berries. Leaf spots are first purple then reddish
brown centers. Pycnidia embedded in tissue with light brown or white centers, 1/8 to 1/4 inch
break through lower surface, spores being formed across. Spots are also present on petioles and fruit
in great numbers. stems, and occasionally there are black spots on
Mycosphaerella caryigena Pecan Downy fruit, with blackened achenes prominent against
Spot. Conidial stage has been listed as a the white of unripe berries. Fruit is poor; total
Pseudocercosporella caryigena. Leaf spots are yield is reduced; runner plants are weakened.
pale yellow when young, turning yellow-brown, Conidia of the Ramularia stage are produced in
brown, or black. Conidia produced in minute clusters of short conidiophores on underside of
acervuli on underside of leaves form a white diseased areas; perithecia are formed in autumn
downy or frosty coating; leaves may drop early. at the edge of the leaf spots where the fungus
Spores are spread in rain, fog, and dew. The fungus winters. New conidia are produced in spring
overwinters in leaves, liberating ascospores in with most infection taking place through stomata.
spring to infect new foliage. Moneymaker and There is a difference in varietal susceptibility.
Stuart varieties are especially susceptible. Control Set healthy plants in well-drained soil;
Control Turn under old leaves before spring remove diseased leaves before planting; spray
(plowing under winter cover in spring takes care with bordeaux mixture before planting and fol-
of this). Spray as for scab, bordeaux mixture low with two or three more applications. The
when leaves are half-grown and bordeaux plus conidia are very sensitive to copper, which pre-
4 pounds of zinc sulfate when tips of small nuts vents sporulation and kills nongerminated spores.
have turned brown. Mycosphaerella fraxinicola (Anamorph,
Mycosphaerella cerasella ▶Cercospora Phyllosticta viridis). Ash Leaf Spot, east of the
circumscissa. Rocky Mountains.
Mycosphaerella (Anamorph, Cercospora) Mycosphaerella juglandis Leaf Spot of black
cercidicola Redbud Leaf Spot, general. Spots walnut.
are circular to angular or irregular with raised Mycosphaerella liriodendri (Phyllosticta
dark brown borders. With age, lesions become liriodendrica). Tulip-Tree Leaf Spot.
grayish above and rusty brown on the undersur- Mycosphaerella louisianae Purple Leaf Spot
face, with the leaf tissue yellow-green outside the of strawberry, in the South. Large, irregular,
borders. Spores are formed on fascicles of conid- reddish purple areas.
iophores projecting through stomata. The fungus Mycosphaerella mori Mulberry Leaf Spot,
winters on fallen leaves, producing perithecia in widespread, with the conidial stage reported
spring. Twigs may be attacked as well as foliage. variously as Cercosporella, Cylindrosporium,
Mycosphaerella citri Leaf Spot or Greasy Spot Phloeospora, Septogloeum, and Septoria. Yellow
on citrus. areas on upper leaf surface are matched by
224 Leaf Spots

whitish patches underneath, the fungus forming Mycosphaerella rosicola ▶Cercospora

a white downy or powdery coating. The disease is rosicola.
most serious in shady locations. Mycosphaerella rubi ▶Septoria rubi.
Mycosphaerella nigromaculans Black Stem Mycosphaerella pyri (Anamorph, Septoria
Spot of cranberry, reported from all cranberry pyricola). Pear Leaf Spot, also on quince, occa-
areas, often associated with red leaf spot. The sional on apple. Spots are small, 1/8 to 1/4 inch,
fungus enters through leaves, grows down the grayish in center, dotted with black fruiting bod-
petioles, and forms elongated black spots on the ies, with a well-defined dark brown margin.
stems, which may be completely girdled, followed There are marked differences in susceptibility in
by defoliation. Fruiting bodies are produced in pear varieties. Flemish Beauty, Duchess, and
autumn on dead stems with ascospores discharged Winter Nellis are moderately, and Kieffer very,
in rainy periods in spring. The anamorph state of resistant. Sprays applied for leaf blight or scab
the fungus is a Ramularia nigromaculans. control leaf spot.
Mycosphaerella nyssicola (Anamorph,
Phyllosticta nyssae). Tupelo Leaf Spot, on sour
gum and water tupelo. Purplish irregular blotches, Mycocentrospora
an inch or more across, are scattered on upper leaf
surface with lower surface dark brown. There may Mycocentrospora verrucosa Leaf Spot on
be heavy defoliation. Perithecia mature in spring euonymus.
on fallen leaves.
Mycosphaerella personata (Anamorph,
Isariopsis clavispora). Leaf Spot, widespread on Myrothecium
muscadine and other grapes after midseason.
Spots are dark brown, 1/4 to 1/2 inch, surrounded Deuteromycetes, Hyphomycetes
by a yellow circle but with a narrow band of
normal green between spot and circle. Sporodochia cushionlike, light or dark; conidio-
Mycosphaerella populicola (Anamorph, phores subhyaline to colored, repeatedly
Septoria populicola); M. populorum branched, bearing conidia terminally; conidia
(S. musiva). Leaf Spot of native poplar; Canker subhyaline to dark, one-celled, ovoid to elongate;
on twigs and branches of hybrid poplars. weakly parasitic or saprophytic.
Mycosphaerella psilospora (Anamorph, Myrothecium roridum Leaf Spot on snap-
Septoria querceti). Oak Leaf Spot on red and dragon, stock, eremurus, gardenia, hollyhock,
other oaks, common in Iowa. Spots very small, aeschynanthus, aglaonema, aphelandra, dieffen-
circular, with strawcolor centers and dark margins. bachia, episcia, fittonia, nematanthus, hoya,
Mycosphaerella ribis Leaf Spot of flowering peperomia, pilea, and sphathiphyllum. Tissues
currant. are dry, brittle, with black sporodochia. Snap-
Mycosphaerella ribis (M. grossulariae, dragon leaves and flowering stems wilt, with
Anamorph, Septoria ribis). Leaf Spot of goose- sunken cracked cankers. Avoid excessive mois-
berry, current. Numerous small brown spots with ture; sterilize soil.
grayish centers are formed on both sides of
leaves; there may be premature defoliation. The
fungus winters in leaves, producing ascospores in
late spring. Two sprays of bordeaux mixture plus Nematostoma
1 pint of self-emulsifying cottonseed oil per 100
gallons have given good control of leaf spot on Ascomycetes, Dothideales
gooseberries in New York. The first application
is about June 1, the second in July right after fruit Nematostoma occidentalis Leaf Hair Discolor-
is picked. ation on Artemisa.
Pestalotia 225

Pestalotia aucubae Aucuba Leaf Spot. The

Neottiospora fungus appears as a weak parasite in sunscald
spots or after other fungi.
Deuteromycetes; Coelomycetes Pestalotia cliftoniae Leaf Spot on buckwheat-
tree. Ashy or pale brown spots. Spores usually
Pycnidia dark, smooth, innate; spores hyaline, curved, constricted at septa, three setae at crest.
one-celled with two to several appendages at the Pestalotia funerea (see ▶Pestalotiopsis
apex. funerea). Leaf Spot, Bark and Cone Spot on
Alpakesa yuccifolia (formerly Neottiospora conifers.
yuccifolia). Yucca Leaf Spot. Pestalotia guepini (see ▶Pestalotiopsis
Neottiospora yuccifolia (see ▶Alpakesa maculans). Camellia Leaf Spot, widespread.
yuccifolia). Yucca Leaf Spot. Pestalotia leucothoës (see ▶Pestalotiopsis
leucothoe¨s). Leucothoë Leaf Spot, apparently